Processos ecol��gicos e filogen��ticos na estrutura����o de comunidades de girinos

P��s-gradua����o em Biologia Animal - IBILCE

Ecologia tr��fica de pequenos mam��feros n��o voadores em uma ��rea cont��nua de Mata Atl��ntica

P��s-gradua����o em Ci��ncias Biol��gicas (Zoologia) - IBRC

Rela����o entre tamanho corporal e estrutura de comunidade em anf��bios anuros do munic��pio de Botucatu/SP

P��s-gradua����o em Ci��ncias Biol��gicas (Zoologia) - IBB

Rela����o entre os atributos funcionais e caracter��sticas do habitat em comunidades de girinos

Funda����o de Amparo �� Pesquisa do Estado de S��o Paulo (FAPESP)

Spatio-temporal distribution of calling male frogs in Morro do Diabo State Park (Southeastern Brazil): implications for conservation and management in a seasonally dry tropical forest

The aims of this study are to determine how anuran calling males are spatially and temporally distributed in the Morro do Diabo State Park (MDSP), and to test for indicator species in different breeding habitats. We found high species turnover among breeding habitats of different hydroperiods. Our results indicate that spatial partitioning is more important than seasonal occurrence in permitting species coexistence at MDSP. Indicator Species Analysis identified one species associated with a local stream, one species associated with permanent ponds, two species associated with temporary ponds, and three species associated with a semipermanent pond. Guilds of species do not appear to stratify their distributions throughout the year, since most species occur during the rainy and warm months and are thus temporally overlapping. The main conservation implication of our work is the need to protect the different areas of the park where these unique breeding habitats, and their associated anuran assemblages, are found.

Morphological patterns and community organization in leaf-litter ant assemblages

We present a new approach to determine the number and composition of guilds, using the hyperdiverse leaf-litter ant fauna as a model, based on appropriate morphological variables and species co-occurrence null models to describe the complex assemblages of interacting Species Community structure at the 1-m(2) scale. We obtained 18 linear morphometric measures from 949 workers of 171 leaf-litter ant species (18762 measurements) surveyed in four Atlantic Forest localities to test whether the assemblages are morphologically structured; the morphological characters were selected to indicate diet and foraging habits. Principal components analysis was used to characterize the morphospace and to describe the guild structure (number of species and composition). The guild proportionality assembly rule (significant tendency toward constant proportion of species in guilds) was assessed at the 1-m(2) scale. Our analysis indicates that the division of leaf-litter ants into guilds is based mainly on microhabitat distribution in the leaf-litter, body size and shape, eye size, and phylogeny. The same guild scheme applied to four more sites shows that different Atlantic Forest areas have the same leaf-fitter ant guilds. The guild proportionality assembly rule was confirmed for most guilds, Suggesting that there are guild-specific limitations on species coexistence within assemblages; on the other hand, in a few cases the variance in guild proportion was greater than expected under the null assumptions. Other studies on ant functional group classification are partially supported by our quantitative morphological analysis. Our results, however, imply that there are more compartments than indicated in previous models, particularly among cryptic species (confined to soil and litter) and tropical climate specialists. We argue that a general null model for the analysis of species association based oil morphology can reveal objectively defined groups and may thus contribute to a robust theory to explain community structure in general and have important consequences on studies of litter ant community ecology in particular.

Seed fate and seedling dynamics after masting in two African rain forest trees

How the effects of biotic factors are moderated by abiotic factors, and their consequences for species interactions, is generally understudied in ecology. A key abiotic feature of forests is regular canopy disturbances that create temporary patches, or ���gaps,��� of above-average light availability. Co-occurring in lowland primary forest of Korup National Park (Cameroon), Microberlinia bisulcata and Tetraberlinia bifoliolata are locally dominant, ectomycorrhizal trees whose seeds share predator guilds in masting years. Here, we experimentally tested the impact of small mammal predators upon seedling abundance, growth, and survivorship. In 2007, we added a fixed density of seeds of each species to exclosures at 48 gap���understory locations across 82.5 ha within a large Microberlinia grove, and at 15 locations outside it. For both species, small mammals removed more seeds in gaps than in understory, whereas this was reversed for seeds killed by invertebrates. Nonetheless, Microberlinia lost twice as many seeds to small mammals, and more to invertebrates in exclosures, than Tetraberlinia, which was more prone to a pathogenic white fungus. After six weeks, both species had greater seedling establishment in gaps than understory, and in exclosures outside compared to exclosures inside the grove. In the subsequent two-year period, seedling growth and survivorship peaked in exclosures in gaps, but Microberlinia had more seedlings' stems clipped by animals than Tetraberlinia, and more than twice the percentage of leaf area damaged. Whereas Microberlinia seedling performance in gaps was inferior to Tetraberlinia inside the grove, outside it Microberlinia had reduced leaf damage, grew taller, and had many more leaves than Tetraberlinia. No evidence was found for ���apparent mutualism��� in the understory as seedling establishment of both species increased away from (>25 m) large stems of either species, pointing to ���apparent competition��� instead. In gaps, Microberlinia seedling establishment was lower near Tetraberlinia than conspecific adults because of context-dependent small mammal satiation. Stage-matrix analysis suggested that protecting Microberlinia from small mammals could increase its population growth rate by 0.06. In the light of prior research we conclude that small mammals and canopy gaps play an important role in promoting species coexistence in this forest, and that their strong interaction contributes to Microberlinia's currently very poor regeneration.

