Polymorphic microsatellites for the study of fragmented populations of Pitcairnia geyskesii L. B. Smith (Bromeliaceae), a specific saxicolous species of inselbergs in French Guiana

Pitcairnia geyskesii is a saxicolous bromeliad endemic to the granitic inselbergs of French Guiana and Surinam. Seven polymorphic microsatellite markers were characterized to investigate the polymorphism among individuals sampled on five inselbergs in French Guiana. The total number of alleles per locus varied from four to 14. Mean observed heterozygosities at each microsatellite locus ranged from 0.114 to 0.629. This set of microsatellites offers an efficient tool for more detailed investigations on the genetic structure of fragmented populations living on such a singular and insular habitat within tropical rainforest.     

A set of polymorphic microsatellite loci for Vriesea gigantea and Alcantarea imperialis (Bromeliaceae) and cross‐amplification in other bromeliad species

Fifteen polymorphic microsatellite markers were isolated and characterized in two species of Bromeliaceae: Vriesea gigantea and Alcantarea imperialis. The number of alleles observed for each locus ranged from three to 16. The loci will be used for studies of the genetic structure of natural populations, reproductive biology, and evolutionary relationships among and within these genera. A cross‐amplification test in 22 taxa suggests that the markers will be useful for similar applications in numerous other bromeliad species.     

Reproductive cycle of Pitcairnia flammea Lindl. (Bromeliaceae/Pitcairnioideae) in an insular Atlantic rainforest area in southeastern Brazil

We studied for 6 years the reproductive cycle of the bromeliad Pitcairnia flammea in the Atlantic Rainforest of Ilha Grande, an island located in the south of Rio de Janeiro state, southeastern Brazil. We marked 150 adult individuals of the species, for which we monthly observed the occurrence of reproductive characteristics. The phenological attributes were related to environmental factors such as photoperiod, rainfall and temperature. Additionally, we estimated a mortality rate that was related to the rainfall in the area. P. flammea possesses a seasonal reproduction pattern, with a relatively fixed and well-defined time span during the winter dry period; flowering precedes fruiting by approx. 1 month. Due to the occurrence of P. flammea preferably on boulders of river channels and banks mortality of the plants is highest when floods from strong rainfall remove the plants mechanically from their growing places.     

The role of hybridization and introgression in maintaining species integrity and cohesion in naturally isolated inselberg bromeliad populations

• Hybridization is a widespread phenomenon present in numerous lineages across the tree of life. Its evolutionary consequences range from effects on the origin and maintenance, to the loss of biodiversity.
• We studied genetic diversity and intra‐ and interspecific gene flow between two sympatric populations of closely‐related species, Pitcairnia flammea and P. corcovadensis (Bromeliaceae), which are adapted to naturally fragmented Neotropical inselbergs, based on nuclear and plastidial DNA.
• Our main results indicate a strong reproductive isolation barrier, although low levels of interspecific gene flow were observed in both sympatric populations. The low rates of intraspecific gene flow observed for both P. corcovadensis and P. flammea populations corroborate the increasing body of evidence that inselberg bromeliad species are maintained as discrete evolutionary units despite the presence of low genetic connectivity. Nuclear patterns of genetic diversity and gene flow revealed that hybridization and introgression might not cause species extinction via genetic assimilation of the rare P. corcovadensis.
• In the face of reduced intraspecific gene exchange, hybridization and introgression may be important aspects of the Pitcairnia diversification process, with a positive evolutionary impact at the bromeliad community level, and thus contribute to increasing and maintaining genetic diversity in local isolated inselberg populations.

     

The Stability of Invertebrate Communities in Bromeliad Phytotelmata in a Rain Forest Subject to Hurricanes

Communities of invertebrate animals in lower canopy and saxicolous tank bromeliads, originally studied in 1993–1997, were resampled along an elevational gradient in tabonuco, palo colorado, and dwarf or cloud forest in Puerto Rico in 2010. These Puerto Rican montane rain forests were impacted strongly by hurricanes in 1989 and 1998, so the surveys in the 1990s represented 4–8 yr of post‐hurricane recovery, whereas our recent survey represents 12 yr of post‐hurricane recovery. At most elevations, species diversity, both within individual bromeliads and at the forest scale, declined between the 1990s and 2010. This decline in diversity between decades is associated with reductions in bromeliad density as the canopy progressively closed during recovery from hurricane damage. The observed decline in alpha and gamma diversity appears to have involved the loss of rarer species, as might be expected from standard metapopulation theory. By contrast, the most common species were remarkably stable in abundance, composition, and frequency of occurrence over the two decades. In the lowermost tabonuco forest, two endemic bromeliad specialists, restricted to bromeliads for their entire life cycle, were not found on resampling. This study also demonstrates that, at least in Puerto Rico, sets of ten plants from each forest were sufficient to monitor bromeliad invertebrate populations and their diversity over time.     

