Tank Bromeliads Research Articles

Influence of isolation on the resilience of tank bromeliad ecosystems to drought in a Neotropical rainforest

AbstractLittle is known of how Neotropical freshwater ecosystems will respond to future climate scenarios. In Neotropical rainforests, a substantial fraction of the freshwater available to the aquatic fauna is found within phytotelmata, plant-held waters that form aquatic islands in a terrestrial matrix. We hypothesized that phytotelmata in close proximity have higher resilience capacity to severe drought than the isolated ones, under the assumption that immigration from nearby sources promotes faster recovery. We used rainshelters to emulate an extreme drought (67 days without rainfall) in tank-forming bromeliads arranged in patches of 1, 3 or 6 plants in a primary forest of French Guiana. Habitat size was a stronger determinant of invertebrate species richness and biomass per bromeliad than patch size. Larger bromeliad patches attenuated the adverse effect of drought on the biomass of predators, probably because short-range migration within dense patches allowed individuals to find moist refuges. However, the recovery of aquatic communities and ecosystem functions was mostly supported by in situ resistance, and a rescue effect of immigration was weak. Whilst environmental management plans tend to focus on dense networks of connected water bodies, our study shows that efforts should not omit the isolated ones.

Tank formation transforms nitrogen metabolism of an epiphytic bromeliad and its phyllosphere bacteria.

Up to half of tropical forest plant species grow on other plants. Lacking access to soils, vascular epiphytes have unique adaptations for mineral nutrition. Among the most distinctive is the tank growth form of certain large bromeliads, which absorb nutrients that are cycled by complex microbial communities in water trapped among their overlapping leaf bases. However, tanks form only after years of growth by juvenile plants, which must acquire nutrients differently. Understanding how nutrient dynamics change during tank bromeliad development can provide key insights into the role of microorganisms in the maintenance of tropical forest biodiversity. We evaluated variations in plant morphology, growth, foliar nitrogen physiology, and phyllosphere bacterial communities along a size gradient spanning the transition to tank formation in the threatened species Tillandsia utriculata. Sequential morphological and growth phases coincided with the transition to tank formation when the longest leaf on plants was between 14 and 19 cm. Before this point, foliar ammonium concentrations were very high, but after, leaf segments absorbed significantly more nitrate. Leaf-surface bacterial communities tracked ontogenetic changes in plant morphology and nitrogen metabolism, with less-diverse communities in tankless plants distinguished by a high proportion of taxa implicated in ureolysis, nitrogen fixation, and methanotrophy, whereas nitrate reduction characterized communities on individuals that could form a tank. Coupled changes in plant morphology, physiology, and microbiome function facilitate the transition between alternative nutritional modes in tank bromeliads. Comparing bromeliads across life stages and habitats may illuminate how nitrogen-use varies across scales.

New insights into the role of the root system of epiphytic bromeliads: comparison of root and leaf trichome functions in acquisition of water and nutrients.

In epiphytic bromeliads, the roots used to be considered poorly functional organs in the processes of absorption and metabolization of water and nutrients, while the leaves always acted as protagonists in both functions. More recent discoveries have been changing this old view of the root system. In this review, we will address the old thoughts of the scientific community regarding the function performed by the roots of epiphytic bromeliads (mere holdfast structures with low physiological activity) and the importance of a reduced or lack of root system for the emergence of epiphytism. We will present indirect and direct evidence that contradicts this older hypothesis. Furthermore, the importance of the root absorptive function mainly for juvenile tankless epiphytic bromeliads and the characteristics of the root absorption process of adult epiphytic tank bromeliads will be thoroughly discussed in physiological aspects. Finally, some factors (species, substrate, environmental conditions) that influence the absorptive capability of the roots of epiphytic tank bromeliads will also be considered in this review, highlighting the importance that the absorptive role of the roots have for the plasticity of bromeliads that live on trees, which is an environment characterized by the intermittent availability of water and nutrients. The roots of tank-forming epiphytic bromeliads play important roles in the absorption and metabolization of nutrients and water. The importance of roots stands out mainly for juvenile tankless bromeliads since the root is the main absorptive organ. In larger plants with tank, although the leaves become the protagonists in the resource acquisition process, the roots complement the absorptive function of the leaf trichomes, resulting in a better growth of the bromeliad. The physiological and biochemical properties of the processes of absorption and distribution of resources in the tissues seem to differ between absorption by trichomes and roots.

