The studies about the negative effect of epiphytes on their phorophytes show contradictory results and are based on limited variables (e.g., shoot survival). On branches of Bursera copallifera, we experimented with the transplantation/removal of Tillandsia recurvata and artificial tussocks, measuring shoot survival, growth, generation of new shoots, and production of inflorescences and fruits. Most single traits did not differ between treatments. The generation of new shoots was lower in the treatments where T. recurvata was present and increased in the branches where it was removed. The lowest shoot survival was in the treatments where T. recurvata was present or was removed. Removing T. recurvata increased plant relative fitness, and it was 43% lower in branches with T. recurvata. Tillandsia recurvata is a structural parasite of B. copallifera. A negative effect of epiphytes on their phorophytes appears counterintuitive since it would not be evolutionarily stable for an epiphyte to shorten the lifespan of its support. Tillandsia recurvata populations are concentrated on B. copallifera branches between 2-4 cm in diameter, while smaller branches are mostly empty, so it is possible that the negative effect of T. recurvata occurs in the smallest branches, explaining why T. recurvata populations are biased to larger branches.

Many bromeliads occur in habitats that are potentially affected by salt. Pitcairnia halophila is a particularly interesting case in this context due to its growing sites in the spray zone of coastal cliffs in Central America. We conducted an experimental/observational study that explored the physiological and ecological basis of the occurrence of this species. Germination of P. halophila was as affected by increasing salinity as that of congeneric species from other natural growing sites, but seedling growth was less impaired in comparison and remained positive even at exposure to half-strength seawater. Interestingly, in situ plant nutrient concentrations documented over an entire growing season suggest that this capacity to tolerate salt stress is not tested at the natural growing site – Na+ concentrations were consistently low in leaves and shoots and comparable to those of seedlings watered with distilled water in the laboratory. We conclude that P. halophila shows the physiological ability to deal with medium levels of salinity, but ecological conditions at the humid tropical growing site allow these plants to avoid salt stress entirely in spite of the close proximity to the sea.

The low availability of pollinators in high-elevation ecosystems can lead to flowering plants showing different adaptive responses in order to assure their reproductive success. Shifts toward autogamy and asexual reproductive rates (the reproductive assurance hypothesis) and the compensatory measures to maintain outcrossing such as flower longevity and more prolonged pistil receptivity (the increased pollination probability hypothesis) are some of these responses. Several studies have tested both hypotheses, but investigations of plants of tropical alpine environments such as paramos that support these assumptions are still scarce. Puya nitida, an endemic Colombian plant species distributed in the paramo and subparamo in the Eastern Cordillera of Cundinamarca department, was used as a case study to test its reproductive characteristics that assure its sexual reproduction. We analysed the species’ floral morphology and development, its phenological patterns and its plant mating-system. We found that Puya nitida showed floral characteristics that promote pollination by birds, herkogamy and dichogamy, flowers and receptive stigmas with 9 and 12 days of longevity, respectively and an index of self-incompatibility that shows that it is mostly self-incompatible. We found a synchronic phenological pattern with an annual frequency and an intermediate duration with a peak in the period of lowest rainfall. Our results suggested that longer floral development, prolonged stigma receptivity, herkogamy and dichogamy and self-incompatibility might assure reproductive success, since the cross-pollination might be favoured when few pollinators are in attendance. Overall, these reproductive mechanisms add evidence to the increased pollination probability hypothesis, specifically for a plant species of a tropical high-elevation ecosystem where pollinators are scarce.

There are a number of studies describing the gross range of morpho-anatomical variability in epiphytic Tillandsia species, but the interspecific variation in seed traits remain largely unexplored, although these play an important role in determining dispersal and establishment success. In order to evaluate interspecific variation in seed morphology, anatomy and germination, we sampled six Tillandsia species from the Yucatan peninsula, Mexico, distributed along a precipitation gradient. We studied morpho-anatomical traits (seed length, seed mass, ratio of coma to seed, ratio of embryo to endosperm), seed terminal velocity in still air, and performed histochemical analyses and germination trials under controlled conditions. Tillandsia recurvata differs from the other five species in the structure of the plumose coma; it was the only species lacking an endosperm and showed distinct seedling development. Among the species, bigger seeds were related to longer comas, and had higher germinability. Overall, seed terminal velocity was invariably slow, compared with reports of other anemochorous species, suggesting a high dispersal potential. Taxonomical and ecological implications of our results are discussed.