Interspecific plant functional variation prevails over intraspecific variation in driving spider beta diversity

1. Non-trophic interactions between plants and animals can affect community structure and species trait composition. However, it is unclear how changes in intra- and interspecific morphological traits of plant species affect non-trophic interactions at a metacommunity scale. Additionally, whether plant evolutionary history determines taxonomic and functional diversity of plant-dwelling predators is an open question. 2. To address these gaps, this study used a published dataset with spiders dwelling exclusively on bromeliads to investigate if: (i) intra- and interspecific variability in host plant morphological traits affects spider taxonomic and functional diversity; and (ii) bromeliad trait evolution determines present-day patterns of spider trait diversity. 3. Spider and bromeliad traits were measured, and a new statistical framework was used to quantify the response of spider beta diversity to intra- and interspecific variation in bromeliad traits. In addition, bromeliad traits were decomposed across its phylogenetic tree to check whether the current variation in morphological traits of bromeliads is a result of either ancestral or recent diversification. 4. Bromeliad intraspecific variation did not affect spiders, but leaf length variation between bromeliad species had a positive effect on spider functional beta diversity. Interestingly, the most ancestral split between two subfamilies explained most of the variation in bromeliad species, which suggests that spider functional diversity could represent an outcome of bromeliad evolutionary history. 5. Overall, the results of this study suggest that interactions between plants and organisms that do not feed directly on their tissues could be shaped by plant evolutionary history, which in turn suggests that non-trophic interactions can be maintained over time.     

Acclimation to future atmospheric CO2 levels increases photochemical efficiency and mitigates photochemistry inhibition by warm temperatures in wheat under field chambers

Urea is an important nitrogen source for some bromeliad species, and in nature it is derived from the excretion of amphibians, which visit or live inside the tank water. Its assimilation is dependent on the hydrolysis by urease (EC: 3.5.1.5), and although this enzyme has been extensively studied to date, little information is available about its cellular location. In higher plants, this enzyme is considered to be present in the cytoplasm. However, there is evidence that urease is secreted by the bromeliad Vriesea gigantea, implying that this enzyme is at least temporarily located in the plasmatic membrane and cell wall. In this article, urease activity was measured in different cell fractions using leaf tissues of two bromeliad species: the tank bromeliad V. gigantea and the terrestrial bromeliad Ananas comosus (L.) Merr. In both species, urease was present in the cell wall and membrane fractions, besides the cytoplasm. Moreover, a considerable difference was observed between the species: while V. gigantea had 40% of the urease activity detected in the membranes and cell wall fractions, less than 20% were found in the same fractions in A. comosus. The high proportion of urease found in cell wall and membranes in V. gigantea was also investigated by cytochemical detection and immunoreaction assay. Both approaches confirmed the enzymatic assay. We suggest this physiological characteristic allows tank bromeliads to survive in a nitrogen‐limited environment, utilizing urea rapidly and efficiently and competing successfully for this nitrogen source against microorganisms that live in the tank water.     

Bromeliad architectural complexity and vertical distribution predict spider abundance and richness

Habitat complexity is a main predictor of the distribution of arthropods on vegetation. However, it remains poorly known whether plant architecture and fine‐scale spatial distribution affect the species richness and composition of associated arthropod guilds. In this study, we extensively sampled bromeliad species with a variety of rosette architectures in a megadiversity region. The aims were to investigate whether (i) possible differences in spider species composition among bromeliad species are related to the distinct architectures of the plants, and (ii) bromeliad architectural complexity (an intrinsic feature) and vertical distribution (an extrinsic feature) are good predictors of spider abundance and richness. Contrary to our expectations, spider species composition did not vary significantly among bromeliad species with different architectures. We found a positive effect of the mean number of leaves on spider abundance and species richness, but it occurred indirectly through spider abundance; factoring out the indirect effect revealed a negative effect of leaf number on species richness. Bromeliad species with wider vertical distributions harboured more spider species. Our results suggest that the dominance of a few spider species and reduced space for orb‐web spiders to attach their webs are the main explanations for lower spider richness on bromeliad species with higher architectural complexity. Our findings highlight the importance of both intrinsic and extrinsic plant features as co‐determinants of predator arthropod diversity.     

