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According to recent taxonomic treatments, Bromeliaceae subfamily Pitcairnioideae comprise the genera Deuterocohnia, Dyckia, Encholirium, Fosterella and Pitcairnia. We present a dated molecular phylogenetic analysis of a comprehensive taxon set to enable inferences of evolution in the subfamily. Phylogenetic relationships in Pitcairnioideae were reconstructed based on three plastid loci (rpl32-trnL, rps16-trnK, 5′ end of matK) and the nuclear single-copy gene PHYC exon 1, using maximum parsimony, maximum likelihood and Bayesian inference. The plastid phylogenetic analysis supports the monophyly of the subfamily, whereas the nuclear DNA data suggest a nested position of Puyoideae in Pitcairnioideae. Fosterella and Dyckia (including Encholirium) are monophyletic in both analyses, whereas Deuterocohnia is paraphyletic and Pitcairnia remains unresolved in the plastid tree. A Bayesian relaxed clock model applied to the plastid data indicates a diversification of Fosterella and Pitcairnia at c. 6-8 Mya, whereas Deuterocohnia and Dyckia might have diversified at about 2-4 Mya. Our data support the concept that Pitcairnioideae originated in the Andes, followed by numerous dispersal events to South and Central America. Plastid capture events might explain the contrasting topologies of plastid and nuclear trees in Deuterocohnia, whereas the position of Puya needs to be re-evaluated with additional nuclear markers.
E apresentada a revisão taxonômica de Encholirium, gênero exclusivamente brasileiro cuja distribuição está intimamente associada a afloramentos rochosos, sendo que a maioria das espécies é encontrada nos afloramentos rochosos do cerrado e caatinga, e nos campos rupestres da Cadeia do Espinhaço de Minas Gerais. Apenas duas espécies ocorrem nos inselbergs dentro dos domínios da floresta atlântica. No presente tratamento são reconhecidas para o gênero 23 espécies. Esta sendo sugerida a transferência de Dyckia bifiora para Encholirium, uma nova espécie esta sendo descrita, dois nomes permaneceram duvidosos e 13 binômios estão sendo colocados em sinonímia. Os caracteres que se mostraram mais relevantes para a circunscrição das espécies de Encholirium foram: hábito (incluindo altura da planta, diâmetro e aspecto geral da roseta), tamanho das brácteas do escapo, tamanho das brácteas florais em relação ás flores, comprimento do pedicelo, forma e coloração das sépalas e pétalas, e posição dos estames e do estilete em relação á corola. São apresentadas chaves de identificação para as espécies de Encholirium, descrições, ilustrações, distribuição geográfica, comentários sobre aspectos taxonômicos e nomenclatórios, bem como alguns aspectos referentes á fenologia e conservação de cada espécie.
Encholirium horridum is a bromeliad that occurs exclusively on inselbergs in the Atlantic Forest biome of Brazil. These rock outcrops form natural islands that isolate populations from each other. We investigated gene flow by pollen through paternity analyses of a bromeliad population in an area of approximately 2 ha in Espírito Santo State, Brazil. To that end, seed rosettes and seedlings were genotyped using nuclear microsatellite loci. A plot was also established from the same population and specimens were genotyped to evaluate their fine-scale spatial genetic structure (SGS) through analyses of spatial autocorrelation and clonal growth. Paternity analysis indicated that 80% of the attributed progenitors of the genotyped seedlings were from inside the study area. The pollen dispersal distances within the area were restricted (mean distance of 45.5 m, varying from 3 to 156 m) and fine-scale SGS was weak (F ij = 0.0122, P < 0.001; Sp = 0.009). Clonal growth was found to be a rare event, supporting the monocarpy of this species.
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Among angiosperms, bromeliads are remarkable for their floral biology, since
most of the species appear to be primarily pollinated by vertebrates, rather than insects
(Krömer et al. 2006). Most other animal-pollinated plants, including the orchids, are
pollinated largely by insects. In the Neotropics, where almost all bromeliad species occur,
hummingbirds and bats are the most important vertebrate pollinators (Fleming et al.
2005). Hummingbirds are the most frequently reported bromeliad-pollinators (Kessler
& Krömer 2000; Carranza-Quiceno & Estévez-Varón 2008), resulting in a ratio of up
to 20 bird-pollinated (“ornithophilous”) to one bat-pollinated (“chiropterophilous”)
species (Sazima et al. 1999). Here, we present a brief overview of the knowledge about
bat-pollination in Bromeliaceae, with special focus on a recent study carried out by
our own working group.
