Quais são os fatores que estruturam as comunidades de girinos no pantanal em diferentes escalas e sub-regiões?

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Unraveling the drivers of community dissimilarity and species extinction in fragmented landscapes

Communities in fragmented landscapes are often assumed to be structured by species extinction due to habitat loss, which has led to extensive use of the species-area relationship (SAR) in fragmentation studies. However, the use of the SAR presupposes that habitat loss leads species to extinction but does not allow for extinction to be offset by colonization of disturbed-habitat specialists. Moreover, the use of SAR assumes that species richness is a good proxy of community changes in fragmented landscapes. Here, we assessed how communities dwelling in fragmented landscapes are influenced by habitat loss at multiple scales; then we estimated the ability of models ruled by SAR and by species turnover in successfully predicting changes in community composition, and asked whether species richness is indeed an informative community metric. To address these issues, we used a data set consisting of 140 bird species sampled in 65 patches, from six landscapes with different proportions of forest cover in the Atlantic Forest of Brazil. We compared empirical patterns against simulations of over 8 million communities structured by different magnitudes of the power-law SAR and with species-specific rules to assign species to sites. Empirical results showed that, while bird community composition was strongly influenced by habitat loss at the patch and landscape scale, species richness remained largely unaffected. Modeling results revealed that the compositional changes observed in the Atlantic Forest bird metacommunity were only matched by models with either unrealistic magnitudes of the SAR or by models ruled by species turnover, akin to what would be observed along natural gradients. We show that, in the presence of such compositional turnover, species richness is poorly correlated with species extinction, and z values of the SAR strongly underestimate the effects of habitat loss. We suggest that the observed compositional changes are driven by each species reaching its individual extinction threshold: either a threshold of forest cover for species that disappear with habitat loss, or of matrix cover for species that benefit from habitat loss.

Tributary and Mainstem Benthic Macroinvertebrate Communities Linked By Direct Dispersal and Indirect Habitat Alteration

Benthic communities in tributary-mainstem networks might interact via downstream drift of invertebrates or material from tributaries and adult dispersal from the mainstem. Depending on the strength of these interactions, mainstem downstream communities are expected to be more similar to tributary communities due to drift or habitat alteration. Communities not connected by flow are expected to be similar due to adult dispersal but decreasing in similarity with distance from the mainstem. We investigated interactions between invertebrate communities of a 7th order river and 5th order tributary by comparing benthic community structure in the river upstream and downstream of the tributary confluence and upstream in the tributary. Non-metric multidimensional scaling showed invertebrate communities and habitat traits from river locations directly downstream of the tributary clustered tightly, intermediate between tributary and mid-channel river locations. In addition, Bray-Curtis dissimilarity increased between the mainstem and tributary with distance upstream in the tributary. Our results indicate that similarities between mainstem and tributary communities are potentially caused by direct mass effects from tributary to downstream mainstem communities by invertebrate drift and indirect mass effects by habitat restructuring via material delivery from the tributary, as well as potential effects of adult dispersal from the river on proximal tributary communities.

