Effect of tank bromeliad micro-environment on <Emphasis Type="Italic">Aedes aegypti</Emphasis> larval mortality

Many species of bromeliads create an aquatic microcosm among their leaves. Besides their native aquatic fauna, these microcosms can be used by larvae of invasive mosquitoes like Aedes aegypti. We compared the mortality among A. aegypti larvae placed inside tanks of Aechmea fasciata bromeliads with larvae placed inside artificial microcosms and with microcosms with low pH (5.4), which simulate the acidic conditions found inside bromeliad tanks. A. aegypti larvae suffered a significantly higher mortality inside bromeliad tanks compared to larvae in control microcosms, but the mortality inside bromeliads did not differ statistically from that found in artificial microcosms simulating bromeliad acidic conditions. We concluded that bromeliad tanks tend to be a less suitable environment for the development of A. aegypti larvae than artificial containers due to the acidification generated by bromeliad physiology.     

Calling site selection by the bromeliad-dwelling treefrog Phyllodytes melanomystax (Amphibia: Anura: Hylidae) in a coastal sand dune habitat

We studied bromeliad selection by calling males of Phyllodytes melanomystax. The study site was a restinga environment in the northeastern state of Bahia, northeastern Brazil. We sampled 202 bromeliads, 101 with and 101 without calling males. We used multiple logistic regression analysis and Wald test to identify which of nine environmental variables investigated could explain the occurrence of calling males within bromeliads. The presence/absence of calling males in bromeliads was influenced by the number of bromeliads in a 2 m radius and the amount of debris inside the rosettes, while physical variables of bromeliads and the volume of stored water inside their rosettes had no influence. The mark-recapture procedure of P. melanomystax revealed site fidelity. This study is the first to explain the pattern of bromeliad selection by a species of the bromeliad-dwelling frog genus Phyllodytes.     

Understorey environments influence functional diversity in tank‐bromeliad ecosystems

1. A substantial fraction of the freshwater available in neotropical forests is impounded within the rosettes of bromeliads that form aquatic islands in a terrestrial matrix. The ecosystem functioning of bromeliads is known to be influenced by the composition of the contained community but it is not clear whether bromeliad food webs remain functionally similar against a background of variation in the understorey environment. 2. We considered a broad range of environmental conditions, including incident light and incoming litter, and quantified the distribution of a very wide range of freshwater organisms (from viruses to macroinvertebrates) to determine the factors that influence the functional structure of bromeliad food webs in samples taken from 171 tank‐bromeliads. 3. We observed a gradient of detritus‐based to algal‐based food webs from the understorey to the overstorey. Algae, rotifers and collector and predatory invertebrates dominated bromeliad food webs in exposed areas, whereas filter‐feeding insects had their highest densities in shaded forest areas. Viruses, bacteria and fungi showed no clear density patterns. Detritus decomposition is mainly due to microbial activity in understorey bromeliads where filter feeders are the main consumers of microbial and particulate organic matter (POM). Algal biomass may exceed bacterial biomass in sun‐exposed bromeliads where amounts of detritus were lower but functional diversity was highest. 4. Our results provide evidence that tank‐bromeliads, which grow in a broad range of ecological conditions, promote aquatic food web diversity in neotropical forests. Moreover, although bromeliad ecosystems have been categorised as detritus‐based systems in the literature, we show that algal production can support a non‐detrital food web in these systems.     

Relationship between invertebrate fauna and bromeliad size.