Diversity and relative abundance of immature mosquitoes (Diptera: Culicidae) in Hohenbergia catingae Ule bromeliads and larvitraps in the Nísia Floresta National Forest, northeast Brazil.

The Nísia Floresta National Forest, located in Rio Grande do Norte (RN), is an important remnant of the Atlantic Forest biome in Brazil. Bromeliad tanks in this forest offer suitable breeding sites for mosquito species that may act as viral vectors, thus posing an epidemiological concern. However, studies investigating the presence of immature Culicidae in natural breeding sites in RN have thus far been restricted to Caatinga vegetation. This study investigated mosquitoes and their natural breeding sites in bromeliads growing in the Nísia Floresta National Forest. From March 2013 to February 2014, monthly samples were collected from the tanks of five randomly selected bromeliads and larvitraps placed in each of the three forest management areas. Hohenbergia catingae Ule (Hohenbergia bromeliad) is an important shelter for immature mosquitoes. Culex (Microculex) was the predominant species, representing 86% of the immature mosquitoes collected. A rare occurrence of Aedes (Stegomyia) aegypti (Linnaeus, 1762) (generally associated with urban areas under high anthropogenic influence) was observed, highlighting the importance of investigating the presence of mosquitoes in different natural habitats. An analysis of species diversity revealed that species such as Culex imitator Theobald, 1903 and Culex davisi Kumm, 1933, have a strong association with bromeliads. In tire traps (larvitraps) Aedes (Stegomyia) albopictus Skuse, 1894 was predominant. Environmental changes, such as deforestation, removal of bromeliads, and climate change in the area, can influence the migration of species and adaptation to new habitats in a peridomiciliary environment around the forest, consequently the possibility of transmission of virus and other pathogens.

A Neoponera villosa (Fabricius, 1804) Worker Pupa with too Many Legs (Hymenoptera: Formicidae).

Several somatic and sexual anomalies such as gynandromorphism and other morphological malformations have been described in ants, but cases of supernumerary legs in natural populations are rare, with only six cases reported to date. Moreover, few cases of abnormalities have been reported for the Ponerinae subfamily, with only three cases of genetic chimeras and five teratological cases worldwide. We report here a new case of teratology in this subfamily, with a seven-legged worker pupa found in a Neoponera villosa colony nesting in an epiphytic tank bromeliad. This is the second case for Mexico of a morphological anomaly in a ponerine ant, but the first report of a teratological case in the subfamily for this country.

Functional traits of predators and decomposer prey determine context dependency in trophic control over ecosystems.

Research Highlight: Piccoli, G. C. d. O., Antiqueira, P. A. P., Srivastava, D. S., & Romero, G. Q. (2024). Trophic cascades within and across ecosystems: The role of anti-predatory defences, predator type and detritus quality. Journal of Animal Ecology, 00, 1-14. https://doi.org/10.1111/1365-2656.14063. Ecosystem functioning is controlled by the interplay between bottom-up supply of limiting nutrients and top-down animal feedback effects. However, the degree of animal versus nutrient control is context-dependent. A key challenge lies in characterizing this context dependency which is hypothesized to depend on differences in animal functional traits. Reporting on an important experiment, Piccoli etal. (2014) evaluate how interactions among functionally different predators and decomposer prey create context dependency in top-down control of a model system-tropical bromeliad tank ecosystems. Bromeliad plants hold water in their tanks supporting microcosm ecosystems containing terrestrial and aquatic insect larvae and arachnids. The ecosystems are supported by nutrients in plant litter that rains down from forest canopies into the tanks. Nutrients are released after litter is decomposed by a functionally diverse community of larval insect decomposers that differ in feeding mode and antipredator defence strategy. This decomposer community is preyed upon by an exclusively narrowly ranging aquatic insect larval predator and widely ranging spider predator that crosses between the aquatic and surrounding terrestrial ecosystems. Experimental manipulation of the animal community to test for the degree of control by predators mediated by the functionally diverse prey community included four treatments: (i) a control with the detritivores composing different function groups but without predators, (ii) the cross-ecosystem spider predator added, (iii) the purely aquatic damselfly larvae predator added and (iv) both predator types added to capture their interacting effect on ecosystem function (decomposition, nutrient release, and plant growth). Notably, the study resolved the causal pathways and strengths of direct and indirect control using structural equation modelling. These findings reveal how context dependency arises due to different capacities of the predators alone and together to overcome prey defences and control their abundances, with attendant cascading effects that diminished as well as enhanced decomposition and nutrient release to support bromeliad plant production. The study reveals that predators have a decided, albeit qualitatively and quantitatively different, hand in shaping the degree of bottom-up control through feedback effect on the release of limiting nutrients. This ground-breaking study provides a way forward in understanding the mechanisms determining context dependency in the control over ecosystem functioning.