The understanding that many aspects of the spatial and temporal patterns of epiphyte communities may be explained by the comportment of early life stages has given rise to a considerable number of germination studies in recent years. Unfortunately, protocols frequently use unproven assumptions and arbitrary experimental conditions. To make future studies as ecologically meaningful as possible we address a number of potential pitfalls with a series of experiments with seeds from a total of 16 species. We show that it is safe to collect capsules for experiments before natural dehiscence – there is afterripening even in the case of very early collections. The application of fluctuating temperatures is not imperative, because there is no consistent difference in the germination response under constant versus fluctuating temperatures. The effects of different osmotic potentials and intermittent drought of varying intensity on germination are qualitatively, but not quantitatively, comparable. Due to the greater ecological realism, we encourage the use of the latter. However, care must be taken to use realistic temperatures – the impact of intermittent drought on germination is modulated by temperature. This highlights the need for data on the in situ temperature regimes during germination as an important prerequisite towards more realistic experiments in the field of germination ecology of vascular epiphytes.

Vertical stratification of vascular epiphytes is frequently attributed to niche partitioning along microclimatic gradients but experimental confirmations of this notion are rare. This study investigates the role of the regeneration phase for the stratification of five bromeliad (Catopsis sessiliflora, Guzmania subcorymbosa, Tillandsia anceps, T. bulbosa and Werauhia gladioliflora) and seven aroid species (Anthurium acutangulum, A. brownii, A. clavigerum, A. durandii, A. friedrichsthalii, A. hacumense and A. scandens) in a Panamanian rain forest. We documented gradients of temperature, vapour pressure deficit and light (n = 10 d) as well as species height distributions (n = 11–120). Microclimatic gradients were substantial (maximal T and RH differences between strata: 5 °C and 18%, respectively) and mean attachment heights of the study species (range = 4–21 m) differed significantly. We tested sensitivity to recurrent drought (four treatments) during germination (all species, cumulative germination of 20 seeds, n = 3) and seedling growth (four aroid species, n = 25). Seedling survival of six aroid species transplanted to three heights (n = 27) was monitored in situ. Some species did not germinate under severe recurrent drought while others germinated at the same rate in all treatments. Seedlings of the most exposed species grew fastest under intermediate recurrent drought while those of the other three species grew fastest when kept constantly wet. Survival of transplanted seedlings did not depend on species attachment height, but this may be attributable to insufficient statistical power. Taken together, the results suggest that the stratification can be explained to a large degree by differential sensitivity to the vertical moisture gradient during the regeneration phase.

Low germination and seedling survival probabilities are reported in various species of epiphytic bromeliad (Benzing 1978, Hietz et al. 2011, Toledo-Aceves & Wolf 2008, Winkler et al. 2005; but see Cascante-Marín et al. 2008). If germination and seedling survival are limiting factors in the life cycle, differential germination and seedling survival between species should be reflected in the relative abundance of established plants (Cascante-Marín et al. 2006, 2008) and in their presence or absence in secondary vegetation (Hietz et al. 2011), while differential germination within the tree would be expected to contribute to a heterogeneous distribution of established plants within the canopy (Hietz et al. 2011, Zotz & Vollrath 2002). Many factors influence the performance and distribution of epiphytes, including forest condition, disturbance type, distance from seed source, tree size and species, microclimate, epiphyte population dynamics and physiology (Cascante-Marín et al. 2009, Hietz et al. 2011, Valencia-Diaz et al. 2010, Zotz & Hietz 2001). In this study, we tested whether germination and seedling survival rates differ between the epiphytic bromeliads Tillandsia multicaulis Steud., T. punctulata Schldl. & Cham. and T. butzii Mez, and whether species abundance reflects the ability to germinate and survive as seedlings within the cloud-forest canopy. We also explore how morphological and physiological traits of the studied species can influence their early establishment.

