Environmental heterogeneity at two spatial scales affects litter diversity-decomposition relationships.

The effects of biodiversity on ecological processes have been experimentally evaluated mainly at the local scale under homogeneous conditions. To scale up experimentally based biodiversity-functioning relationships, there is an urgent need to understand how such relationships are affected by the environmental heterogeneity that characterizes larger spatial scales. Here, we tested the effects of an 800-m elevation gradient (a large-scale environmental factor) and forest habitat (a fine-scale factor) on litter diversity-decomposition relationships. To better understand local and landscape scale mechanisms, we partitioned net biodiversity effects into complementarity, selection, and insurance effects as applicable at each scale. We assembled different litter mixtures in aquatic microcosms that simulated natural tree holes, replicating mixtures across blocks nested within forest habitats (edge, interior) and elevations (low, mid, high). We found that net biodiversity and complementarity effects increased over the elevation gradient, with their strength modified by forest habitat and the identity of litter in mixtures. Complementarity effects at local and landscape scales were greatest for combinations of nutrient-rich and nutrient-poor litters, consistent with nutrient transfer mechanisms. By contrast, selection effects were consistently weak and negative at both scales. Selection effects at the landscape level were due mainly to nonrandom overyielding rather than spatial insurance effects. Our findings demonstrate that the mechanisms by which litter diversity affects decomposition are sensitive to environmental heterogeneity at multiple scales. This has implications for the scaling of biodiversity-ecosystem function relationships and suggests that future shifts in environmental conditions due to climate change or land use may impact the functioning of aquatic ecosystems.

Different plant compartments, different yeasts: The example of the bromeliad phyllosphere.

The plant phyllosphere is one of the largest sources of microorganisms, including yeasts. In bromeliads, the knowledge of yeasts is dispersed and still incipient. To understand the extent of our knowledge of the subject, this review proposes to compile and synthesize existing knowledge, elucidating possible patterns, biotechnological and taxonomic potentials, bringing to light new knowledge, and identifying information gaps. For such, we systematically review scientific production on yeasts in bromeliads using various databases. The results indicated that the plant compartments flowers, fruits, leaves, and water tank (phytotelma) have been studied when focusing on the yeast community in the bromeliad phyllosphere. More than 180 species of yeasts and yeast-like fungi were recorded from the phyllosphere, 70% were exclusively found in one of these four compartments and only 2% were shared among all. In addition, most of the community had a low frequency of occurrence, and approximately half of the species had a single record. Variables such as bromeliad subfamilies and functional types, as well as plant compartments, were statistically significant, though inconclusive and with low explanatory power. At least 50 yeast species with some biotechnological potentials have been isolated from bromeliads. More than 90% of these species were able to produce extracellular enzymes. In addition, other biotechnological applications have also been recorded. Moreover, new species have been described, though yeasts were only exploited in approximately 1% of the existing bromeliads species, which highlights that there is still much to be explored. Nevertheless, it appears that we are still far from recovering the completeness of the diversity of yeasts in this host. Furthermore, bromeliads proved to be a good ecological model for prospecting new yeasts and for studies on the interaction between plants and yeasts. In addition, the yeast community diverged among plant compartments, establishing bromeliads as a microbiologically complex and heterogeneous mosaic.

Tremella ananatis sp. nov. and Tremella lamprococci sp. nov., two yeast species associated with bromeliads.

Eight yeast isolates with an affinity to the genus Tremella were obtained from bromeliads from different locations in Brazil. Although the formation of basidia and basidiocarp were not observed, on the basis of the results of sequence analysis of the D1/D2 domain of the large subunit (LSU) rRNA gene and internal transcribed spacer (ITS) region, we suggest that these isolates represent two novel species of the genus Tremella. These yeasts are phylogenetically related to Tremella saccharicola and Tremella globispora. Therefore, we propose Tremella ananatis sp. nov. and Tremella lamprococci sp. nov. as novel yeast species of the order Tremellales (Agaricomycotina, Basidiomycota). Sequence analysis revealed that Tremella ananatis sp. nov. differs by 11 and 28 nucleotide substitutions from Tremella saccharicola in the D1/D2 sequence and ITS region, respectively. Moreover, Tremella lamprococci sp. nov. differs by 15 and 29 nucleotide substitutions from Tremella globispora in the D1/D2 sequence and ITS region, respectively. The holotypes of Tremella ananatis sp. nov. and Tremella lamprococci sp. nov. are CBS 14568T and CBS 14567T, and the MycoBank numbers are MB840480 and MB840481, respectively.