Interactive effects of drought and deforestation on multitrophic communities and aquatic ecosystem functions in the Neotropics-a test using tank bromeliads.

Together with the intensification of dry seasons in Neotropical regions, increasing deforestation is expected to exacerbate species extinctions, something that could lead to dramatic shifts in multitrophic communities and ecosystem functions. Recent studies suggest that the effects of habitat loss are greater where precipitation has decreased. Yet, experimental studies of the pure and interactive effects of drought and deforestation at ecosystem level remain scarce. Here, we used rainshelters and transplantation from rainforest to open areas of natural microcosms (the aquatic ecosystem and microbial-faunal food web found within the rainwater-filled leaves of tank bromeliads) to emulate drought and deforestation in a full factorial experimental design. We analysed the pure and interactive effects of our treatments on functional community structure (including microorganisms, detritivore and predatory invertebrates), and on leaf litter decomposition in tank bromeliad ecosystems. Drought or deforestation alone had a moderate impact on biomass at the various trophic level, but did not eliminate species. However, their interaction synergistically reduced the biomass of all invertebrate functional groups and bacteria. Predators were the most impacted trophic group as they were totally eliminated, while detritivore biomass was reduced by about 95%. Fungal biomass was either unaffected or boosted by our treatments. Decomposition was essentially driven by microbial activity, and did not change across treatments involving deforestation and/or drought. Our results suggest that highly resistant microorganisms such as fungi (plus a few detritivores) maintain key ecosystem functions in the face of drought and habitat change. We conclude that habitat destruction compounds the problems of climate change, that the impacts of the two phenomena on food webs are mutually reinforcing, and that the stability of ecosystem functions depends on the resistance of a core group of organisms. Assuming that taking global action is more challenging than taking local-regional actions, policy-makers should be encouraged to implement environmental action plans that will halt habitat destruction, to dampen any detrimental interactive effect with the impacts of global climate change.

Trophic cascades within and across ecosystems: The role of anti-predatory defences, predator type and detritus quality.

Species in one ecosystem can indirectly affect multiple biodiversity components and ecosystem functions of adjacent ecosystems. The magnitude of these cross-ecosystem effects depends on the attributes of the organisms involved in the interactions, including traits of the predator, prey and basal resource. However, it is unclear how predators with cross-ecosystem habitat interact with predators with single-ecosystem habitat to affect their shared ecosystem. Also, unknown is how such complex top-down effects may be mediated by the anti-predatory traits of prey and quality of the basal resource. We used the aquatic invertebrate food webs in tank bromeliads as a model system to investigate these questions. We manipulated the presence of a strictly aquatic predator (damselfly larvae) and a predator with both terrestrial and aquatic habitats (spider), and examined effects on survival of prey (detritivores grouped by anti-predator defence), detrital decomposition (of two plant species differing in litter quality), nitrogen flux and host plant growth. To evaluate the direct and indirect effects each predator type on multiple detritivore groups and ultimately on multiple ecosystem processes, we used piecewise structural equation models. For each response variable, we isolated the contribution of different detritivore groups to overall effects by comparing alternate model formulations. Alone, damselfly larvae and spiders each directly decreased survival of detritivores and caused multiple indirect negative effects on detritus decomposition, nutrient cycling and host plant growth. However, when predators co-occurred, the spider caused a negative non-consumptive effect on the damselfly larva, diminishing the net direct and indirect top-down effects on the aquatic detritivore community and ecosystem functioning. Both detritivore traits and detritus quality modulated the strength and mechanism of these trophic cascades. Predator interference was mediated by undefended or partially defended detritivores as detritivores with anti-predatory defences evaded consumption by damselfly larvae but not spiders. Predators and detritivores affected ecosystem decomposition and nutrient cycling only in the presence of high-quality detritus, as the low-quality detritus was consumed more by microbes than invertebrates. The complex responses of this system to predators from both recipient and adjacent ecosystems highlight the critical role of maintaining biodiversity components across multiple ecosystems.