Biotechnological techniques comprise useful tools for the conservation of endangered plant genetic resources. In the present work, polymorphism and usefulness of amplified fragment length polymorphism (AFLP) markers in assessing the genetic diversity in populations of Billbergia zebrina were investigated in nodular cultures and adult plants of the species. AFLP markers revealed moderate-to-high genetic diversity based on the estimations of Nei's gene diversity (mean He = 0.28), Shannon index of diversity (mean HS = 0.48) and the number of polymorphic fragments (mean of 56.17 polymorphic fragments over six primer pairs). In comparison to published studies of population genetics performed in other bromeliad species, the present study suggests that natural populations of B. zebrina likely maintain high levels of genetic diversity, an important feature towards conservation of plant genetic resources. The results obtained reveal that AFLP markers comprise a powerful tool in order to assess the levels of genetic diversity in natural populations of this endangered species. Integrating AFLP markers with in vitro propagation techniques is understood as an adequate strategy for conservation programmes of this species.

Tree species are potential hosts for epiphytes; however in some forests epiphytes have a biased distribution among hosts. In a tropical dry forest of Mexico, previous research showed that there are trees with few epiphytes. It is possible that the bark of these hosts contain allelochemicals that influence epiphyte seed germination. The aims of this study were (1) to determine whether hosts with low epiphyte abundance (Ipomoea murucoides, I. pauciflora and Lysiloma acapulcense) would inhibit seed germination of Tillandsia recurvata through aqueous and organic bark extracts, (2) to determine whether germination of T. recurvata would differ among the hosts with low epiphyte abundance and a host with high epiphyte abundance (Bursera copallifera) and (3) to relate the chemical composition of organic bark extracts with inhibition of T. recurvata seed germination. Hexanic and dichloromethanic extracts were partially chemically characterized. Total phenolics and flavonoids concentrations of methanolic extracts were analysed. Aqueous and organic bark extracts from hosts with few epiphytes inhibited T. recurvata seed germination. Aqueous and dichloromethanic extracts of B. copallifera inhibited slightly the germination of T. recurvata. There was a positive correlation between concentration of flavonoids and inhibition of seed germination. Results suggest that a combination of compounds may be responsible for affecting the germination of T. recurvata. This study demonstrates the chemical effect of aqueous and organic bark extracts from hosts on germination of an epiphytic bromeliad.

It has been suggested that vascular epiphyte composition and abundance may be influenced by forest host composition. We studied the epiphyte species distribution among host species of a tropical dry forest in Mexico. All the epiphyte stands supported by woody plants (dbh > 3 cm) of ten forest plots (0.1 ha each) were counted. We measured the dbh of all the hosts, noted their bark characteristics (texture and peeling behaviour), and measured the bark thickness of the most abundant host species. Epiphyte distribution was biased toward a high concentration of epiphytes in three host species and a lower abundance of epiphytes on five host species. This was consistent among epiphyte taxa and host species, allowing us to classify hosts as preferred (with more epiphyte stands than expected by chance) and limiting species (with fewer epiphyte stands than expected by chance), at a community level. Host quality did not relate to mean phorophyte size (measured as basal area) or to bark characteristics (peeling behaviour, thickness and texture) between species. For some epiphyte taxa, the observed distribution indicated that the forest contained preferred and limiting host species mainly. Our data suggest that the epiphyte species in the forest studied could be host limited. We concluded that neither host size nor obvious bark characteristics can be used to predict epiphyte distribution and that further research is necessary.

Nectarivorous flower mites can reduce the volume of nectar available to pollinators. The effects of the flower mite Proctolaelaps sp. on nectar availability in flowers of a melittophilous bromeliad Neoregelia johannis (Bromeliaceae) was evaluated in a coastal rain forest in south-eastern Brazil. In a randomized block experiment utilizing 18 flower pairs, one per bromeliad ramet, pollinators (Bombus morio) and mites were excluded, and then nectar volume, sugar concentration and sugar mass were quantified over the anthesis period. Mites significantly reduced nectar volume early in the morning (6h00–8h00), but not later (10h00–12h00). Mites decreased total volume of nectar available up to 22%. Sugar concentration in nectar was higher earlier in the morning, and decreased between 10h00–12h00. The pronounced consumption of nectar by mites during the period of higher sugar concentration reduced the total amount of sugar available to pollinators by 31%. This is the first study showing that flower mites decrease nectar rewards in a melittophilous plant. Because nectar volume by itself incompletely describes nectar production rates and the effects of nectar removal by flower mites on the availability of sugar, our study highlights the inclusion of sugar content in future studies assessing the effects of thieves on nectar production rates.