In bromeliad phytotelma, anthropic disturbance does not affect the nematode trophic structure.

Phytotelmata (sing. phytotelma) are plant-associated reservoirs of rainwater and organic debris. These freshwater ecosystems are found in tree and bamboo holes, pitcher plants, and tank-forming bromeliads. Some studies suggest that anthropic disturbance (AD) may change the physico-chemical properties (PCPs) of the water retained in the phytotelma, and indirectly impact its biota. Hence, new AD-bioindicators could be found in the phytotelma biota. To test this hypothesis, three areas of Atlantic Forest were selected, distinct only by the level of long-term AD. In these areas, we monitored the nematode trophic structure and the water PCPs in the bromeliad Neoregelia cruenta during two years (eight seasons). Significant differences among areas were found in some seasons for total nematode abundance and/or the abundance of some trophic groups, but no pattern emerged relative to the level of AD. Anthropic disturbance did not impact nematode trophic structure possibly because the water PCPs remained fairly similar in all three areas. Our results do not corroborate previous reports that AD alters phytotelma water. On the other hand, our findings support previous studies suggesting that nematodes inhabiting bromeliad phytotelma are not good candidates for AD-bioindicators.

Nematode trophic structure in the phytotelma of Neoregelia cruenta (Bromeliaceae) in relation to microenvironmental and climate variables.

The term phytotelma (pl. phytotelmata) designates a plant-associated reservoir of fresh water and organic debris. Phytotelmata in tank bromeliads are abundant in tropical forests, and they provide shelter, food, and water for many metazoans. Among the invertebrates known to inhabit phytotelmata, nematodes are the least studied, despite their important role in nutrient and energy cycles in freshwater ecosystems. This study was conceived to characterize the nematode trophic structure in the phytotelma of the bromeliad N. cruenta, and to identify climate and microenvironmental variables that impact it. Nematode abundance (total and per trophic group), rainfall, air temperature, the amount of organic debris fallen into the phytotelma, and eight physico-chemical properties (PCPs) of the water retained in the bromeliad tank - volume; temperature; pH; dissolved organic carbon, nitrogen, oxygen, and solids; and electrical conductivity - were monitored during two years in a natural reserve in Brazil. Bacterial and hyphal feeder nematodes predominated over other trophic groups. Nematode abundance (total and per trophic group) was not impacted by fluctuations in rainfall or air temperature. The amount of organic debris fallen into the phytotelma correlated positively with nematode abundance (total and per trophic group). Regarding the PCPs of water, the only significant correlation - positive - was between the amount of dissolved oxygen and the abundance of hyphal feeder nematodes. These results bring a clearer understanding of the ecology of nematodes inhabiting phytotelmata, which are peculiar and understudied freshwater ecosystems.

Inquiline predator increases nutrient-cycling efficiency of Nepenthes rafflesiana pitchers.