Physiological characterisation of aquatic traps in the epiphytic carnivorous plant Utricularia humboldtii

The suction traps of aquatic Utricularia species are discoid bladders, 1-6 mm long, with their lumen isolated by a mobile trapdoor from the ambient medium. Water is pumped out of the traps to maintain the negative pressure in the reset traps. When a prey animal touches the sensory hairs on the trapdoor it opens, the ambient water along with the prey is sucked inside and the trapdoor is closed again. Utricularia humboldtii is a robust, semiaquatic-epiphytic or terrestrial species from the generic section Orchidioides from South America. The efficiency of its aquatic traps was measured based on trap firing and resetting rates as well as changes in trap thickness due to both mechanically stimulated and spontaneous firings using an electronic sensor. The U. humboldtii traps exhibited relatively low firing and resetting rates. These values are 2 to 20 times lower than those in other aquatic Utricularia species reported in the literature. These results together with the low aerobic respiration rate of sliced traps indicate that the less efficient aquatic traps in U. humboldtii are specialised for catching fine prey items in bromeliad water tanks. A marked lag-period in trap resetting was found to occur during the first 30 min after firing, contradicting the accepted concept of continuous water pumping. Spontaneous trap firings were also observed with the same magnitude as that in stimulated firings. This implies that spontaneous firings occur consistently in all the tested aquatic Utricularia species. However, although they relate to the continuous water pumping mechanism of the traps, their physiological importance has not been fully elucidated.

Tank Bromeliads as a Water Reservoir Used by Humans: An Important Overlooked Ecosystem Service in Xerophytic Forests

The Bromeliaceae are known for providing numerous ecosystem services. Tank bromeliads that collect water in the bases of their leaves have been an important source of water for people in semiarid regions such as the Gran Chaco. In this large area characterized by xerophytic vegetation grows Aechmea distichantha, the only Chaquenian terrestrial tank bromeliad. We carried out a literature review on water consumption from this tank bromeliad by people in this region. Then, we analyzed the variations in the amount of water stored in the tanks of plants from observational studies as well as the effects of the amount and frequency of water addition on the hydrology of water retained in the tank from an irrigation experiment study. We found 85 reports on the consumption of water from A. distichantha plants by people, including several Indigenous Groups, Criollos, and Soldiers. According to these reports, the tank accumulates about a liter of relatively fresh and clear water that lasts for several weeks. Our observational study showed a large variation in the amount of water held in the tank among years and sites, and our irrigation experiment showed that even plants receiving low and unevenly distributed rainfall conserved some water in their tanks. All these results support the importance of this tank bromeliad for surviving when moving in Eastern Chaco during dry periods until the beginning of the twentieth century. At present, it seems that people living in sedentarism no longer frequently use it for water provision, highlighting that the relative importance of a plant species for providing ecosystem services depends on the historical context.

Urbanization and Water Containers Influence the Mosquito Community with Consequences for Aedes aegypti.

We aimed to determine how the degree of urbanization in a Neotropical city influences Aedes aegypti (L.), a pantropical vector of urban yellow fever, dengue, Zika and Chikungunia, via other mosquito species, whether they are competitors or predators, native to the area or invasive. We conducted experiments twice a month during one year in the city of Kourou, French Guiana, on three sites characterized by increasing percentages of imperviousness (i.e., 0.65%, 33.80% and 86.60%). These sites were located in a ≈5ha forest fragment, a residential area with gardens, and in the older part of the city, respectively, and correspond to slightly, moderately and highly urbanized sites. There, we monitored twice a month during one year a total of 108 mosquito communities inhabiting four types of containers (i.e., a tank bromeliad, dry stumps of bamboo, ovitraps and car tires) installed in a random block design. In the tanks of the bromeliad, likely due to the acidity of the water, the immatures of native mosquito species prevailed, particularly Wyeomyia pertinans (Williston) in the slightly urbanized site. The general pattern was very similar in the three other types of containers where Limatus durhamii Théobald dominated in the slightly urbanized site, so that the abundance of Ae. aegypti immatures was low compared to those of native species. Yet, Ae. aegypti strongly dominated in the two more urbanized sites. These findings open up perspectives for vector management, including the conservation and/or the augmentation of natural enemies through modifications to landscape features.