Almost 50% of the estimated 2500 species in the Bromeliaceae grow epiphytically in a remarkably wide range of habitats from inhospitable deserts to tropical rain forests (Benzing 2000). The degree of dependence on the host varies (Benzing 1990, Laube & Zotz 2006), and in some cases, epiphytic bromeliads may dispense completely with living hosts, and may thrive on artificial substrates such as electrical cables. This is not entirely surprising because this family provides examples of particularly remarkable adaptations to the epiphytic habitat like water-impounding leaf bases and water- and nutrient-absorbing scales (Benzing 2000). The so-called atmospheric forms in this family (e.g. Tillandsia recurvata) possess a dense covering of these scales, and use roots entirely as holdfasts, while leaves perform photosynthesis and take up water and nutrients. Although quite regularly mentioned in the literature (Benzing 1990, 2000; Brighigna et al. 1997, Lüttge 1989), we are not aware of any study investigating the vigour of plants growing on cables compared with plants growing on trees. The only functional aspects that have already been studied are related to nutrients, i.e. presence of nitrogen-fixing endophytes in Tillandsia recurvata plants from natural hosts and from electrical cables in Mexico (Puente & Bashan 1994) and differences in chemical composition of plant tissues and nutrient retention by leaves in Tillandsia capillaris growing on cables and trees in Argentina (Abril & Bucher 2009). Intermittent water supply, but also low nutrient supply, play an important role in epiphyte ecology (Zotz & Hietz 2001), which leads to the following alternative hypotheses: (1) Plants on electrical cables are usually close to roads.

The habit, shade-tolerance and dispersal agent of 19 bromeliad species were studied in a rainforest community in order to relate specific traits to the spatial occurrence of the species. Highest density was found in riparian forest (RF), followed by restinga scrub (RE), rocky shore (RS) and dense canopy forest (DF). Terrestrial bromeliads occurred in open sites (RE and RS), epiphytes were absent in RS and facultative species occupied all four habitats. Bird-dispersed bromeliads occurred in the upper canopy, whereas those dispersed by mammals were low in the forest profile. Bromeliads which are dispersed by similar fauna fruit sequentially through the year. Differences in traits may allow the sharing of habitats and strata among species.

Plant establishment was studied in a swamp forest in the lowland Atlantic rainforest in southeastern Brazil. A topographic gradient allowed a clear distinction between a periodically flooded and a permanently flooded site in the study area. Both sites were mosaics of patches: the former showed soil patches of varying sizes subject to distinct flooding durations (from 0 to 300 d per year) and in the latter, the tank bromeliads which densely occupied the understorey, trap litter and provided ‘suspended soil’ patches. In the periodically flooded site, the occurrence of regenerants (< 1.0 m tall) of the actively regenerating tree populations of Calophyllum brasiliense, Symphonia globulifera and Tovomitopsis paniculata (all Clusiaceae), originating from seed or vegetative organs, was recorded for 1 m2 quadrats placed in flooded (n = 82) and unflooded (n = 103) patches. In the permanently flooded site, dominated by Tabebuia cassinoides (Bignoniaceae), the presence of germinated seedlings was counted for 400 rosettes of the terrestrial tank bromeliad Nidularium procerum. The results showed that unflooded patches, including tank bromeliads, favoured seedling establishment. However, T. paniculata and T. cassinoides colonize patches where flooding lasts longer or is permanent mostly by reproducing vegetatively, while the seeds of C. brasiliense, which showed no form of vegetative reproduction, and S. globulifera are successful in colonizing flooded patches. Germination tests and field observations indicated that, for C. brasiliense, pre- and post-dispersal seed dormancy, a two-phase seed dispersal (water and bats) and seedling tolerance to flooding accounted for this pattern.

Aechmea magdalenae is a terrestrial bromeliad that dominates areas of forest understorey on Barro Colorado Island, Panama. Nocturnal CO2 uptake via crassulacean acid metabolism was the primary means of carbon gain under well-watered conditions and all light regimes. The ability to maintain a positive carbon balance under conditions of very low light was demonstrated by laboratory measurements of photosynthesis and forest measurements of growth. Low-light-grown juvenile rosettes had the same daily net assimilation whether tested at photon flux densities of 15 or 300 μmol m−2 s−1 Growth rates of rosettes in treefall gaps were similar to those of plants in closed canopy forest. Growth rates of forest plants were increasingly correlated with canopy openness as the wet season progressed due to increased growth by gap plants, suggesting that water availability rather than light may limit growth during the annual dry season.