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’insularità, la stabilità termica, la struttura ecotonale “a mosaico”, la frequente presenza di specie rare ed endemiche, o l’elevata diversità in taxa) le rendono laboratori naturali utili allo studio dei processi ecologici, tra cui i processi di assembly. Al fine di studiare queste dinamiche è necessario un approccio multi-scala, per questo motivi sono state prese in considerazione tre scale spaziali. A scala locale è stato compiuto un campionamento stagionale su sette sorgenti (quattro temporanee e tre permanenti) del Monte Prinzera, un affioramento ofiolitico vicino alla città 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 comunità. A scala più 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 comunità. 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 più accurati e completi, ma anche significativi disturbi sulle componenti biotiche e abiotiche delle sorgenti. Questo metodo è quindi raccomandato solo se il campionamento ha come scopo un’approfondita analisi della biodiversità. 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 comunità. Questo lavoro ha confermato che i processi niche-based sono determinanti nello strutturare le comunità di ambienti sorgentizi, e che i driver ambientali spiegano una rilevante percentuale della variabilità delle comunità. Infatti le comunità di invertebrati del Monte Prinzera sono influenzate da fattori legati al chimismo delle acque, alla composizione e all’eterogeneità 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 conduttività, il pH e la temperatura dell’acqua sono più stabili. È probabile che sia la stabilità termica di questi ambienti a spiegare l’assenza di variazioni stagionali nella struttura delle comunità di macroinvertebrati. L’azione di niche-filtering delle sorgenti è stata analizzata tramite lo studio della diversità funzionale delle comunità di ostracodi dell’Emilia-Romagna. Le sorgenti ospitano più del 50% del pool di specie regionale, e numerose specie sono state rinvenute esclusivamente in questi habitat. Questo è il primo studio che analizza la diversità funzionale degli ostracodi, è stato quindi necessario stilare una lista di tratti funzionali. Analizzando il pool di specie regionale, la diversità funzionale nelle sorgenti non è significativamente diversa da quella misurata in comunità assemblate in maniera casuale. Le sorgenti non limitano quindi la diversità funzionale tra specie coesistenti, ma si può 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 comunità 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 comunità emiliano-romagnole i fattori spaziali spiegano solo una ridotta percentuale della variabilità biologica totale, mentre tutte le comunità risultano influenzate maggiormente dalle variabili ambientali. Il controllo ambientale è quindi prevalente rispetto a quello attuato dai fattori spaziali. Questo risultato dimostra che, nonostante le dinamiche stocastiche siano importanti in tutte le comunità studiate, a questa scala spaziale i fattori deterministici ricoprono un ruolo prevalente. I processi stocastici diventano più influenti invece nei climi aridi, dove il disturbo collegato ai frequenti eventi di disseccamento delle sorgenti provoca una dinamica source-sink tra le diverse comunità. Si è infatti notato che la variabilità spiegata dai fattori ambientali diminuisce all’aumentare dell’aridità 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 quantità di variabilità spiegata dai fattori ambientali. Si può quindi concludere che processi deterministici e stocastici non si escludono mutualmente, ma contribuiscono contemporaneamente a strutturare le comunità di invertebrati sorgentizi. Infine, a scala continentale, le comunità di ostracodi sorgentizi mostrano chiari pattern biogeografici e sono organizzate lungo gradienti ambientali principalmente collegati altitudine, latitudine, temperatura dell’acqua e conducibilità. Anche la tipologia di sorgente (elocrena, reocrena o limnocrena) è influente sulla composizione delle comunità. La presenza di specie rare ed endemiche inoltre caratterizza specifiche regioni geografiche.

Crayfish assemblage shifts in a large drought-prone wetland: the roles of hydrology and competition

1. Faster growing, larger and/or more aggressive crayfish species are predicted to dominate permanent waterbodies. We tested this prediction using a 9 year dataset for two species of crayfish (Procambarus alleni and Procambarus fallax) co-existing in a sub-tropical flowing slough in southern Florida. Using a series of laboratory and mesocosm experiments we also compared life history traits and performance of the respective species to test mechanisms that could explain dominance shifts in the local crayfish assemblages. 2. Over the 9-year period, P. alleni densities were the greatest in shallower, shorterhydroperiod areas bordering the slough, while P. fallax densities were higher in deeper, longer-hydroperiod central areas. These areas were separated by 0.8–2 km of continuous wetland with no apparent barriers to movement between them. 3. Density of P. fallax was not strongly affected by any measures of hydrological variation, while P. alleni density increased with more severe drought conditions. Following the strongest droughts, P. alleni colonized areas in the centre of the slough where they had been absent or scarce in wetter years. 4. We conducted experiments to compare growth rates, drought tolerance, and competitive dominance of these species. P. alleni survived drought conditions better, had higher growth rates, and was the dominant competitor for space and food. While drought probably limits P. fallax in the drier slough habitats, neither drought sensitivity nor interspecific competition with P. fallax can explain decreases of P. alleni with wetter conditions. 5. Our results indicate that a competition-colonization tradeoff cannot explain the crayfish compositional dynamics in this wetland because P. alleni is both the best competitor and the best at surviving in and colonizing areas with the strongest droughts. Future attention should focus on the potential for selective effects of predators that co-vary with hydrology. 6. The traits (large size, fast growth, competitive dominance) exhibited by P. alleni, which is absent in long-hydroperiod wetlands, are those exhibited by dominant crayfish in permanent lakes and streams containing fish. Although these traits make crayfish less vulnerable to fish in some lakes and streams, life-history models of community structure across permanence gradients suggest the opposite traits should be favoured for co-existence with fish.