Light intensity mediates phenotypic plasticity and leaf trait regionalization in a tank bromeliad.

Phenotypic plasticity allows plants to cope with environmental variability. Plastic responses to the environment have mostly been investigated at the level of individuals (plants) but can also occur within leaves. Yet the latter have been underexplored, as leaves are often treated as functional units with no spatial structure. We investigated the effect of a strong light gradient on plant and leaf traits and examined whether different portions of a leaf show similar or differential responses to light intensity. We measured variation in 27 morpho-anatomical and physiological traits of the rosette and leaf portions (i.e. base and apex) of the tank bromeliad Aechmea aquilega (Bromeliaceae) when naturally exposed to a marked gradient of light intensity. The light intensity received by A. aquilega had a strong effect on the structural, biochemical and physiological traits of the entire rosette. Plants exposed to high light intensity were smaller and had wider, shorter, more rigid and more vertical leaves. They also had lower photosynthetic performance and nutrient levels. We found significant differences between the apex and basal portions of the leaf under low-light conditions, and the differences declined or disappeared for most of the traits as light intensity increased (i.e. leaf thickness, adaxial trichome density, abaxial and adaxial trichome surface, and vascular bundle surface and density). Our results reveal a strong phenotypic plasticity in A. aquilega, particularly in the form of a steep functional gradient within the leaf under low-light conditions. Under high-light conditions, trait values were relatively uniform along the leaf. This study sheds interesting new light on the functional complexity of tank bromeliad leaves, and on the effect of environmental conditions on leaf trait regionalization.

Underestimated diversity and range size of diving beetles in tank bromeliads—Coleoptera of ‘hygrofloric’ lifestyle (Dytiscidae)

Abstract Tank bromeliads provide a vast amount of stagnant water high up in the Neotropical forest canopy. However, the aquatic macroinvertebrate diversity in this specialized habitat remains poorly explored. Here, we study obligatorily bromeliadicolous species of the diving beetle genus Copelatus Erichson, 1832. We review the known species Copelatus bromeliarum Scott, 1912 (Trinidad and Venezuela) and Copelatus bimaculatus Resende & Vanin, 1991 (Brazil). We describe three new species: Copelatus espinhasso sp. nov. (Brazil), Copelatus florae sp. nov. (Brazil, Colombia, Costa Rica, Ecuador and Panama) including its larva, and Copelatus panguana sp. nov. (Peru). Based on their male genital morphology, these species form a distinct clade within the genus. Other shared characters such as compact drop-shaped habitus, shortened appendages, and reduction of natatory setae on legs are interpreted as adaptations for what we call the ‘hygrofloric’ lifestyle. We document the thin water layer between leaf axils as the preferred habitat of both Copelatus adults and larvae, and not necessarily the central tank.

Bromeliads compete with microorganisms for nutrients in their phytotelm

Tank bromeliads impound water between their leaves, and have been successfully used as a model system to develop ecological theory. One fundamental interaction remains, however, unexplored in this system: the potential for bromeliad plants to compete with aquatic microorganisms for nutrients in their water-filled tanks. Here we conduct the first experimental test of this interaction by directly blocking bromeliads from accessing phytotelm water and quantifying change in the concentrations of dissolved inorganic nitrogen, phosphate, autotrophic algae and heterotrophic bacteria. To understand any context-dependency of responses, we crossed this manipulation with different combinations of light exposure and allochthonous resource type. We found that bromeliads reduced the concentrations of dissolved inorganic nitrogen, especially when the resource was nutrient-rich. Bromeliads also reduced the concentration of heterotrophic bacteria within their tank regardless of environmental context and algal concentrations particularly under high light, nutrient-rich resource conditions. Our results demonstrate that bromeliads can strongly impact their phytotelm by reducing the concentration of key nutrients, concomitantly reducing densities of algae under optimal growth conditions. This strong competition with algae for nutrients may explain why high abundances of algae within wild tank bromeliads are only found under specific field conditions. Bacteria may have a more commensal relationship with bromeliads, as they tend to increase bromeliad nutrition through decomposition. In conclusion, our study underlines that tank bromeliads do not only provide habitats for other species, but also interact strongly with the aquatic ecosystem that they create.