Several bromeliads species store water and organic substrates, allowing the establishment of phytotelmata and associated fauna on their leaves. In this study, we sampled 70 individuals of Vriesea sp. (Carrière) (Bromeliaceae), in rupestrian fields in the Serra de Ouro Branco-MG, Brazil. The relationships between invertebrate species richness and abundance and size of bromeliads were tested using multiple regression. We found 19 species associated with bromeliads, mainly Diptera larvae. The abundance of the phytotelmate fauna increased principally in relation to the volume of water in the bromeliad reservoir. Phytotelmata richness was affected principally by diameter of the reservoir. There was a significant relationship between the abundance and richness of invertebrates associated with leaves with diameter and height of the plant. Invertebrate richness was better explained by abundance of individuals. These results suggest that the increase of richness was attended by higher numbers of microhabitats and more space for colonization of bigger bromeliads. Additionally, there was more chance of sampling different species in locales with greater abundance of individuals.     

Changes in rainfall level and litter stoichiometry affect aquatic community and ecosystem processes in bromeliad phytotelmata

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Aquatic invertebrate communities in tank bromeliads: how well do classic ecological patterns apply?

Tank bromeliads (Bromeliaceae) often occur in high densities in the Neotropics and represent a key freshwater habitat in montane forests, housing quite complex invertebrate communities. We tested the extent to which there are species richness–altitude, richness–environment, richness–size, richness–habitat complexity and richness–isolation relationships for the aquatic invertebrate communities from 157 bromeliads in Cusuco National Park, Honduras. We found that invertebrate species richness and abundance correlated most strongly, and positively, with habitat size, which accounted for about a third of the variance in both. Apart from bromeliad size (equivalent of the species–area relationship), we found remarkably little evidence of classic biogeographic and ecological relationships with species richness in this system. Community composition correlated with altitude, bromeliad size and position, though less than 20% of the variation was accounted for by the tested variables. The turnover component of dissimilarity between the communities correlated with altitude, while the nestedness-resultant component was related to bromeliad size. The unexplained variance could reflect a large stochastic component in the system, associated with the ephemerality of the habitat patches (both the plants themselves and the fluctuations in their water content) and stochasticity due to the dispersal dynamics in the system. We conclude that there is a small contribution of classic biogeographic factors to the diversity and community composition of aquatic invertebrates communities in bromeliads. This may be due to the highly dynamic nature of this system, with small patch sizes and high emigration rates. The patterns may mostly be driven by factors affecting colonisation success.     

Oviposition site selection by the bromeliad-dweller harvestman <Emphasis Type="Italic">Bourguyia</Emphasis><Emphasis Type="Italic">hamata</Emphasis> (Arachnida: Opiliones)

Bourguyia hamata females oviposit almost exclusively inside the rosette formed by the curled leaves of the epiphytic bromeliad Aechmea nudicaulis. We investigated whether the architecture of the individual bromeliads influences oviposition site selection by this harvestman species. We collected data on the presence of clutches inside bromeliads, rosette length, rosette slope in relation to tree trunks, and the amount of debris inside the rosette. Additionally, we measured the water volume inside the rosettes as well as the variation in the humidity inside and outside bromeliads with long and short rosettes. Longer rosettes were preferred as oviposition site possibly because they accumulate more water and maintain lower internal humidity variation than the external environment. Although the slope of the rosettes did not influence the occurrence of oviposition, the probability of debris accumulation inside the rosettes increased with their slope, and the frequency of clutches was greater in bromeliads with small amounts of debris. A field experiment showed that bromeliads with water inside the rosette were more frequently used as oviposition sites than bromeliads without water. In conclusion, females oviposit predominantly in bromeliads that accumulate more water and have small amounts of debris inside the rosettes, probably because these characteristics promote a more adequate microhabitat for egg development.     

Environmental drivers of invertebrate population dynamics in Neotropical tank bromeliads

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Host bromeliads and their associated frog species: Further considerations on the importance of species interactions for conservation

Bromeliads constitute a good example of symbiosis with organisms that spend their entire life cycle inside the plants, and often depend on them to breed. The bromeliads benefit from this interaction by increasing their nutrients intake. Conservation efforts tend to focus on a single endangered species, but in symbiotic associations, the viability of one species depends on that of the other. Based on IUCN criteria, any species that depends on another to complete its life cycle should be assigned a conservation status equivalent to that of the host taxon, where appropriate. We gathered published plus fieldwork data on the frog-bromeliad mutualism and compiled a checklist of 99 bromeligenous frogs species associated to 69 bromeliad hosts, and found threatened bromeliads hosting non-threatened frogs. We found that 62% bromeligenous frogs inhabit unspecified bromeliads. Finally, we propose strategies for improving understanding and conservation of the frog-bromeliad mutualism.     