The modified-leaf pitchers of Nepenthes rafflesiana pitcher plants are aquatic, allochthonous ecosystems that are inhabited by specialist inquilines and sustained by the input of invertebrate prey. Detritivorous inquilines are known to increase the nutrient-cycling efficiency (NCE) of pitchers but it is unclear whether predatory inquilines that prey on these detritivores decrease the NCE of pitchers by reducing detritivore populations or increase the NCE of pitchers by processing nutrients that may otherwise be locked up in detritivore biomass. Nepenthosyrphus is a small and poorly studied genus of hoverflies and the larvae of one such species is a facultatively detritivorous predator that inhabits the pitchers of N. rafflesiana. We fitted a consumer-resource model to experimental data collected from this system. Simulations showed that systems containing the predator at equilibrium almost always had higher NCEs than those containing only prey (detritivore) species. We showed using a combination of simulated predator/prey exclusions that the processing of the resource through multiple pathways and trophic levels in this system is more efficient than that accomplished through fewer pathways and trophic levels. Our results thus support the vertical diversity hypothesis, which predicts that greater diversity across trophic levels results in greater ecosystem functioning.

Digestive mutualism in a pitcher plant supports the monotonic rather than hump-shaped stress-gradient hypothesis model.

The stress-gradient hypothesis (SGH) predicts that the strength and frequency of facilitative interactions increase monotonically with increasing environmental stress, but some empirical studies have found this decrease at extreme stress levels, suggesting a hump-shaped SGH instead. However, empirical studies of the SGH are often hindered by confounding resource and non-resource stress gradients. Nepenthes pitcher plants trap animal prey using modified-leaf pitfall traps which are also inhabited by organisms known as inquilines. Inquilines may assist pitchers in the digestion of trapped prey. This interaction is known as a digestive mutualism and is both mutualistic and facilitative by definition. Inquiline species may also facilitate each other via processing chain commensalisms. We used in vitro experiments to examine the isolated effect of resource stress on the outcomes of two facilitative interactions: (i) digestive mutualism-facilitation of pitcher nutrient sequestration by two inquiline dipteran larvae, culicids and phorids; (ii) processing chain commensalism-facilitation between these two inquiline taxa. The net nutritional benefit of phorids on N. gracilis was found to conform more to a monotonic rather than hump-shaped SGH model. However, the effect of culicids on N. gracilis and the effects of culicids and phorids on each other were weak. These findings provide compelling evidence that changes in facilitation along an isolated resource stress gradient conform to the predictions of the monotonic SGH model rather than that of the revised hump-shaped model, and highlight the importance of isolating stress gradients in empirical tests of the SGH.

A dipteran larva-pitcher plant digestive mutualism is dependent on prey resource digestibility.

Positive species interactions tend to be context dependent. However, it is difficult to predict how benefit in a mutualism changes in response to changing contexts. Nepenthes pitcher plants trap animal prey using leaf pitfall traps known as pitchers. Many specialized inquiline organisms inhibit these pitchers, and are known to facilitate the digestion of prey carcasses in them. Nepenthes gracilis traps diverse arthropod prey taxa, which are likely to differ greatly in the ease with which they may be digested, independently of inquilines, by plant enzymes. In this study, we used in vitro experiments to compare the nutritional benefit provided by phorid (scuttle fly) and culicid (mosquito) dipteran larvae to their host, N. gracilis, and to each other. The effects of phorids on N. gracilis nutrient sequestration were very variable, being positive for large prey which have low digestibility, but negative for small prey which are highly digestible. However, the effect of culicids on N. gracilis and the effects of culicids and phorids on each other were not significantly altered by prey type. These results show that a digestive mutualism is highly dependent on the digestibility of the resource-a context dependency that conforms well to the predictions of the stress-gradient hypothesis in facilitation research. Our findings have significant implications for many other digestive mutualisms, and also suggest that greater insights may be gained from the synthesis of concepts between the fields of mutualism and facilitation research.

Predatory dipteran larva contributes to nutrient sequestration in a carnivorous pitcher plant.

