Deforestation is the turning point for the spreading of a weedy epiphyte: an IBM approach.

The rapid spread of many weeds into intensely disturbed landscapes is boosted by clonal growth and self-fertilization strategies, which conversely increases the genetic structure of populations. Here, we use empirical and modeling approaches to evaluate the spreading dynamics of Tillandsia recurvata (L.) L. populations, a common epiphytic weed with self-reproduction and clonal growth widespread in dry forests and deforested landscapes in the American continent. We introduce the TRec model, an individual-based approach to simulate the spreading of T. recurvata over time and across landscapes subjected to abrupt changes in tree density with the parameters adjusted according to the empirical genetic data based on microsatellites genotypes. Simulations with this model showed that the strong spatial genetic structure observed from empirical data in T. recurvata can be explained by a rapid increase in abundance and gene flow followed by stabilization after ca. 25 years. TRec model's results also indicate that deforestation is a turning point for the rapid increase in both individual abundance and gene flow among T. recurvata subpopulations occurring in formerly dense forests. Active reforestation can, in turn, reverse such a scenario, although with a milder intensity. The genetic-based study suggests that anthropogenic changes in landscapes may strongly affect the population dynamics of species with 'weedy' traits.

Air quality biomonitoring of trace elements in the metropolitan area of Huancayo, Peru using transplanted Tillandsia capillaris as a biomonitor.

The air quality and distribution of trace elements in a metropolitan area of the Peruvian Andes were evaluated using transplanted epiphytic Tillandsia capillaris as biomonitors. Biomonitors were collected from the non-contaminated area and exposed to five sites with different types of contamination for three months in 2017. After exposure, the content of twenty-one elements were determined by ICP-MS analysis. Datasets were evaluated by one-way ANOVA, exposed-to-baseline (EB), hierarchical cluster analysis (HCA) and principal component analysis (PCA). Results showed significant differences among sampling sites for several elements. According to EF ratios for Ba, Cr, Cu, Pb, Sb, and Zn EB ratios value greater than 1.75 were found around urban areas, indicating anthropogenic influence, which can be attributed to vehicular sources. The highest values of As and Cd were found in areas of agricultural practices, therefore their presence could be related to the employment of agrochemicals (pesticides, herbicides, and phosphate fertilizers). HCA shows that most elements come from vehicular sources and lower from agricultural and natural sources.

Design of a unidirectional water valve in Tillandsia.

The bromeliad Tillandsia landbeckii thrives in the Atacama desert of Chile using the fog captured by specialized leaf trichomes to satisfy its water needs. However, it is still unclear how the trichome of T. landbeckii and other Tillandsia species is able to absorb fine water droplets during intermittent fog events while also preventing evaporation when the plant is exposed to the desert's hyperarid conditions. Here, we explain how a 5800-fold asymmetry in water conductance arises from a clever juxtaposition of a thick hygroscopic wall and a semipermeable membrane. While absorption is achieved by osmosis of liquid water, evaporation under dry external conditions shifts the liquid-gas interface forcing water to diffuse through the thick trichome wall in the vapor phase. We confirm this mechanism by fabricating artificial composite membranes mimicking the trichome structure. The reliance on intrinsic material properties instead of moving parts makes the trichome a promising basis for the development of microfluidics valves.

Unravelling intricate interactions among atmospheric bromeliads with highly overlapping niches in seasonal systems.

Biotic and abiotic interactions are important factors that explain community assembly. For example, epiphytic communities are shaped by tree traits that can act as environmental filters, but also by positive and/or negative interactions among coexisting epiphytes on a tree. Here, we studied interactions among three widespread atmospheric bromeliads with overlapping niches (Tillandsia recurvata, T. tricholepis and T. pohliana), using experimental data about facilitation through seed capture, interspecific interaction during seed germination and competition among adult individuals. We aim to understand how species interactions are reflected in the natural coexisting patterns of epiphytes in communities at high and low tree densities. Tillandsia pohliana showed higher facilitation by capturing almost all wind-dispersing seeds, and had the largest reduction in seed germination in the presence of any adult individual, also presenting a relatively high rate of adult mortality in the presence of other individuals. Our results indicate higher colonisation ability for T. pohliana and distinct strategies of rapid exploitation of T. recurvata and T. tricholepis individuals. In natural communities, the coexistence among atmospheric bromeliads may be hampered by dispersal limitations in wind-dispersed epiphytes at high tree density conditions, but a negative effect of T. recurvata on T. pohliana is still reflected in their reduced coexistence. However, competitive patterns observed in the experimental data may be overshadowed by a possible mass effect driving large communities under such conditions. Our results show the joint effect of positive interactions and high dispersal levels on the community patterns of atmospheric bromeliads.