In comparing the population structure of epiphytic and saxicolous Tillandsia species I found no significant differences between polycarpic species. Epiphytic T. ionochroma populations had a higher seedling:adult ratio than saxicolous T. ionochroma populations, but the difference was not statistically significant. Tillandsia ultriculala, a monocarpic epiphyte, had a significantly higher seedling:adult ratio than the polycarpic species. T. utriculata's higher ratio may result from its greater seed production and lack of vegetative propagation. Unlike the polycarpic species, all adult-size rosettes grow from seeds in T. utriculata. Tillandsia sphaerocephala, an obligate saxicole, occurred in larger clusters (i.e. more rosettes per genet) than T. ionochroma. The increased tendency for vegetative reproduction in T. sphaerocephala suggests that spatial patchiness is less of a constraint than with epiphytes.

Vascular epiphytes can be quite abundant in vegetation close to the ocean surf, where they are exposed to a more or less continuous input of salt spray. The ecophysiological basis of their occurrence, i.e. salt tolerance or avoidance, is unresolved, because all previous studies were observational and conclusions thus circumstantial. Here, the effect of varying concentrations of salt water on germination, and growth and survival of seedlings and established plants was investigated in a growth cabinet study under controlled conditions. Seeds (1500), seedlings (750) and small tank plants (336) were from four populations of Werauhia sanguinolenta that were growing either close to the sea or inland in Panama. Changes of Na+ and K+ concentrations in plant tissue were also determined. No differences in the sensitivity to salt were found among populations, nor among life stages. External concentrations (Cext) of up to 15% sea water (c. 0.5% Na+) allowed complete germination as well as positive growth and survival in both seedlings and established plants over short periods (8–10 wk). After longer exposure (12 wk) of established plants visible damage and increased mortality were observed at lower Cext, but critical tissue Na+ levels were similar: c. 50 mg gDW−1. It is concluded that this common epiphyte does not meet the definition of a halophyte, but still possesses a rather high tolerance to sodium.

The reciprocal importance of bromeliads and hummingbirds has been proposed for many years, even suggesting coevolution between these two groups. Nevertheless, data are lacking that allow a better test of the relationships involved. Here we investigate the relationship between bromeliads and hummingbirds in an area of secondary Atlantic rain forest in southern Brazil. The study examined the interactions among 12 species of bromeliad and 10 of hummingbird at Reserva Natural Salto Morato, Paraná state. The number of flowering species of bromeliad and the species richness and abundance of hummingbirds were quantified monthly between November 2004 and October 2005. Focal observations on each bromeliad species were made to determine the hummingbird visitors. Neither species richness nor abundance of hummingbirds were related to bromeliad phenology. Together with the monthly variation in visit frequency by a given pollinator to a given plant, these factors indicate a generalization in the use of bromeliads by hummingbirds and argue against tight coevolution. Ramphodon naevius and Thalurania glaucopis were the main pollinators in the community. Aechmea nudicaulis was the most generalist bromeliad species. The generalist species interacted with other generalists or with asymmetric specialists and there was no specialist–specialist interaction. This produced a strongly organized and nested matrix of interactions. This nestedness is similar to other plant-pollinators networks, supporting the hypothesis that the evolutionary relationship between bromeliads and hummingbirds is no stronger than that of other pollination networks.

We studied the interaction between the bromeliad Werauhia gladioliflora and its flower visitors in the Caribbean lowland forest of Costa Rica, in order to quantify the mutual benefits to both partners. Over 6 y, the bromeliads flowered mainly between October and December; with an individual inflorescence flowering for an average of 34 d (n = 233 inflorescences). The bromeliad showed a flexible breeding system with autogamy occurring in addition to cross-pollination. Exclusive pollinators were small nectar-feeding bats (Phyllostomidae: Glossophaginae). The average volume of nectar produced per flower was 1.1 ml (n = 25 flowers). The main visitor was the bat Glossophaga commissarisi, which approached the flowers exclusively using hovering flight. Visitation by bats, measured by infrared light sensors, occurred throughout the night with an activity peak after midnight. Median hovering duration of the bats at the flowers was 320 ms (n = 1246 visits). Hourly mean of hovering duration was negatively correlated with hourly nectar secretion rate. The flower visitation behaviour of a bat over the night seems to be shaped by a combination of intrinsic physiological factors and by nectar availability. Size of both flowers and visitors make Werauhia gladioliflora a very accessible system for quantification of factors affecting evolution of bat–plant interactions.