Changes in biodiversity and vegetation composition in the central Swiss Alps during the transition from pristine forest to first farming

Aim: We investigate the response of vegetation composition and plant diversity to increasing land clearance, burning and agriculture at the Mesolithic���Neolithic transition (c. 6400���5000 bc) when first farming was introduced. Location: The Valais, a dry alpine valley in Switzerland. Methods: We combine high-resolution pollen, microscopic charcoal and sedimentological data to reconstruct past vegetation, fire and land use. Pollen evenness, rarefaction-based and accumulation-based palynological richness analyses were used to reconstruct past trends in plant diversity. Results: Our results show that from c. 5500 cal. yr bc, slash-and-burn activities created a more open landscape for agriculture, at the expense of Pinus and Betula forests. Land clearance by slash-and-burn promoted diverse grassland ecosystems, while on the long term it reduced woodland and forest diversity, affecting important tree species such as Ulmus and Tilia. Main conclusions: Understanding the resilience of Alpine ecosystems to past disturbance variability is relevant for future nature conservation plans. Our study suggests that forecasted land abandonment in the Alps will lead to pre-Neolithic conditions, with significant biodiversity losses in abandoned grassland ecosystems. Thus, management measures for biodiversity, such as ecological compensation areas, are needed in agricultural landscapes with a millennial history of human impact, such as the non-boreal European lowlands. Our study supports the hypothesis that species coexistence is maximized at an intermediate level of disturbances. For instance, species richness decreased when fire exceeded the quasi-natural variability observed during the Mesolithic times. Under a more natural disturbance regime, rather closed Pinus sylvestris and mixed oak forests would prevail.

Chapter Six ��� Individual Trait Variation and Diversity in Food Webs

In recent years, there has been a renewed interest in the ecological consequences of individual trait variation within populations. Given that individual variability arises from evolutionary dynamics, to fully understand eco-evolutionary feedback loops, we need to pay special attention to how standing trait variability affects ecological dynamics. There is mounting empirical evidence that intra-specific phenotypic variation can exceed species-level means, but theoretical models of multi-trophic species coexistence typically neglect individual-level trait variability. What is needed are multispecies datasets that are resolved at the individual level that can be used to discriminate among alternative models of resource selection and species coexistence in food webs. Here, using one the largest individual-based datasets of a food web compiled to date, along with an individual trait-based stochastic model that incorporates Approximate Bayesian computation methods, we document intra-population variation in the strength of prey selection by different classes or predator phenotypes which could potentially alter the diversity and coexistence patterns of food webs. In particular, we found that strongly connected individual predators preferentially consumed common prey, whereas weakly connected predators preferentially selected rare prey. Such patterns suggest that food web diversity may be governed by the distribution of predator connectivity and individual trait variation in prey selection. We discuss the consequences of intra-specific variation in prey selection to assess fitness differences among predator classes (or phenotypes) and track longer term food web patterns of coexistence accounting for several phenotypes within each prey and predator species.

Herbivores differentially limit the seedling growth and sapling recruitment of two dominant rain forest trees