Two coexisting tank bromeliads host distinct algal communities on a tropical inselberg

The tank bromeliads Aechmea aquilega (Salisb.) and Catopsis berteroniana (Schultes f.) coexist on a sun‐exposed Neotropical inselberg in French Guiana, where they permit conspicuous freshwater pools to form that differ in size, complexity and detritus content. We sampled the algal communities (both eukaryotic and cyanobacterial taxa, including colourless forms) inhabiting either A. aquilega (n = 31) or C. berteroniana (n = 30) and examined differences in community composition and biomass patterns in relation to several biotic and abiotic variables. Chlorella sp. and Bumilleriopsis sp. were the most common taxa and dominated the algal biomass in A. aquilega and C. berteroniana, respectively. Using a redundancy analysis, we found that water volume, habitat complexity and the density of phagotrophic protozoa and collector‐gatherer invertebrates were the main factors explaining the distribution of the algal taxa among the samples. Hierarchical clustering procedures based on abundance and presence/absence data clearly segregated the samples according to bromeliad species, revealing that the algal communities in the smaller bromeliad species were not a subset of the communities found in the larger bromeliad species. We conclude that, even though two coexisting tank bromeliad populations create adjacent aquatic habitats, each population hosts a distinct algal community. Hence, bromeliad diversity is thought to promote the local diversity of freshwater algae in the Neotropics.     

Heterogeneity of terrestrial bromeliad colonies and regeneration of Acacia praecox (Fabaceae) in a humid-subtropical-Chaco forest, Argentina

In several tropical and subtropical forests, plants of the understorey act as an ecological filter that differentially affects woody species regeneration. In convex sectors of the Schinopsis balansae (Anacardiaceae) forests of the Southeastern Chaco there are dense colonies of terrestrial bromeliads. These may influence forest regeneration by intercepting rain water and propagules in their tanks. Within colonies, the spatial distribution of bromeliads is clumped because their clonal growth leaves numerous internal gaps. In this study we describe the internal heterogeneity of three bromeliad colonies (plots) and analyze how this heterogeneity affects Acacia praecox regeneration (i.e. seedling recruitment and survival). In January 1996, we randomly placed three transects with 150 contiguous quadrats of 100 cm(2) in each plot. For each quadrat we recorded the type of floor cover (i.e. bromeliads, herbs, litter, or bare soil) and the presence of A. praecox seeds or seedlings. In July 1996 we relocated the transects and recorded seedling survival. Bromeliad colonies showed a high internal heterogeneity. Almost half of the 450 quadrats were covered by two terrestrial bromeliads. Aechmea distichantha was recorded in 81% of all quadrats with bromeliads, and Bromelia serra in the others. All quadrats with bromeliads were covered by litter. Half of them were occupied by the bases of bromeliads and the others were covered by their leaves. In contrast, where bromeliads were not present, soil surface was covered by litter in 83% and by herbaceous vegetation in 11% of the quadrats; very few quadrats were covered by bare soil. In January 1996, we recorded 127 seeds and 176 seedlings of A. praecox. Seed and seedling densities of A. praecox were similar in quadrats with and without bromeliads, but variability in seedling density of A. praecox was higher within than among plots. Seed density was higher in quadrats covered by bromeliad leaves than inside the tanks. Seedling survival of A. praecox was slightly higher in quadrats with bromeliads in only one of the three plots. No seedling survived inside the bromeliad tanks. Apparently. bromeliad colonies have no effect on seedling regeneration of A. praecox.     