High resilience to extreme climatic changes in the CAM epiphyte Tillandsia utriculata L. (Bromeliaceae).

Climate change is expected to increase the frequency of extreme climatic events, yet few studies have addressed the capacity of plant species to deal with such events. Species that are widespread are predicted to be highly plastic and able to acclimate to highly changing conditions. To study the plasticity in physiological responses of the widely distributed epiphyte Tillandsia utriculata, we transplanted individuals from a coastal scrub and broadleaf evergreen forest to a similar coastal scrub site and forest. After a 45-day acclimation, the plants were moved to a semi-controlled greenhouse at each site, and then subjected to a 20-day drought. Physiological variables were measured during the acclimation and the drought. The individuals of scrub and forest populations had similar relative water content and carbon assimilation in the contrasting conditions of the two transplantation sites despite the high discrepancy between the environments at their original site. Electron transport rates were higher in individuals from the scrub population. Electron transport rates were also higher than estimated from carbon assimilation, suggesting that photorespiration was present. The individuals of the coastal scrub population had a higher capacity to dissipate excess energy this way. The relative distance index of plasticity was high overall, indicating that some traits are highly plastic (titratable acidity, carbon assimilation) in order to maintain the stability of others (maximum quantum yield Fv /Fm and relative water content). We conclude that T. utriculata is a highly plastic species with a high capacity to tolerate extreme environmental changes over a short time.

Dispersal limitation of Tillandsia species correlates with rain and host structure in a central Mexican tropical dry forest.

Seed dispersal permits the colonization of favorable habitats and generation of new populations, facilitating escape from habitats that are in decline. There is little experimental evidence of the factors that limit epiphyte dispersion towards their hosts. In a tropical dry forest in central Mexico, we monitored the phenology of dispersion of epiphyte species of the genus Tillandsia; we tested experimentally whether precipitation could cause failures in seed dispersal and whether seed capture differs among vertical strata and between host species with high (Bursera copallifera) and low (Conzattia multiflora) epiphyte loads. With the exception of one species that presents late dispersion and low abundance, all of the species disperse prior to the onset of the rainy season. However, early rains immobilize the seeds, affecting up to 24% of the fruits in species with late dispersion. We observed that Tillandsia seeds reach both Bursera and Conzattia hosts, but found that adherence to the host is 4-5 times higher in Bursera. Furthermore, seeds liberated from Bursera travel shorter distances and up to half may remain within the same crown, while the highest seed capture takes place in the upper strata of the trees. We conclude that dispersion of Tillandsia seeds is limited by early rains and by the capture of seeds within the trees where populations concentrate. This pattern of capture also helps to explain the high concentrations of epiphytes in certain hosts, while trees with few epiphytes can be simultaneously considered deficient receivers and efficient exporters of seeds.

Trade-off between soluble protein production and nutritional storage in Bromeliaceae.