Resource heterogeneity may influence how plants are attacked and respond to consumers in multiple ways. Perhaps a better understanding of how this interaction might limit sapling recruitment in tree populations may be achieved by examining species��� functional responses to herbivores on a continuum of resource availability. Here, we experimentally reduced herbivore pressure on newly established seedlings of two dominant masting trees in 40 canopy gaps, across c. 80 ha of tropical rain forest in central Africa (Korup, Cameroon). Mesh cages were built to protect individual seedlings, and their leaf production and changes in height were followed for 22 months. With more light, herbivores increasingly prevented the less shade-tolerant Microberlinia bisulcata from growing as tall as it could and producing more leaves, indicating an undercompensation. The more shade-tolerant Tetraberlinia bifoliolata was much less affected by herbivores, showing instead near to full compensation for leaf numbers, and a negligible to weak impact of herbivores on its height growth. A stage-matrix model that compared control and caged populations lent evidence for a stronger impact of herbivores on the long-term population dynamics of M. bisulcata than T. bifoliolata. Our results suggest that insect herbivores can contribute to the local coexistence of two abundant tree species at Korup by disproportionately suppressing sapling recruitment of the faster-growing dominant via undercompensation across the light gradient created by canopy disturbances. The functional patterns we have documented here are consistent with current theory, and, because gap formations are integral to forest regeneration, they may be more widely applicable in other tropical forest communities. If so, the interaction between life-history and herbivore impact across light gradients may play a substantial role in tree species coexistence.

Intraspecific Aggregation Alters Competitive Interactions in Experimental Plant Communities

We tested the prediction from spatial competition models that intraspecific aggregation may promote coexistence and thus maintain biodiversity with experimental communities of four annual species. Monocultures, three-species mixtures, and the four-species mixture were sown at two densities and with either random or intraspecifically aggregated distributions. There was a hierarchy of competitive abilities among the four species. The weaker competitors showed higher aboveground biomass in the aggregated distribution compared to the random distribution, especially at high density. In one species, intraspecific aggregation resulted in an 86% increase in the number of flowering individuals and a 171% increase in the reproductive biomass at high density. The competitively superior species had a lower biomass in the aggregated distribution than in the random distribution at high density. The data support the hypothesis that the spatial distribution of plants profoundly affects competition in such a way that weaker competitors increase their fitness while stronger competitors are suppressed when grown in the neighborhood of conspecifics. This implies that the spatial arrangement of plants in a community can be an important determinant of species coexistence and biodiversity.

Density-dependent dynamics of a dominant rain forest tree change with juvenile stage and time of masting

Although negative density dependence (NDD) can facilitate tree species coexistence in forests, the underlying mechanisms can differ, and rarely are the dynamics of seedlings and saplings studied together. Herein we present and discuss a novel mechanism based on our investigation of NDD predictions for the large, grove-forming ectomycorrhizal mast fruiting tree, Microberlinia bisulcata (Caesalpiniaceae), in an 82.5-ha plot at Korup, Cameroon. We tested whether juvenile density, size, growth and survival decreases with increasing conspecific adult basal area for 3245 ���new��� seedlings and 540 ���old��� seedlings (< 75-cm tall) during an approximately 4-year study period (2008���2012) and for 234 ���saplings��� (��� 75-cm tall) during an approximately 6-year study period (2008���2014). We found that the respective densities of new seedlings, old seedlings and saplings were positively, not and negatively related to increasing BA. Maximum leaf numbers and heights of old seedlings were negatively correlated with increasing basal areas, as were sapling heights and stem diameters. Whereas survivorship of new seedlings decreased by more than one-half with increasing basal area over its range in 2010���2012, that of old seedlings decreased by almost two-thirds, but only in 2008���2010, and was generally unrelated to conspecific seedling density. In 2010���2012 relative growth rates in new seedlings��� heights decreased with increasing basal area, as well as with increasing seedling density, together with increasing leaf numbers, whereas old seedlings��� growth was unrelated to either conspecific density or basal area. Saplings of below-average height had reduced survivorship with increasing basal area (probability decreasing from approx. 0.4 to 0.05 over the basal area range tested), but only sapling growth in terms of leaf numbers decreased with increasing basal area. These static and dynamic results indicate that NDD is operating within this system, possibly stabilizing the M. bisulcata population. However, these NDD patterns are unlikely to be caused by symmetric competition or by consumers. Instead, an alternative mechanism for conspecific adult���juvenile negative feedback is proposed, one which involves the interaction between tree phenology and ectomycorrhizal linkages.