The fluids of Nepenthes pitcher plants are habitats to many specialized animals known as inquilines, which facilitate the conversion of prey protein into pitcher-absorbable nitrogen forms such as ammonium. Xenoplatyura beaveri (Diptera: Mycetophilidae) is a predatory dipteran inquiline that inhabits the pitchers of Nepenthes ampullaria Larvae of X. beaveri construct sticky webs over the fluid surface of N. ampullaria to ensnare emerging adult dipteran inquilines. However, the interaction between X. beaveri and its host has never been examined before, and it is not known if X. beaveri can contribute to nutrient sequestration in N. ampullaria. Xenoplatyura beaveri individuals were reared in artificial pitchers in the laboratory on a diet of emergent Tripteroides tenax mosquitoes, and the ammonium concentration of the pitcher fluids was measured over time. Fluid ammonium concentration in tubes containing X. beaveri was significantly greater than those of the controls. Furthermore, fluid ammonium concentrations increased greatly after X. beaveri larvae metamorphosed, although the cause of this increase could not be identified. Our results show that a terrestrial, inquiline predator can contribute significantly to nutrient sequestration in the phytotelma it inhabits, and suggest that this interaction has a net mutualistic outcome for both species.

The metagenome of bromeliads phytotelma in Puerto Rico.

Bromeliads tank water or phytotelma is an eutrophic microenvironment where microorganisms have evolved to resist sudden changes in pH and nutritional competition. Metagenomics studies have been poorly studied in bromeliads and environmental DNA (eDNA) characterization for its microenvironment is deficient in Puerto Rico. Therefore, the data represents the microbial communities inhabiting bromeliads phytotelma. eDNA was extracted using Metagenomic DNA Isolation Kit for Water. Next-Generation-Sequencing technology (Illumina MiSeq) was used for sequencing the isolated eDNA. This data provides an insight about diversity and functional depiction of microorganisms inhabiting bromeliads phytotelma. The data of this metagenome is available in the BioSample Submission Portal as Bioproject PRJNA39461 and Sequence Read Archive (SRA) accession number SRP114300. MG-RAST metagenomic analysis server is located under the study ID mgp79812.

Aquatic invertebrates associated with bromeliads in Atlantic Forest fragments / Invertebrados associados a bromélias em fragmentos de Mata Atlântica

Abstract: Forest fragments in the state of Minas Gerais contain a large number of Bromeliaceae genera, whose cisterns accumulate water and organic matter, providing shelter and food for a wide range of organisms. However, these fragments often consist only of small patches of vegetation, making the species more vulnerable to the effects of this landscape changes. This study aimed to test the effect of the distance to the edge and of the matrix type on the structure of aquatic invertebrate communities in four morphospecies of bromeliads. Samples were collected in Atlantic Forest fragments adjacent to pasture and planted forest areas, in a region of Serra da Mantiqueira, Brazil. Of the 147 bromeliads investigated, we found 35 taxa, among which the most abundant groups were Ostracoda (4,962 individuals), Culicidae (2,358), Tanypodinae (1,164) and Scirtidae (1,043). The richness of taxa and Shannon diversity of invertebrates were similar at different distances from the edge, with variation of richness between 17 and 23 taxa and diversity between 1.25 and 1.52. The composition of the fauna in the cisterns changed between some collection sections (A vs. C, C vs. D and D vs. F), irrespective of being close to or far from the edge. There was no variation of richness (t=-1.145, df=106, p= 0.341) and diversity (t= 1.376, df= 106, p= 0.429) among samples collected from fragments next to planted forest and pasture, likely because the bromeliads studied were located on hillsides above the canopy of planted forests, and subject to similar conditions to those found in the fragments next to pastures. The results demonstrate the importance of bromeliads in maintaining the richness and diversity of the invertebrates they harbor, even in altered landscapes exposed to extreme conditions such as fragment edges.