Seasonal Fish Dispersal in Ephemeral Wetlands of the Florida Everglades

We hypothesized that fishes in short-hydroperiod wetlands display pulses in activity tied to seasonal flooding and drying, with relatively low activity during intervening periods. To evaluate this hypothesis, sampling devices that funnel fish into traps (drift fences) were used to investigate fish movement across the Everglades, U.S.A. Samples were collected at six sites in the Rocky Glades, a seasonally flooded karstic habitat located on the southeastern edge of the Everglades. Four species that display distinct recovery patterns following drought in long-hydroperiod wetlands were studied: eastern mosquitofish (Gambusia holbrooki) and flagfish (Jordanella floridae) (rapid recovery); and bluefin killifish (Lucania goodei) and least killifish (Heterandria formosa) (slow recovery). Consistent with our hypothesized conceptual model, fishes increased movement soon after flooding (immigration period) and just before drying (emigration period), but decreased activity in the intervening foraging period. We also found that eastern mosquitofish and flagfish arrived earlier and showed stronger responses to hydrological variation than either least killifish or bluefin killifish. We concluded that these fishes actively colonize and escape ephemeral wetlands in response to flooding and drying, and display species-specific differences related to flooding and drying that reflect differences in dispersal ability. These results have important implications for Everglades fish metacommunity dynamics.

Ecological and evolutionary effects of dispersal on freshwater zooplankton

A recent focus on contemporary evolution and the connections between communities has sought to more closely integrate the fields of ecology and evolutionary biology. Studies of coevolutionary dynamics, life history evolution, and rapid local adaptation demonstrate that ecological circumstances can dictate evolutionary trajectories. Thus, variation in species identity, trait distributions, and genetic composition may be maintained among ecologically divergent habitats. New theories and hypotheses (e.g., metacommunity theory and the Monopolization hypothesis) have been developed to understand better the processes occurring in spatially structured environments and how the movement of individuals among habitats contributes to ecology and evolution at broader scales. As few empirical studies of these theories exist, this work seeks to further test these concepts. Spatial and temporal dispersal are the mechanisms that connect habitats to one another. Both processes allow organisms to leave conditions that are suboptimal or unfavorable, and enable colonization and invasion, species range expansion, and gene flow among populations. Freshwater zooplankton are aquatic crustaceans that typically develop resting stages as part of their life cycle. Their dormant propagules allow organisms to disperse both temporally and among habitats. Additionally, because a number of species are cyclically parthenogenetic, they make excellent model organisms for studying evolutionary questions in a controlled environment. Here, I use freshwater zooplankton communities as model systems to explore the mechanisms and consequences of dispersal and to test these nascent theories on the influence of spatial structure in natural systems. In Chapter one, I use field experiments and mathematical models to determine the range of adult zooplankton dispersal over land and what vectors are moving zooplankton. Chapter two focuses on prolonged dormancy of one aquatic zooplankter, Daphnia pulex. Using statistical models with field and mesocosm experiments, I show that variation in Daphnia dormant egg hatching is substantial among populations in nature, and some of that variation can be attributed to genetic differences among the populations. Chapters three and four explore the consequences of dispersal at multiple levels of biological organization. Chapter three seeks to understand the population level consequences of dispersal over evolutionary time on current patterns of population genetic differentiation. Nearby populations of D. pulex often exhibit high population genetic differentiation characteristic of very low dispersal. I explore two alternative hypotheses that seek to explain this pattern. Finally, chapter four is a case study of how dispersal has influenced patterns of variation at the community, trait and genetic levels of biodiversity in a lake metacommunity.