Simulation Model of Benthic Antarctic Assemblages (SIMBAA), link to PC-executable

SIMBAA is a spatially explicit, individual-based simulation model. It was developed to analyse the response of populations of Antarctic benthic species and their diversity to iceberg scouring. This disturbance is causing a high local mortality providing potential space for new colonisation. Traits can be attributed to model species, e.g. in terms of reproduction, dispersal, and life span. Physical disturbances can be designed in space and time, e.g. in terms of size, shape, and frequency. Environmental heterogeneity can be considered by cell-specific capacities to host a certain number of individuals. When grid cells become empty (after a disturbance event or due to natural mortality of of an individual), a lottery decides which individual from which species stored in a pool of candidates (for this cell) will recruit in that cell. After a defined period the individuals become mature and their offspring are dispersed and stored in the pool of candidates. The biological parameters and disturbance regimes decide on how long an individual lives. Temporal development of single populations of species as well as Shannon diversity are depicted in the main window graphically and primary values are listed. Examples for simulations can be loaded and saved as sgf-files. The results are also shown in an additional window in a dimensionless area with 50 x 50 cells, which contain single individuals depicted as circles; their colour indicates the assignment to the self-designed model species and the size represents their age. Dominant species per cell and disturbed areas can also be depicted. Output of simulation runs can be saved as images, which can be assembled to video-clips by standard computer programs (see GIF-examples of which "Demo 1" represents the response of the Antarctic benthos to iceberg scouring and "Demo 2" represents a simulation of a deep-sea benthic habitat).

Community attributes determine facilitation potential in a semi-arid steppe

Studies on positive plant���plant relations have traditionally focused on pair-wise interactions. Conversely, the interaction with other co-occurring species has scarcely been addressed, despite the fact that the entire community may affect plant performance. We used woody vegetation patches as models to evaluate community facilitation in semi-arid steppes. We characterized biotic and physical attributes of 53 woody patches (patch size, litter accumulation, canopy density, vegetation cover, species number and identity, and phylogenetic distance), and soil fertility (organic C and total N), and evaluated their relative importance for the performance of seedlings of Pistacia lentiscus, a keystone woody species in western Mediterranean steppes. Seedlings were planted underneath the patches, and on their northern and southern edges. Woody patches positively affected seedling survival but not seedling growth. Soil fertility was higher underneath the patches than elsewhere. Physical and biotic attributes of woody patches affected seedling survival, but these effects depended on microsite conditions. The composition of the community of small shrubs and perennial grasses growing underneath the patches controlled seedling performance. An increase in Stipa tenacissima and a decrease in Brachypodium retusum increased the probability of survival. The cover of these species and other small shrubs, litter depth and community phylogenetic distance, were also related to seedling survival. Seedlings planted on the northern edge of the patches were mostly affected by attributes of the biotic community. These traits were of lesser importance in seedlings planted underneath and in the southern edge of patches, suggesting that constraints to seedling establishment differed within the patches. Our study highlights the importance of taking into consideration community attributes over pair-wise interactions when evaluating the outcome of ecological interactions in multi-specific communities, as they have profound implications in the composition, function and management of semi-arid steppes.

Assembly processes of invertebrate communities in springs across different spatial scales