The genus Dyckia (Bromeliaceae) comprises more than 150 terrestrial or epilithic species with a strongly xeromorphic habit. Most of its members belong to the azonal rock vegetation of Neotropical savannas and forests of Brazil and adjacent countries. Dyckia is relatively spe-cies-rich compared with its closest relatives Encholirium (27 species) and Deuterocohnia (17 species). Here, we present the first molecular phylogenetic analysis of Dyckia using DNA sequence data from six plastid loci (matK gene, rps16 intron, petD intron, rpl32-trnL, rps16-trnK and trnD-trnT) and a portion of the nuclear gene phyC. A total of 124 accessions were included, corresponding to 79 taxa from six genera. Phylogenetic trees were generated using par-simony, likelihood and Bayesian methods. DNA sequence variation among Dyckia species turned out to be extremely low, and phylogenies were poorly resolved. The mono-phyly of Dyckia is supported, whereas evidence is provided that Encholirium is paraphyletic. Based on a dated plastid DNA tree, Dyckia experienced a recent radiation starting around 2.9 million years ago. Four major clades could be identified that roughly correspond to the geographic origin of the samples. A parsimony network based on plastid DNA haplotypes shows a star-like pattern, indicating recent range expansions. Our data are compatible with a scenario where Dyckia and Encholirium diverged in northeastern Brazil, whereas one lineage of Dyckia dis-persed to southern Brazil from where a rapid colonization of suitable habitats was initiated. We discuss our results in relation to species delimitation in Dyckia.
Summary: A new species of Pilosocereus subg.
Gounellea, P. frewenii, is described and illustrated
for the first time. It is a dwarf, geographically disjunct
relative of P. gounellei and, in contrast to
other members of its subgenus, occurs in dense
dry forest (on limestone rock outcrops) in an area
situated between the Atlantic Forest and savanna
(cerrado) biomes, far distant from the caatinga.
Its phytogeographical significance is discussed
and conservation status assessed as Critically
Endangered (CR).
Foram estudadas plantas ornitófilas em uma área de caatinga da Reserva Particular do Patrimônio Natural Cantidiano Valgueiro (8º36'00" S e 38º34'5" W) em Pernambuco, Nordeste do Brasil, no período de julho de 2002 a junho de 2003. Oito espécies ornitófilas foram registradas no período de estudo, distribuídas em sete gêneros e cinco famílias. Cactaceae foi a família com maior número de espécies polinizadas por beija-flores, sendo representada por três espécies, seguida de Bromeliaceae, com duas espécies. Foram registradas espécies em floração durante todo o ano. Com exceção de Bromelia laciniosa Mart. ex Schult. f. e Neoglaziovia variegata (Arruda) Mez (Bromeliaceae), todas as demais espécies tiveram pico de floração no período seco. Metade das espécies ornitófilas da comunidade estudada apresentou hábito herbáceo. A maioria das espécies apresentou flores vermelhas (62,5%), sendo o tipo tubo registrado em todas as espécies. O comprimento médio do tubo da corola foi 20,2 ± 5,6 mm, a concentração de açúcares no néctar variou de 18% a 33,5% e o volume de 22 a 41 µL. Cinco espécies de beija-flores foram registradas visitando as flores da comunidade estudada, das quais apenas uma foi residente. Chlorostilbon aureoventris (d'Orbigny & Lafresnaye, 1838), devido ao seu comportamento e freqüência de visita, foi considerada como a espécie dominante da comunidade. Comparações com estudos semelhantes evidenciaram que o número de espécies de plantas ornitófilas que floresceram no período de estudo na Reserva foi expressivamente menor do que o encontrado em estudos em remanescentes de Mata Atlântica e áreas neotropicais em geral. Além disso, uma espécie de Trochilinae, e não de Phaethornithinae, atuou como organizadora da comunidade.
The many-flowered, brush-like spikes ofEncholirium glaziovii, a ground-dwelling pitcairnioid bromeliad of the campo rupestre formation of southeastern Brazil, was observed being pollinated by the glossophagine bat,Lonchophylla bokermanni, in the Serra do Cip (Minas Gerais). Nectar feeding was while hovering, and the pollen was preferentially transferred by the bat''s snout. The floral pattern is chiropterophilous; unlike known tillandsioid bat flowers, stamens and style are protrusive beyond a small, persistent perigon, and anthesis, apparently protogynous, extends over several nights, with gradual onset and cease. Although various other flower-visitingChiroptera are known to occur in the area and are not mechanically precluded from exploitingEncholirium glaziovii, no one seems to compete withLonchophylla bokermanni. This species, a trap-liner, habitually forages in open habitats, a possible reason for its exclusiveness.—Flower details of two other species ofEncholirium from the same locality including one with the bat pollination syndrome, are given.