BACKGROUND AND AIMS: Bromeliads are able to occupy some of the most nutrient-poor environments especially because they possess absorptive leaf trichomes, leaves organized in rosettes, distinct photosynthetic pathways [C3, Crassulacean acid metabolism (CAM) or facultative C3-CAM], and may present an epiphytic habit. The more derived features related to these traits are described for the Tillandsioideae subfamily. In this context, the aims of this study were to evaluate how terrestrial predators contribute to the nutrition and performance of bromeliad species, subfamilies and ecophysiological types, whether these species differ in their ecophysiological traits and whether the physiological outcomes are consistent among subfamilies and types (e.g. presence/absence of tank, soil/tank/atmosphere source of nutrients, trichomes/roots access to nutrients). METHODS: Isotopic (15N-enriched predator faeces) and physiological methods (analyses of plant protein, amino acids, growth, leaf mass per area and total N incorporated) in greenhouse experiments were used to investigate the ecophysiological contrasts between Tillandsioideae and Bromelioideae, and among ecophysiological types when a predatory anuran contributes to their nutrition. KEY RESULTS: It was observed that Bromelioideae had higher concentrations of soluble protein and only one species grew more (Ananas bracteatus), while Tillandsioideae showed higher concentrations of total amino acids, asparagine and did not grow. The ecophysiological types that showed similar protein contents also had similar growth. Additionally, an ordination analysis showed that the subfamilies and ecophysiological types were discrepant considering the results of the total nitrogen incorporated from predators, soluble protein and asparagine concentrations, relative growth rate and leaf mass per area. CONCLUSIONS: Bromeliad subfamilies showed a trade-off between two strategies: Tillandsioideae stored nitrogen into amino acids possibly for transamination reactions during nutritional stress and did not grow, whereas Bromelioideae used nitrogen for soluble protein production for immediate utilization, possibly for fast growth. These results highlight that Bromeliaceae evolution may be directly associated with the ability to stock nutrients.

From dusk till dawn: nocturnal and diurnal pollination in the epiphyte Tillandsia heterophylla (Bromeliaceae).

In order to compare the effectiveness of diurnal and nocturnal pollinators, we studied the reproductive biology and pollinators of Tillandsia heterophylla E. Morren, an epiphytic tank bromeliad endemic to southeastern Mexico. Since anthesis in T. heterophylla is predominantly nocturnal but lasts until the following day, we hypothesised that this bromeliad would receive visits from both diurnal and nocturnal visitors, but that nocturnal visitors would be the most effective pollinators, since they arrive first to the receptive flower, and that bats would be the most frequent nocturnal visitors, given the characteristics of the nectar. Flowering of T. heterophylla began in May and lasted until July. The species is fully self-compatible, with an anthesis that lasts for ca. 15-16 h. Mean volume of nectar produced per flower was 82.21 µl, with a mean sugar concentration of 6.33%. The highest volume and concentration of nectar were found at 20:00 h, with a subsequent decline in both to almost zero over the following 12-h period. T. heterophylla has a generalist pollination system, since at least four different morphospecies of visitors pollinate its flowers: bats, moths, hummingbirds and bees. Most of the pollinating visits corresponded to bats and took place in the early evening, when stigma receptivity had already begun; making bats the probable pollinator on most occasions. However, diurnal pollinators may be important as a 'fail-safe' system by which to guarantee the pollination of T. heterophylla.

High but not dry: diverse epiphytic bromeliad adaptations to exposure within a seasonally dry tropical forest community.

• Vascular epiphytes have developed distinct lifeforms to maximize water uptake and storage, particularly when delivered as pulses of precipitation, dewfall or fog. The seasonally dry forest of Chamela, Mexico, has a community of epiphytic bromeliads with Crassulacean acid metabolism showing diverse morphologies and stratification within the canopy. We hypothesize that niche differentiation may be related to the capacity to use fog and dew effectively to perform photosynthesis and to maintain water status. • Four Tillandsia species with either 'tank' or 'atmospheric' lifeforms were studied using seasonal field data and glasshouse experimentation, and compared on the basis of water use, leaf water δ(18) O, photosynthetic and morphological traits. • The atmospheric species, Tillandsia eistetteri, with narrow leaves and the lowest succulence, was restricted to the upper canopy, but displayed the widest range of physiological responses to pulses of precipitation and fog, and was a fog-catching 'nebulophyte'. The other atmospheric species, Tillandsia intermedia, was highly succulent, restricted to the lower canopy and with a narrower range of physiological responses. Both upper canopy tank species relied on tank water and stomatal closure to avoid desiccation. • Niche differentiation was related to capacity for water storage, dependence on fog or dewfall and physiological plasticity.