Il presente lavoro ha lo scopo di comprendere i processi sottesi ai pattern di coesistenza tra le specie di invertebrati sorgentizi, distinguendo tra dinamiche stocastiche e deterministiche. Le sorgenti sono ecosistemi complessi e alcune loro caratteristiche (ad esempio l���insularit��, la stabilit�� termica, la struttura ecotonale ���a mosaico���, la frequente presenza di specie rare ed endemiche, o l���elevata diversit�� in taxa) le rendono laboratori naturali utili allo studio dei processi ecologici, tra cui i processi di assembly. Al fine di studiare queste dinamiche �� necessario un approccio multi-scala, per questo motivi sono state prese in considerazione tre scale spaziali. A scala locale �� stato compiuto un campionamento stagionale su sette sorgenti (quattro temporanee e tre permanenti) del Monte Prinzera, un affioramento ofiolitico vicino alla citt�� di Parma. In questa area sono stati valutati l���efficacia e l���impatto ambientale di diversi metodi di campionamento e sono stati analizzati i drivers ecologici che influenzano le comunit��. A scala pi�� ampia sono state campionate per due volte 15 sorgenti della regione Emilia Romagna, al fine di identificare il ruolo della dispersione e la possibile presenza di un effetto di niche-filtering. A scala continentale sono state raccolte informazioni di letteratura riguardanti sorgenti dell���area Paleartica occidentale, e sono stati studiati i pattern biogeografici e l���influenza dei fattori climatici sulle comunit��. Sono stati presi in considerazione differenti taxa di invertebrati (macroinvertebrati, ostracodi, acari acquatici e copepodi), scegliendo tra quelli che si prestavano meglio allo studio dei diversi processi in base alle loro caratteristiche biologiche e all���approfondimento tassonomico raggiungibile. I campionamenti biologici in sorgente sono caratterizzati da diversi problemi metodologici e possono causare impatti sugli ambienti. In questo lavoro sono stati paragonati due diversi metodi: l���utilizzo del retino con un approccio multi-habitat proporzionale e l���uso combinato di trappole e lavaggio di campioni di vegetazione. Il retino fornisce dati pi�� accurati e completi, ma anche significativi disturbi sulle componenti biotiche e abiotiche delle sorgenti. Questo metodo �� quindi raccomandato solo se il campionamento ha come scopo un���approfondita analisi della biodiversit��. D���altra parte l���uso delle trappole e il lavaggio della vegetazione sono metodi affidabili che presentano minori impatti sull���ecosistema, quindi sono adatti a studi ecologici finalizzati all���analisi della struttura delle comunit��. Questo lavoro ha confermato che i processi niche-based sono determinanti nello strutturare le comunit�� di ambienti sorgentizi, e che i driver ambientali spiegano una rilevante percentuale della variabilit�� delle comunit��. Infatti le comunit�� di invertebrati del Monte Prinzera sono influenzate da fattori legati al chimismo delle acque, alla composizione e all���eterogeneit�� dell���habitat, all���idroperiodo e alle fluttuazioni della portata. Le sorgenti permanenti mostrano variazioni stagionali per quanto riguarda le concentrazioni dei principali ioni, mentre la conduttivit��, il pH e la temperatura dell���acqua sono pi�� stabili. �� probabile che sia la stabilit�� termica di questi ambienti a spiegare l���assenza di variazioni stagionali nella struttura delle comunit�� di macroinvertebrati. L���azione di niche-filtering delle sorgenti �� stata analizzata tramite lo studio della diversit�� funzionale delle comunit�� di ostracodi dell���Emilia-Romagna. Le sorgenti ospitano pi�� del 50% del pool di specie regionale, e numerose specie sono state rinvenute esclusivamente in questi habitat. Questo �� il primo studio che analizza la diversit�� funzionale degli ostracodi, �� stato quindi necessario stilare una lista di tratti funzionali. Analizzando il pool di specie regionale, la diversit�� funzionale nelle sorgenti non �� significativamente diversa da quella misurata in comunit�� assemblate in maniera casuale. Le sorgenti non limitano quindi la diversit�� funzionale tra specie coesistenti, ma si pu�� concludere che, data la soddisfazione delle esigenze ecologiche delle diverse specie, i processi di assembly in sorgente potrebbero essere influenzati da fattori stocastici come la dispersione, la speciazione e le estinzioni locali. In aggiunta, tutte le comunit�� studiate presentano pattern spaziali riconoscibili, rivelando una limitazione della dispersione tra le sorgenti, almeno per alcuni taxa. Il caratteristico isolamento delle sorgenti potrebbe essere la causa di questa limitazione, influenzando maggiormente i taxa a dispersione passiva rispetto a quelli a dispersione attiva. In ogni caso nelle comunit�� emiliano-romagnole i fattori spaziali spiegano solo una ridotta percentuale della variabilit�� biologica totale, mentre tutte le comunit�� risultano influenzate maggiormente dalle variabili ambientali. Il controllo ambientale �� quindi prevalente rispetto a quello attuato dai fattori spaziali. Questo risultato dimostra che, nonostante le dinamiche stocastiche siano importanti in tutte le comunit�� studiate, a questa scala spaziale i fattori deterministici ricoprono un ruolo prevalente. I processi stocastici diventano pi�� influenti invece nei climi aridi, dove il disturbo collegato ai frequenti eventi di disseccamento delle sorgenti provoca una dinamica source-sink tra le diverse comunit��. Si �� infatti notato che la variabilit�� spiegata dai fattori ambientali diminuisce all���aumentare dell���aridit�� del clima. Disturbi frequenti potrebbero provocare estinzioni locali seguite da ricolonizzazioni di specie provenienti dai siti vicini, riducendo la corrispondenza tra gli organismi e le loro richieste ambientali e quindi diminuendo la quantit�� di variabilit�� spiegata dai fattori ambientali. Si pu�� quindi concludere che processi deterministici e stocastici non si escludono mutualmente, ma contribuiscono contemporaneamente a strutturare le comunit�� di invertebrati sorgentizi. Infine, a scala continentale, le comunit�� di ostracodi sorgentizi mostrano chiari pattern biogeografici e sono organizzate lungo gradienti ambientali principalmente collegati altitudine, latitudine, temperatura dell���acqua e conducibilit��. Anche la tipologia di sorgente (elocrena, reocrena o limnocrena) �� influente sulla composizione delle comunit��. La presenza di specie rare ed endemiche inoltre caratterizza specifiche regioni geografiche.