Endemism and ecological islands: the ostracods from Jamaican bromeliads

1. The inhabitants of ecological islands, such as the phytolemata of bromeliad plants, provide an opportunity to examine the genetic patterns resulting from island radiations and draw inferences about modes of speciation.
2. Allozyme electrophoresis, as well as mitochondrial DNA and morphological analyses, were employed to delimit species boundaries and assess the population genetic structure of the ostracods from bromeliads on the island of Jamaica.
3. Nine species from the genus Elpidium were both sexually reproducing and endemic to the island. The other commonly encountered ostracod genus, Candonopsis , was represented by at least two sexually reproducing species.
4. Most Elpidium species showed very restricted distributions, low heterozygosity, and marked gene pool fragmentation, suggesting that population bottlenecks have occurred frequently. Whether founder events played a causal role in the diversification of these ostracods remains uncertain, but bromeliad habitats seem to be a source of high biodiversity, and may represent an ideal setting for extensive allopatric speciation.     

Endemism and ecological islands: the ostracods from Jamaican bromeliads

1. The inhabitants of ecological islands, such as the phytolemata of bromeliad plants, provide an opportunity to examine the genetic patterns resulting from island radiations and draw inferences about modes of speciation.
2. Allozyme electrophoresis, as well as mitochondrial DNA and morphological analyses, were employed to delimit species boundaries and assess the population genetic structure of the ostracods from bromeliads on the island of Jamaica.
3. Nine species from the genus Elpidium were both sexually reproducing and endemic to the island. The other commonly encountered ostracod genus, Candonopsis , was represented by at least two sexually reproducing species.
4. Most Elpidium species showed very restricted distributions, low heterozygosity, and marked gene pool fragmentation, suggesting that population bottlenecks have occurred frequently. Whether founder events played a causal role in the diversification of these ostracods remains uncertain, but bromeliad habitats seem to be a source of high biodiversity, and may represent an ideal setting for extensive allopatric speciation.     

Geographic Range,Habitats, and Host Plants of Bromeliad‐living Jumping Spiders (Salticidae)1

Although spiders are a very diverse group on vegetation, their associations with plants are poorly known. Some salticid species specifically use Bromeliaceae as host plants in some regions of South America. In this study, I report the geographic range of these spider‐bromeliad associations, and whether the spiders inhabit particular bromeliad species and vegetation types, as well as open areas or interior of forests. Nine salticid species were found to be associated with up to 23 bromeliad species in cerrados (savanna‐like vegetation), semideciduous and seasonal forests, coastal sand dune vegetation, restingas, inselbergs, highland forests, chacos, and rain forests at 47 localities in Brazil, Paraguay, Bolivia, and Argentina. Some species were typically specialists, inhabiting almost exclusively one bromeliad species over a large geographic range (e.g., Psecas chapoda on Bromelia balansae), whereas others were generalists, occurring on up to 7–8 bromeliad species (e.g., Psecas sp., Eustiromastix nativo, and Coryphasia sp. 1). The regional availability of bromeliad species among habitats may explain this pattern of host plant use. More jumping spiders were found on bromeliads in open areas than on bromeliads in the interior of forests. These results show that several jumping spider species may be strictly associated with the Bromeliaceae in the Neotropics. This is one of the few studies to show host‐specific associations for spiders on a particular plant type over a wide geographic range.     