It takes two to tango: self incompatibility in the bromeliad Tillandsia streptophylla (Bromeliaceae) in Mexico.

Floral phenology and breeding system of Tillandsia streptophylla (Bromeliaceae) were studied in a low inundated forest in Yucatan, Mexico. During the flowering season, from March to August, terminal scapose 1-branched, paniculate inflorescences are produced with one flower per branch opening per day, over a period of 11-29 days. Flowers are tubular, light violet, with the stigma placed below the anthers, both protruding above the corolla. Flowers are protandrous, with anthers releasing pollen from 0500 hours and stigma becoming receptive around 0900 hours. Controlled experimental crosses suggest that Tillandsia streptophylla is self incompatible and therefore, pollinator-dependent.

It takes two to tango: self incompatibility in the bromeliad Tillandsia streptophylla (Bromeliaceae) in Mexico

Floral phenology and breeding system of Tillandsia streptophylla (Bromeliaceae) were studied in a low inundated forest in Yucatan, Mexico. During the flowering season, from March to August, terminal scapose 1-branched, paniculate inflorescences are produced with one flower per branch opening per day, over a period of 11-29 days. Flowers are tubular, light violet, with the stigma placed below the anthers, both protruding above the corolla. Flowers are protandrous, with anthers releasing pollen from 0500 hours and stigma becoming receptive around 0900 hours. Controlled experimental crosses suggest that Tillandsia streptophylla is self incompatible and therefore, pollinator-dependent. Rev. Biol. Trop. 57 (3): 761-770. Epub 2009 September 30.

Estudiamos la fenología floral y el sistema de cruzamiento de la bromelia Tillandsia streptophylla (Bromeliaceae) en una selva baja inundable en Yucatán, México. Durante la estación de floración (marzo a agosto), las plantas producen una inflorescencia terminal, escaposa, paniculada, 1-dividida, con una flor abriendo por rama por día para un período de floración de 11-29 días por inflorescencia. Las flores son tubulares, de corola violeta claro, con el estigma y anteras exertos, pero las anteras más largas que el estigma en antesis. Las flores son protandras, con las anteras liberando el polen desde las 0500 horas y la receptividad del estigma comenzando a las 0900 horas. Los cruces experimentales controlados sugieren que Tillansdia streptophylla es auto incompatible y por ende, dependiente de los polinizadores.

Comparison of the air pollution biomonitoring ability of three Tillandsia species and the lichen Ramalina celastri in Argentina.

Bioaccumulation ability and response to air pollution sources were evaluated for Tillandsia capillaris Ruíz and Pav. f. capillaris, T. recurvata L., T. tricholepis Baker and the lichen Ramalina celastri (Spreng.) Krog. and Swinsc. Epiphyte samples collected from a non contaminated area in the province of Córdoba were transplanted to a control site and three areas categorised according to agricultural, urban and industrial (metallurgical and metal-mechanical) emission sources. Bioindicators were exposed for 3-, 6- and 9-month periods. A foliar damage index was established for Tillandsia and a pollution index for the lichen, and S, Fe, Mn and Zn concentrations were determined. An order of efficiency for the species and conditions studied is proposed taking into account heavy metal accumulation: T. recurvata >T. tricholepis >R. celastri >T. capillaris. All species studied showed Mn to be related to agricultural activity and Fe to industries and soil particles, and Zn was related to urban and industrial sources. As far as physiological response is concerned, T. tricholepis and T. capillaris were more sensitive to agricultural activities, whereas T. recurvata was sensitive to urban and industrial sources, and only partially to agricultural sources. No relationship was found for R. celastri.

The narrow-leaf syndrome: a functional and evolutionary approach to the form of fog-harvesting rosette plants.