Ecosystem services provided by bromeliad plants: A systematic review

The unprecedented loss of biological diversity has negative impacts on ecosystems and the associated benefits which they provide to humans. Bromeliads have high diversity throughout the Neotropics, but they have been negatively affected by habitat loss and fragmentation, climate change, invasive species, and commercialization for ornamental purpose. These plants provide direct benefits to the human society, and they also form microecosystems in which accumulated water and nutrients support the communities of aquatic and terrestrial species, thus maintaining local diversity. We performed a systematic review of the contribution of bromeliads to ecosystem services across their native geographical distribution. We showed that bromeliads provide a range of ecosystem services such as maintenance of biodiversity, community structure, nutrient cycling, and the provisioning of food and water. Moreover, bromeliads can regulate the spread of diseases, and water and carbon cycling, and they have the potential to become important sources of chemical and pharmaceutical products. The majority of this research was performed in Brazil, but future research from other Neotropical countries with a high diversity of bromeliads would fill the current knowledge gaps and increase the generality of these findings. This systematic review identified that future research should focus on provisioning, regulating, and cultural services that have been currently overlooked. This would enhance our understanding of how bromeliad diversity contributes to human welfare, and the negative consequences that loss of bromeliad plants can have on communities of other species and the healthy functioning of the entire ecosystems.     

How effective are non-destructive sampling methods to assess aquatic invertebrate diversity in bromeliads?

With the growing interest in small aquatic water bodies, especially as naturally replicated model systems for ecological research, aquatic invertebrate communities in phytotelmata are increasingly receiving attention these days. The recognition of the substantial contribution to the regional species pool of specialised species draws further attention to these small and often temporary habitats. The methods currently used for studying communities in some types of phytotelma, such as bromeliads, tend to be destructive, typically involving complete dissection of the plant. The expected increase in sampling intensity associated with the increasing interest in phytotelmata may result in a negative impact on plant populations in some areas, decreasing numbers in an unsustainable way, especially in locations with ongoing, intensive research. We therefore aimed to investigate whether less-destructive sampling methods can achieve sufficient data quality to allow their use as alternatives to complete plant dissection. We tested the effectiveness of three such methods in measuring the aquatic invertebrate communities in tank bromeliads (Tillandsia guatemalensis) in Cusuco National Park, Honduras. The three methods were pipetting the water out of the bromeliad, turning the bromeliad upside down and dissecting only the outer part of the plant (the oldest, often deteriorating leaves). Overall, we found that these methods were poor predictors of richness and abundance of the organisms in communities. However, we found big differences between taxonomic groups, depending in part on the ecology of the organisms, and we suggest that some less-destructive alternative methods may be appropriate for studying some specific groups (e.g. Culicidae). Based on these results and a rapid survey of the abundance of bromeliads in the national park, we question whether intensive, ongoing research into aquatic invertebrate communities in similar phytotelma populations is sustainable. From the point of view of conservation, alternative model systems need to be found.     

Selecting a Hiding Place: Anuran Diversity and the use of Bromeliads in a Threatened Coastal Sand Dune Habitat in Brazil

Among vertebrates, anuran amphibians represent the highest number of species associated with bromeliads and possess a range of ecological, behavioral, and morphological specializations to life in these plants. Despite the importance of bromeliads as biodiversity amplifiers, and their diversity in some habitats, studies of the relationship between anurans and these plants are scarce in Brazil. Here, we investigated the way anurans select and use bromeliads in a threatened coastal habitat. We analyzed data from 23 standardized samples of the anurans associated with the bromeliad Neoregelia cruenta in the Restinga de Maricá, State of Rio de Janeiro, Southeastern Brazil. We found nine anuran species using these bromeliads, representing the highest richness reported for a Brazilian restinga. We identified a general pattern of bromeliad usage, where plants located at the edges of scrub patches (exposed to the sun) were more frequently occupied by anurans than those located more to the center (in the shade). There is strong evidence of an active selective process based on the quality of the water stored in the rosette, which differs between plants depending on their position in the scrub patch. Although the number of individuals varied during the period of study, the frequency of bromeliads used was constant, indicating that plant occupation follows a regular pattern throughout the year. Furthermore, the high frequency of bromeliads used by anurans during the whole year highlights the importance of considering these plants in developing conservation programs concerning the protection of anurans.     