Reproductive biology and natural hybridization between two endemic species of Pitcairnia (Bromeliaceae)

We investigated pollination biology and breeding systems in hybridizing populations of Pitcairnia albiflos and P. staminea; both species are endemic to rocky outcrops at Rio de Janeiro, Brazil. These species are morphologically distinct and easily recognized by floral color: white in P. albiflos and red in P. staminea. Putative hybrids show a large range of intermediate pink floral colors. The showy hermaphroditic flowers offer pollen and nectar that attract many visitors including bees, butterflies, hawk moths, and bats. Although the flowers of both parental species and hybrids open at night, only P. albiflos had other adaptations for nocturnal pollination. Flowering times overlapped during three consecutive years of observation. Bees visited both species and putative hybrids. Cross-pollinations were performed within and among parental species and hybrids in a greenhouse using plants transplanted from the field. Pitcairnia staminea and hybrids are self-compatible and could be spontaneously self-pollinated, whereas P. albiflos, though self-compatible, needs pollinators' services for self-pollination. Facultative agamospermy was found in the parental species. Prezygotic and postzygotic reproductive barriers between these taxa were weak. Reciprocal hand-pollinations between parental species and with hybrids yielded high fruit sets with viable seeds. Evaluations of fruit set, seed set, seed germination, and pollen viability were undertaken to compare the fitness of the hybrids relative to their parents. The hybrids showed equivalent fitness, except for lower pollen viability. Some conservation implications are noted.     

Distribution and Flowering Ecology of Bromeliads along Two Climatically Contrasting Elevational Transects in the Bolivian Andes1

We compared the diversity, taxonomic composition, and pollination syndromes of bromeliad assemblages and the diversity and abundance of hummingbirds along two climatically contrasting elevational gradients in Bolivia. Elevational patterns of bromeliad species richness differed noticeably between transects. Along the continuously wet Carrasco transect, species richness peaked at mid‐elevations, whereas at Masicurí most species were found in the hot, semiarid lowlands. Bromeliad assemblages were dominated by large epiphytic tank bromeliads at Carrasco and by small epiphytic, atmospheric tillandsias at Masicurí. In contrast to the epiphytic taxa, terrestrial bromeliads showed similar distributions across both transects. At Carrasco, hummingbird‐pollination was the most common pollination mode, whereas at Masicurí most species were entomophilous. The proportion of ornithophilous species increased with elevation on both transects, whereas entomophily showed the opposite pattern. At Carrasco, the percentage of ornithophilous bromeliad species was significantly correlated with hummingbird abundance but not with hummingbird species richness. Bat‐pollination was linked to humid, tropical conditions in accordance with the high species richness of bats in tropical lowlands. At Carrasco, mixed hummingbird/bat‐pollination was found especially at mid‐elevations, i.e., on the transition between preferential bat‐pollination in the lowlands and preferential hummingbird‐pollination in the highlands. In conclusion, both richness patterns and pollination syndromes of bromeliad assemblages varied in distinct and readily interpretable ways in relation to environmental humidity and temperature, and bromeliad pollination syndromes appear to follow the elevational gradients exhibited by their pollinators.     

Micro-environmental factors and the endemism of bromeliad aquatic fauna

Tank bromeliads harbour aquatic microcosms with many endemic species among their leaves. We performed a set of experiments to determine which factors maintain the bromeliad aquatic fauna in isolation from neighbouring ponds. We cultivated three invertebrates species (an ostracod, an annelid and a cladoceran) from a pond surrounded by terrestrial bromeliads in Southeastern Brazil and introduced them inside cleaned bromeliads, using recipients with the same volume as controls. The pH, conductivity and organism densities were monitored in the bromeliad samples and controls for 41 days. The samples introduced inside the cleaned bromeliads showed a significant decrease in pH and conductivity compared to the controls. The pond organism populations introduced in the bromeliads presented a high extinction rate and a significant population decrease when compared to the ones introduced in the controls. We attributed the population decline experienced by the pond organisms to the oligotrophic conditions generated inside the tanks by the bromeliads due to the nutrient absorption. We suggest that the changes in water chemistry induced by the bromeliads could play an important role in isolating their microcosm communities from other freshwater systems. Other mechanisms that could produce the high rates of endemism in bromeliad fauna are discussed. Handling editor: K. Martens     

Genetic variability of an endangered Bromeliaceae species (Pitcairnia albiflos) from the Brazilian Atlantic rainforest