Temperature-dependent resistance to starvation of three contrasting freshwater ciliates

We investigated the temperature-dependent response to starvation of three contrasting freshwater ciliates (Ciliophora). The cyst-forming algivorous species Meseres corlissi and the bactivorous species Glaucomides bromelicola, which cannot form cysts, co-occur in the reservoirs (tanks) of tree bromeliads. The mixotrophic species Coleps spetai is common in many lakes. We hypothesized that the ciliates’ different traits and life strategies would affect their survival rates and temperature sensitivity under food depleted conditions. We measured the decline of the ciliate populations in microcosm experiments at different temperatures for several days. We used an imaging flow cytometer to size the ciliates and documented their morphological and physiological changes in response to starvation. We found that the cyst-forming species had the highest mortality rates but may endure long-term starvation by encystment. The sympatric, non-encysting species suffered the lowest mortality rates and could survive for more than three weeks without food. The mixotrophic species had intermediate mortality rates but showed the highest phenotypic plasticity in response to starvation. A significant fraction of the C. spetai population appeared unaffected by starvation, suggesting that the endosymbionts provided some resources to the host cells. The mean mortality rate per day of all three species increased with temperature by 0.09 °C−1.

Effects of warming on the structure of aquatic communities in tropical bromeliad microecosystems.

Freshwaters are among the most vulnerable ecosystems to climate warming, with projected temperature increases over the coming decades leading to significant losses of aquatic biodiversity. Experimental studies that directly warm entire natural ecosystems in the tropics are needed, for understanding the disturbances on aquatic communities. Therefore, we conducted an experiment to test the impacts of predicted future warming on density, alpha diversity, and beta diversity of freshwater aquatic communities, inhabiting natural microecosystems-Neotropical tank bromeliads. Aquatic communities within the tanks bromeliads were experimentally exposed to warming, with temperatures ranging from 23.58 to 31.72°C. Linear regression analysis was used to test the impacts of warming. Next, distance-based redundancy analysis was performed to assess how warming might alter total beta diversity and its components. This experiment was conducted across a gradient of habitat size (bromeliad water volume) and availability of detrital basal resources. A combination of the highest detritus biomass and higher experimental temperatures resulted in the greatest density of flagellates. However, the density of flagellates declined in bromeliads with higher water volume and lower detritus biomass. Moreover, the combination of the highest water volume and high temperature reduced density of copepods. Finally, warming changed microfauna species composition, mostly through species substitution (β repl component of total beta-diversity). These findings indicate that warming strongly structures freshwater communities by reducing or increasing densities of different aquatic communities groups. It also enhances beta-diversity, and many of these effects are modulated by habitat size or detrital resources.

Acid waters in tank bromeliads: Causes and potential consequences.

The consequences of acidity for plant performance are profound, yet the prevalence and causes of low pH in bromeliad tank water are unknown despite its functional relevance to key members of many neotropical plant communities. We investigated tank water pH for eight bromeliad species in the field and for the widely occurring Guzmania monostachia in varying light. We compared pH changes over time between plant and artificial tanks containing a solution combined from several plants. Aquaporin transcripts were measured for field plants at two levels of pH. We investigated relationships between pH, leaf hydraulic conductance, and CO2 concentration in greenhouse plants and tested proton pump activity using a stimulator and inhibitor. Mean tank water pH for the eight species was 4.7 ± 0.06 and was lower for G. monostachia in higher light. The pH of the solution in artificial tanks, unlike in plants, did not decrease over time. Aquaporin transcription was higher for plants with lower pH, but leaf hydraulic conductance did not differ, suggesting that the pH did not influence water uptake. Tank pH and CO2 concentration were inversely related. Fusicoccin enhanced a decrease in tank pH, whereas orthovanadate did not. Guzmania monostachia acidified its tank water via leaf proton pumps, which appeared responsive to light. Low pH increased aquaporin transcripts but did not influence leaf hydraulic conductance, hence may be more relevant to nutrient uptake.