Does the strength of cross‐ecosystem trophic cascades vary with ecosystem size? A test using a natural microcosm

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Water-filled bromeliad as roost site of a tropical lizard,Urostrophus vautieri (Sauria: Leiosauridae)

Bromeliads are a conspicuous component of tropical forests. Whereas several amphibian species are closely associated with bromeliads, reptiles are much less frequently observed in bromeliads and only a few species use bromeliads for egg deposition or as roost site. We report on an adult Urostrophus vautieri that was sleeping in a water-filled bromeliad. The individual was submerged except for head and shoulder. To our knowledge, it is the first time that such behavior has been observed in an arboreal Neotropical lizard.     

Phoretic Behaviour of Bromeliad Annelids (<Emphasis Type="Italic">Dero</Emphasis>) and Ostracods (<Emphasis Type="Italic">Elpidium</Emphasis>) using Frogs and Lizards as Dispersal Vectors

The phoretic behaviour of ostracods (Elpidium bromeliarum) andannelids (Dero superterrenus) that inhabit tank bromeliads was studied. Our previous field observations had shown that bromeliad ostracods can be found attached to the skin of amphibians and reptiles that move among bromeliads, probably allowing the ostracods to colonise new tanks. In this paper, we present the first record of bromeliad annelids found attached to frogs moving among bromeliads in the field. We have also enlarged the database on bromeliad ostracods engaged in phoretic association with terrestrial vertebrates in three locations in southeastern Brazil. In our laboratory experiments bromeliad annelids show a strong significant tendency to climb onto papers that had been in contact with frog skin when compared with control papers, indicating a kind of chemically oriented behaviour. Bromeliad ostracods, on the other hand, attached themselves to treated and untreated papers with same frequency. When brought into contact with various species of frogs and lizards, the bromeliad annelids and ostracods both presented preference to attach themselves to frogs, but the annelids showed a stronger preference to attach to frogs and to avoid attachment to lizards. Another experiment demonstrated that bromeliad annelids are much more prone to dehydration than are ostracods. We suggest that the chemically oriented behaviour presented by bromeliad annelids toward frogs could diminish the risk of death by dehydration during the transport among bromeliads due to the moist characteristic of frog skins.     

Two bromeliad species with contrasting functional traits partition the understory space in a South American xerophytic forest: correlative evidence of environmental control and limited dispersal

We examined the spatial distribution of two bromeliad species with contrasting functional traits in the understory of a xerophytic South American Chaco forest. Aechmea distichantha is a facultative terrestrial species with well-developed phytotelma and short rhizomes, whereas Bromelia serra is a strictly terrestrial species with soil-exploring roots and long rhizomes. Both bromeliads develop colonies on relatively elevated patches in Schinopsis balansae forests. We evaluated the roles of environmental controls, limited dispersal, and interspecific competition as drivers of the different distribution of these bromeliads. We mapped the overstory, understory and topography of 16 forest plots with bromeliads (400 m² each, subdivided in 100 4-m² subplots). We sampled soil characteristics on sectors dominated by each bromeliad species. We used structural equation modeling to assess direct and indirect associations of each bromeliad species cover with environmental conditions, abundance of conspecifics in the vicinity, and local abundance of the other species. A. distichantha cover increased on elevated subplots with high tree/shrub basal area, whereas B. serra cover showed the opposite pattern. In addition, A. distichantha cover was negatively associated with B. serra cover, but not vice versa, and cover of both species increased with the abundance of nearby conspecifics, suggesting that limited vegetative dispersal partly accounted for their distribution. Sectors dominated by A. distichantha had lower soil bulk density and higher organic matter content than those dominated by B. serra. According to our model, influences of competition and limited vegetative dispersal reinforce the association between distribution of these bromeliads and environmental heterogeneity of the forest understory.     