Encholirium is a Brazilian genus of Bromeliaceae that occurs exclusively in rocky landscapes. This work aimed to generate basic information for the conservation of three Encholirium species that are endemic to the rocky mountains of Cadeia do Espinhaço, employing population genetic analyses. E. pedicellatum and E. biflorum have only one very small population each, both occurring in unprotected, private land sites, being critically endangered. E. subsecundum is more widespread, and some of its populations dwell in protected areas. Five Random Amplified Polymorphic DNA (RAPD) primers generated approximately 60 polymorphic bands for each species. This technique demonstrated the presence of a single RAPD profile for every individual sampled (except for one clone found in E. biflorum). High levels of genetic variability were not expected, due to the clonal habit of the plants and small size of the populations. Populations of E. biflorum and E. pedicellatum presented, respectively, 16.06% (Φst
= 0.16, p<0.001) and 8.44% (Φst
= 0.08, p<0.001) of the total genetic diversity attributable to genetic differences among groups within the populations. In E. subsecundum, 14.52% (Φst
= 0.15, p<0.001) of the total genetic diversity was found among populations. Estimates of the Shannon’s Diversity Index provided similar results. These results are valuable for the development of conservation strategies.
A few years after the publication of the revision of Encholirium by the first author, two new species have been found in neighbouring municipalities on the west side of the Serra do Cipó,
Cadeia do Espinhaço, in Minas Gerais. Both have a quite restricted habitat, growing on quartzitic/arenitic rocks above 1000m
alt., within 10 – 12km of each other. Encholirium agavoides Forzza & Zappi has striking silvery leaves and a small, agave-like habit while Encholirium ctenophyllum Forzza & Zappi also has small rosettes that can be distinguished by their narrow, strongly pectinate and reflexed leaves.
A key for the species in the region is presented.
Passados apenas alguns anos após a publicação da revisão do gênero Encholirium estas duas espécies aqui apresentadas foram encontradas em municípios vizinhos na encosta oeste da Serra do Cipó, na Cadeia
do Espinhaço, Minas Gerais, sendo ambas restritas aos afloramentos de quartzito e arenito, acima de 1000m s.n.m., a aproximadamente
10 – 12km uma da outra. Encholirium agavoides Forzza & Zappi possui pequenas folhas prateadas agrupadas em rosetas com aspecto de Agave, enquanto que as rosetas, também relativamente pequenas de Encholirium ctenophyllum Forzza & Zappi apresentam folhas estreitas, fortemente pectinadas e reflexas. Uma chave de identificação para as espécies
da região é apresentada.
Key wordsBromeliaceae–Eastern Brazil–
Encholirium
–IUCN categories–new species–Pitcairnioideae–Serra do Cipó–taxonomy
We studied hummingbirds and their food plants in an area of caatinga vegetation. We specifically examined their seasonal use of this habitat, migratory and non-migratory status, their foraging strategies and community roles The study was conducted in an area of arboreal-shrub caatinga, located in the Serra do Pará, municipality of Santa Cruz do Capibaribe, state of Pernambuco in northeastern Brazil. Field work was undertaken during 12 expeditions on a monthly basis between June, 2007 and May, 2008. Five species of hummingbirds were recorded visiting flowers in the community studied. Three were considered residents: Chlorostilbon lucidus (Shaw, 1812), Eupetomena macroura (Gmelin, 1788), and Heliomaster squamosus (Temminck, 1823). Hummingbirds visited 31 species of plants, of which only five presented attributes related to ornithophily. C. lucidus visited 29 plant species, including all ornithophilous species, and it was the most aggressive, defending territories. Among hummingbirds, C. lucidus may be considered the principal pollinator. Hummingbirds may also be acting as pollen vectors for some of the plant species not identified as ornithophilous. The hummingbird guilds varied among the plant species used as floral resources, as well as in their frequency of visits. Differences in plant species abundance, hummingbird preference, competitive exclusion or flowering seasonality are factors likely to influence those variations.
Premise of the study:
Inselbergs are terrestrial, island-like rock outcrop environments that present a highly adapted flora. The epilithic bromeliad Encholirium spectabile is a dominant species on inselbergs in the Caatinga of northeastern Brazil. We conducted a population genetic analysis to test whether the substantial phenotypic diversity of E. spectabile could be explained by limited gene flow among populations and to assess the relative impact of pollen vs. seed dispersal on the genetic structure of the species.