Pitcairnia albiflos is a Bromeliaceae species endemic to Brazil that has been included as data-deficient in the extinction risk list of Brazilian flora. We analyzed genetic variability in P. albiflos populations using RAPD markers to investigate population structure and reproductive mechanisms and also to evaluate the actual extinction risk level of this species. Leaves of 56 individuals of P. albiflos from three populations were collected: Urca Hill (UH, 20 individuals), Chacrinha State Park (CSP, 24 individuals) and Tijuca National Park (TNP, 12 individuals). The RAPD technique was effective in characterizing the genetic diversity in the P. albiflos populations since it was possible to differentiate the populations and to identify exclusive bands for at least two of them. Even if there is low genetic diversity among them (CSP-UH = 0.463; CSP-TNP = 0.440; UH-TNP = 0.524), the populations seem to be isolated according to the low genetic diversity observed within them (H(pop) CSP = 0.060; H(pop) UH = 0.042; H(pop) TNP = 0.130). This fact might be the result of clonal and self-reproduction predominance and also from environmental degradation around the collection areas. Consequently, it would be important to protect all populations both in situ and ex situ to prevent the decrease of genetic variability. The low genetic variability among individuals of the same population confirms the inclusion of this species as critically endangered in the risk list for Brazilian flora.     

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.     

Sympatric bromeliad species (Pitcairnia spp.) facilitate tests of mechanisms involved in species cohesion and reproductive isolation in Neotropical inselbergs

The roles of intra- and interspecific gene flow in speciation and species evolution are topics of great current interest in molecular ecology and evolutionary biology. Recent modelling studies call for new empirical data to test hypotheses arising from the recent shift from a 'whole-genome reproductive isolation' view to a 'genic' view of species and speciation. Particularly scarce (and thus of particular interest) are molecular genetic data on recently radiated, naturally hybridizing species in strongly structured and species-rich environments. Here, we studied four sympatric plant species (Pitcairnia spp.; Bromeliaceae) adapted to Neotropical inselbergs (isolated outcrops resembling habitat 'islands' in tropical rainforests) using nuclear and plastid DNA. Patterns of plastid DNA haplotype sharing and nuclear genomic admixture suggest the presence of both, incomplete lineage sorting and interspecific gene flow over extended periods of time. Integrity and cohesion of inselberg species of Pitcairnia are maintained despite introgression and in the face of extremely low within-species migration rates (N(e)m < 1 migrant per generation). Cross-evaluation of our genetic data against published pollination experiments indicate that species integrity is maintained by the simultaneous action of multiple prezygotic barriers, including flowering phenology, pollinator isolation and divergent mating systems. Postzygotic Bateson-Dobzhansky-Muller incompatibilities appear to contribute to isolation, as suggested by asymmetric introgression rates of single loci. Our results suggest that incomplete lineage sorting, hybridization and introgression form integral aspects of adaptive radiation in Neotropical inselberg 'archipelagos'. Inselbergs with multiple closely related co-occurring species should be of special interest to students of speciation in mountain systems, and to ongoing conservation programmes in the Atlantic Rainforest biodiversity hotspot.     

Genetic variation in Aechmea winkleri,a bromeliad from an inland Atlantic rainforest fragment in Southern Brazil

We investigated the genetic diversity and structure of Aechmea winkleri Reitz, an endemic bromeliad found in Southern Brazilian Atlantic rainforest. Seven nuclear microsatellite markers were used to analyze 162 samples from four localities sampled throughout the entire geographic distribution of the species. Results indicate relatively high levels of genetic diversity with an average of allelic richness of 3.57, and observed and expected heterozygosity of 0.559 and 0.608, respectively. The within-inbreeding coefficient was low, ranging from −0.011 to 0.094. All localities significantly deviated from Hardy–Weinberg equilibrium with three of them showing heterozygosites deficiency. Most of the genetic variation (96.64%) was found within localities. No reduction in population size (bottleneck) was detected. Low levels of genetic differentiation among localities were found with pairwise F_ST comparisons varying from 0.021 to 0.075. Bayesian analyses revealed that A. winkleri is composed by two genetic groups. The number of migrants per generation was high (>1), which maintain localities' cohesion and gene flow. Despite genetic erosion was not detected in the present study, our results revealed that the four localities sampled actually represent one population, the only known so far in nature. Management strategies for A. winkleri conservation should be undertaken as it is an endemic species which occurs in a biome that has gone through major deforestation and fragmentation. This would avoid the increase of inbreeding rates and the loss of genetic diversity.