Are Algae Relevant to the Detritus-Based Food Web in Tank-Bromeliads?

We assessed the occurrence of algae in five species of tank-bromeliads found in contrasting environmental sites in a Neotropical, primary rainforest around the Nouragues Research Station, French Guiana. The distributions of both algal abundance and biomass were examined based on physical parameters, the morphological characteristics of bromeliad species and with regard to the structure of other aquatic microbial communities held in the tanks. Algae were retrieved in all of the bromeliad species with mean densities ranging from ～10(2) to 10(4) cells/mL. Their biomass was positively correlated to light exposure and bacterial biomass. Algae represented a tiny component of the detrital food web in shaded bromeliads but accounted for up to 30 percent of the living microbial carbon in the tanks of Catopsis berteroniana, located in a highly exposed area. Thus, while nutrient supplies are believed to originate from wind-borne particles and trapped insects (i.e., allochtonous organic matter), our results indicate that primary producers (i.e., autochtonous organic matter) are present in this insectivorous bromeliad. Using a 24-h incubation of size-fractionated and manipulated samples from this plant, we evaluated the impact of mosquito foraging on algae, other microorganisms and rotifers. The prey assemblages were greatly altered by the predation of mosquito larvae. Grazing losses indicated that the dominant algal taxon, Bumilleriopsis sp., like protozoa and rotifers, is a significant part of the diet of mosquito larvae. We conclude that algae are a relevant functional community of the aquatic food web in C. berteroniana and might form the basis of a complementary non-detrital food web.     

Metacommunity size influences aquatic community composition in a natural mesocosm landscape

Ecosystems are often arranged in naturally patchy landscapes with habitat patches linked by dispersal of species in a metacommunity. The size of a metacommunity, or number of patches, is predicted to influence community dynamics and therefore the structure and function of local communities. However, such predictions have yet to be experimentally tested using full food webs in natural metacommunities. We used the natural mesocosm system of aquatic macroinvertebrates in bromeliad phytotelmata to test the effect of the number of patches in a metacommunity on species richness, abundance, and community composition. We created metacommunities of varying size using fine mesh cages to enclose a gradient from a single bromeliad up to the full forest. We found that species richness, abundance, and biomass increased from enclosed metacommunities to the full forest size and that diversity and evenness also increased in larger enclosures. Community composition was affected by metacommunity size across the full gradient, with a more even detritivore community in larger metacommunities, and taxonomic groups such as mosquitoes going locally extinct in smaller metacommunities. We were able to divide the effects of metacommunity size into aquatic and terrestrial habitat components and found that the importance of each varied by species; those with simple life cycles were only affected by local aquatic habitat whereas insects with complex life cycles were also affected by the amount of terrestrial matrix. This differential survival of obligate and non‐obligate dispersers allowed us to partition the beta‐diversity between metacommunities among functional groups. Our study is one of the first tests of metacommunity size in a natural metacommunity landscape and shows that both diversity and community composition are significantly affected by metacommunity size. Synthesis Natural food webs are sensitive to meta‐community size, i.e. the number of patches connected through dispersal. We provide an empirical test using the aquatic foodweb associated within bromeliads as a model system. When we reduced the number of bromeliad patches connect through dispersal, we found a clear change of the foodweb in terms of population sizes, beta diversity, community composition and predator‐prey ratios. The response of individual taxa was predictable based on species traits including dispersal modes, life cycle, and adult resource requirements. Our study demonstrates that community structure is strongly influenced by the interplay of species traits and landscape properties.