Resumo: Remanescentes florestais em Minas Gerais concentram grande número de gêneros de Bromeliaceae, cujos taques acumulam água e a matéria orgânica que servem de abrigo e fonte de alimento para uma grande variedade organismos. Entretanto, tais áreas têm sido reduzidas a pequenas manchas de vegetação, tornando as espécies mais vulneráveis aos efeitos decorrentes dessa modificação da paisagem. O presente estudo visou caracterizar a estrutura das comunidades de invertebrados aquáticos em quatro morfoespécies de bromélias. As amostras foram coletadas em fragmentos de Mata Atlântica em matriz de pastagem e floresta plantada, em uma região da Serra da Mantiqueira, Brasil. Nas 147 bromélias investigadas foram encontrados 35 táxons, entre os quais os grupos mais abundantes foram Ostracoda (4,962 individuals), Culicidae (2,358), Tanypodinae (1,164) e Scirtidae (1,043). A riqueza de táxons e diversidade de Shannon foram semelhantes em diferentes distâncias da borda, com variação entre 17 e 23 táxons e diversidade entre 1.25 e 1.52. A composição da fauna nos tanques variou entre algumas faixas de coleta (A vs. C, C vs. D e D vs. F), independentemente de estarem próximas ou distantes da borda. Não houve variação da riqueza (t=-1.145, df=106, p= 0.341) e diversidade (t= 1.376, df= 106, p= 0.429) entre as amostras coletadas em fragmentos próximos à floresta plantada e pastagem, provavelmente devido às bromélias estudadas estarem localizadas em encostas acima do dossel das florestas plantadas, e sujeitas a condições semelhantes às encontradas nos fragmentos ao lado de pastagens. Foi possível demonstrar a importância das bromélias na manutenção da riqueza e da diversidade de invertebrados que abrigam, mesmo em paisagens alteradas e expostas a condições extremas, tais como nas bordas dos fragmentos.

Flagellate protist abundance in phytotelmata of Aechmea distichantha Lem. (Bromeliaceae) in the upper Paraná river basin / Abundância de protozoários flagelados em fitotelmata de Aechmea distichantha Lem. (Bromeliaceae) na bacia do alto rio Paraná

In some Bromeliaceae species the leaf sheaths form a tank or phytotelma, accumulating water and organic detritus, conditions that allow the establishment of different communities. Thus, this study evaluated the relationship between diameter and volume of water in the tank of the bromeliad Aechmea distichantha Lem. and density of flagellates found in the phytotelmata, as well as the influence of the hydrological periods and proximity of sampled strata on the phytotelma patterns of abundance distribution. Samplings were conducted in two hydrological periods in 2010. In each sampling, four plants were collected from three different strata, classified according to the distance from the river level. A one-way ANOVA identified significant differences in flagellate density between hydrological periods. The influence of the water volume in the phytotelmata on the flagellate density was evidenced by regression analyses, as well as the lack of relationship between the density and perimeter of the tank. Moreover, the distance of the strata from the river level did not influence the pattern of flagellate abundance. Thus, our results suggested that the hydrological period and the volume of water in the tanks of Aechmea distichantha are determinant on the abundance of heterotrophic flagellate in the phytotelmata.

Em algumas Bromeliaceae as bainhas foliares formam uma cisterna ou fitotelma, onde ocorre acúmulo de água e detritos orgânicos, que permitem o estabelecimento de diferentes comunidades. Assim, este estudo avaliou a relação entre o diâmetro e volume de água no tanque de Aechmea distichantha Lem. e a abundância de flagelados encontrados nestes fitotelmata, bem como a influência do período hidrológico e a proximidade dos estratos amostrados sobre os padrões de distribuição desse atributo. As coletas foram realizadas em dois períodos hidrológicos no ano de 2010. Foram coletadas quatro plantas em três estratos diferentes de acordo com a distância em relação ao rio. Foram identificadas diferenças significativas entre a densidade dos flagelados e os períodos hidrológicos. Além disso, verificou-se a influência do volume de água presente nos fitotelmata sobre a densidade dos flagelados, bem como a ausência de relação entre a densidade desses protistas e o perímetro do tanque das bromélias. A distância dos distintos estratos em relação ao rio não influenciou a abundância dos flagelados. Assim, os resultados sugerem que o período hidrológico bem como o volume de água presente nos tanques de Aechmea distichantha atuam como um fator determinante sobre a abundância da comunidade de flagelados heterotróficos nos fitotelmata.