Plants that use fog as an important water-source frequently have a rosette growth habit. The performance of this morphology in relation to fog interception has not been studied. Some first-principles from physics predict that narrow leaves, together with other ancillary traits (large number and high flexibility of leaves, caudices, and/or epiphytism) which constitute the "narrow-leaf syndrome" should increase fog-interception efficiency. This was tested using aluminum models of rosettes that differed in leaf length, width and number and were exposed to artificial fog. The results were validated using seven species of Tillandsia and four species of xerophytic rosettes. The total amount of fog intercepted in rosette plants increased with total leaf area, while narrow leaves maximized interception efficiency (measured as interception per unit area). The number of leaves in the rosettes is physically constrained because wide-leafed plants can only have a few blades. At the limits of this constraint, net fog interception was independent of leaf form, but interception efficiency was maximized by large numbers of narrow leaves. Atmospheric Tillandsia species show the narrow-leaf syndrome. Their fog interception efficiencies were correlated to the ones predicted from aluminum-model data. In the larger xerophytic rosette species, the interception efficiency was greatest in plants showing the narrow-leaf syndrome. The adaptation to fog-harvesting in several narrow-leaved rosettes was tested for evolutionary convergence in 30 xerophytic rosette species using a comparative method. There was a significant evolutionary tendency towards the development of the narrow-leaf syndrome the closer the species grew to areas where fog is frequently available. This study establishes convergence in a very wide group of plants encompassing genera as contrasting as Tillandsia and Agave as a result of their dependence on fog.

Effect of canopy position on germination and seedling survival of epiphytic bromeliads in a Mexican humid montane forest.

BACKGROUND AND AIMS: Seeds of epiphytes must land on branches with suitable substrates and microclimates to germinate and for the resulting seedlings to survive. It is important to understand the fate of seeds and seedlings in order to model populations, but this is often neglected when only established plants are included in analyses. METHODS: The seeds of five bromeliad species were exposed to different canopy positions in a Mexican montane forest, and germination and early seedling survival were recorded. Additionally, the survival of naturally dispersed seedlings was monitored in a census over 2.5 years. Survival analysis, a procedure rarely used in plant ecology, was used to study the influence of branch characteristics and light on germination and seedling survival in natural and experimental populations. KEY RESULTS: Experimental germination percentages ranged from 7.2 % in Tillandsia deppeana to 33.7 % in T. juncea, but the seeds of T. multicaulis largely failed to germinate. Twenty months after exposure between 3.5 and 9.4 % of the seedlings were still alive. There was no evidence that canopy position affected the probability of germination, but time to germination was shorter in less exposed canopy positions indicating that higher humidity accelerates germination. More experimental seedlings survived when canopy openness was high, whereas survival in census-seedlings was influenced by moss cover. While mortality decreased steadily with age in juveniles of the atmospheric Tillandsia, in the more mesomorphic Catopsis sessiliflora mortality increased dramatically in the dry season. CONCLUSIONS: Seedling mortality, rather than the failure to germinate, accounts for the differential distribution of epiphytes within the canopy studied. With few safe sites to germinate and high seedling mortality, changes of local climate may affect epiphyte populations primarily through their seedling stage.

Generous-like flowers: nectar production in two epiphytic bromeliads and a meta-analysis of removal effects.

Animal-pollinated angiosperm plants that respond positively to nectar removal by replenishment invest energy that can entail a reproductive cost. We investigated whether or not nectar removal stimulates replenishment in two hummingbird-pollinated bromeliad species. Nectar replenishment rates were also assessed by removing nectar from manually pollinated flowers because pollination events might be used as signals to save energy by preventing allocation to post-pollination nectar production. Then we synthesized the current understanding of nectar removal effects by reviewing existing published studies with a meta-analysis. The magnitude and significance of estimated nectar removal effects and factors associated with variation in size and direction of nectar removal effects were elucidated with the meta-analysis. We found that both Tillandsia species strongly respond to repeated nectar removal by producing >3 times additional nectar. Nectar secretion patterns were not altered by pollination (stigmatic pollen deposition) and we found no evidence of nectar reabsorption. Although the effect size varied widely across systems and/or environmental conditions, the meta-analysis showed that nectar removal had overall a positive effect on nectar replenishment (mainly among species inhabiting wet tropical habitats such as Tillandsia), and a negative effect on the secretion of additional sugar, suggesting that those plants are resource limited and conservative in the secretion of additional sugar.