Effect of substrate fertility on tank-bromeliad performances

Members of the plant family Bromeliaceae can uptake nutrients directly from their leaves via leaf absorbing trichomes and their roots have long been reduced to anchorage function, thus overlooked. Recently, evidence has accumulated for a significant role for the roots of some species of tank bromeliads in both water and nutrients absorption. However, to date, little attention has been paid to the importance of the substrate fertility for the structure of the roots and the growth and performances of tank bromeliads. This study investigated the effect of substrate fertility on Aechmea aquilega regarding leaf and root traits, nutrient content, and growth. Seeds of this tank bromeliad were sowed in a greenhouse in French Guiana in three different substrates: a nutrient-poor, a nutrient-rich and a mixed substrate. The performances of 15-month-old A. aquilega were assessed by measuring leaf and root traits related to nutrient acquisition and resources capture. We show that plants growing in nutrient-poor substrate grew twice slower and were smaller than plants grown on the nutrient-rich substrate with fewer leaves and roots, lower total dry mass, and smaller leaves and root length. Overall, 70% of measured traits responded significantly to the experimental treatments indicating that the response of A. aquilega to nutrient availability is a combination of physiological processes, leaf and root structure, and chemistry. This study is the first to show that the fertility of the substrate on which the bromeliad A. aquilega grows has a strong and lasting effect on the plant performances and may be a relevant factor for bromeliad ecology.

New Proposal of Epiphytic Bromeliaceae Functional Groups to Include Nebulophytes and Shallow Tanks.

The Bromeliaceae family has been used as a model to study adaptive radiation due to its terrestrial, epilithic, and epiphytic habits with wide morpho-physiological variation. Functional groups described by Pittendrigh in 1948 have been an integral part of ecophysiological studies. In the current study, we revisited the functional groups of epiphytic bromeliads using a 204 species trait database sampled throughout the Americas. Our objective was to define epiphytic functional groups within bromeliads based on unsupervised classification, including species from the dry to the wet end of the Neotropics. We performed a hierarchical cluster analysis with 16 functional traits and a discriminant analysis, to test for the separation between these groups. Herbarium records were used to map species distributions and to analyze the climate and ecosystems inhabited. The clustering supported five groups, C3 tank and CAM tank bromeliads with deep tanks, while the atmospheric group (according to Pittendrigh) was divided into nebulophytes, bromeliads with shallow tanks, and bromeliads with pseudobulbs. The two former groups showed distinct traits related to resource (water) acquisition, such as fog (nebulophytes) and dew (shallow tanks). We discuss how the functional traits relate to the ecosystems inhabited and the relevance of acknowledging the new functional groups.

Diversity and putative metabolic function of prokaryotic communities in tank bromeliads along an elevation gradient in tropical Mexico.

Tank bromeliads are unique canopy microhabitats that offer freshwater and organic nutrient-rich substrates in the Neotropics. In them it is possible to thoroughly characterize environmental factors and species composition of terrestrial and aquatic biota. Therefore, these plants have been used as natural models to study how communities are distributed and assembled. Here we used amplicon sequencing of the 16S rRNA gene and their functional annotations to study the diversity and metabolic potential of prokaryotic communities in tank bromeliads in five different forests along an elevation gradient in tropical Mexico. Furthermore, we analyzed the effects of vegetation type and environmental factors inside the tanks on prokaryotic composition. We found a high prokaryotic diversity in tank bromeliads along the elevation gradient. Prokaryotes commonly observed in acidic environments rich in organic carbon, and the potential pathogen Pasteurella multocida, were present in all samples, but few amplicon sequence variants were shared between forests. The prokaryotic composition was affected by forest type, and comparisons against null models suggest that it was shaped by non-neutral processes. Furthermore, prokaryotic community changes significantly covaried with tank water temperature, pH, and inorganic carbon. We found a high diversity of putative metabolic groups dominated by chemoheterotrophs and fermenters, but taxonomic groups involved in nitrogen and sulfur cycling were also present in all samples. These results suggest that tank bromeliads promote taxonomic and metabolic diversity of the prokaryotic community at a local and regional scale and play an important role in the biogeochemistry of forest canopies in the Neotropics.