Methods:
Nuclear and chloroplast microsatellite markers were used to genotype E. spectabile individuals from 20 rock outcrop locations, representing four geographic regions: northern Espinhaço Range, Borborema Plateau, southwestern Caatinga and southeastern Caatinga. F-statistics, structure, and other tools were applied to evaluate the genetic makeup of populations.
Key results:
Considerable levels of genetic diversity were revealed. Genetic structuring among populations was stronger on the plastid as compared with the nuclear level, indicating higher gene flow via bat pollination as compared with seed dispersal by wind. structure and AMOVA analyses of the nuclear data suggested a high genetic differentiation between two groups, one containing all populations from the southeastern Caatinga and the other one comprising all remaining samples.
Conclusions:
The strong genetic differentiation between southeastern Caatinga and the remaining regions may indicate the occurrence of a cryptic species in E. spectabile. The unique genetic composition of each inselberg population suggests in situ conservation as the most appropriate protection measure for this plant lineage.
Famine Foods of Brazil’s Seasonal Dry Forests: Ethnobotanical and Nutritional Aspects. Famine foods are used by people in times of food scarcity. Although the northeast of Brazil regularly faces periods of drought that cause a shortage of traditional foods, the use of famine foods is insufficiently recorded. The purpose of this study was to record the knowledge of famine food in two rural Caatinga communities, Carão and Cachoeira, in the states of Pernambuco and Paraiba, respectively, and to determine the chemical composition of the major famine foods utilized. The ethnobotanical aspect was divided into two stages: free lists and semistructured interviews. Based on the interviews, the main famine foods known by communities were selected for nutritional analysis. The study revealed that certain foods are used by communities only in times of shortages. Carão showed a higher species richness than Cachoeira, but the food knowledge was more widespread among the interviewees in Cachoeira. In Carão, the main species mentioned were Dioclea grandiflora and Manihot dichotoma, whereas in Cachoeira, Encholirium spectabile and Pilosocereus gounellei were the major species indicated. The species examined had a high carbohydrate content, especially D. grandiflora, Manihot glaziovii, and M. dichotoma, and could be used for complementary feeding of the population if adverse effects could be minimized. Mandevilla tenuiflora is indicated for future studies of potential agro-industrial applications and the beneficial aspects of its root, which could represent an alternative source of income for the population.
Shifts in pollen vectors favour diversification of floral traits, and differences in pollination strategies between congeneric sympatric species can contribute to reproductive isolation. Divergence in flowering phenology and selfing could also reduce interspecific crossing between self-compatible species. We investigated floral traits and visitation rates of pollinators of two sympatric Encholirium species on rocky outcrops to evaluate whether prior knowledge of floral characters could indicate actual pollinators. Data on flowering phenology, visitation rates and breeding system were used to evaluate reproductive isolation. Flowering phenology overlapped between species, but there were differences in floral characters, nectar volume and concentration. Several hummingbird species visited flowers of both Encholirium spp., but the endemic bat Lonchophylla bokermanni and an unidentified sphingid only visited E. vogelii. Pollination treatments demonstrated that E. heloisae and E. vogelii were partially self-compatible, with weak pollen limitation to seed set. Herbivores feeding on inflorescences decreased reproductive output of both species, but for E. vogelii the damage was higher. Our results indicate that actual pollinators can be known beforehand through floral traits, in agreement with pollination syndromes stating that a set of floral traits can be associated with the attraction of specific groups of pollinators. Divergence on floral traits and pollinator assemblage indicate that shifts in pollination strategies contribute to reproductive isolation between these Encholirium species, not divergence on flowering phenology or selfing. We suggest that hummingbird pollination might be the ancestral condition in Encholirium and that evolution of bat pollination made a substantial contribution to the diversification of this clade.
Fragments of the Atlantic forest of Northeast Brazil: Biodiversity, conservation and the Bromeliads
- J A Siqueiro Filho
- E M C Leme
- JA Siqueiro Filho
Ein [sic!] neue felsbewohnende Art der Gattung Encholirium aus dem äussersten Norden von Minas Gerais, Brasilien/A new rupicolous species of Encholirium from the uppermost north of Minas Gerais, Brazil: Encholirium viride
- P J Braun
- E Pereira
- PJ Braun
Eine neue, isoliert wachsende Encholirium Art aus dem Norden von Goiás (Zentral-Brasilien)/A new isolated growing species of Encholirium from northern Goiás (central Brazil): Encholirium josinoi-narcisae
- P J Braun
- E Pereira
- PJ Braun