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The Re-Establishment of Andrea (Bromeliaceae: Bromelioideae), a Monotypic Genus from Southeastern Brazil Threatened with Extinction
Abstract
A cladistic analysis of morphological data for the genus Canistropsis is presented. With the exception of two species, C. selloana and C. correia-araujoi, results support Canistropsis as monophyletic. Canistropsis correia-araujoi is a suspected, natural bigeneric hybrid. Canistropsis selloana is transferred back into the monotypic genus Andrea. A taxonomic treatment, including a greatly emended description, and discussion of the distribution, habitat, and conservation of Andrea selloana is presented.
... Brown & Leme, 2005). Leme (1997) described Edmundoa based on species of Canistrum and Nidularium with remarkably lanate peduncles and inflorescences. ...
... Canistropsis. A few years later, one of the species of this new genus, Canistropsis selloana (Baker) Leme, was considered morphologically singular enough (Brown & Leme, 2005;Leme et al., 2008) to be segregated as the monotypic genus Eduandrea, positioned outside the Nidularioid Complex. As a result of this turbulent taxonomic history, the Nidularioid Complex currently comprises six genera: Canistrum, Canistropsis, Edmundoa, Neoregelia, Nidularium and Wittrockia (Fig. 1). ...
... In order to assess the circumscription of and relationships within the Nidularioid Complex, we also sampled two species of Canistropsis, three of Canistrum, three of Edmundoa, six of Nidularium, and three of Wittrockia, including their respective type species (Appendix 1). Due to their intricate taxonomic history and formerly assumed relationship with the Nidularioid Complex (Leme, 1997(Leme, , 1998Brown & Leme, 2005;Leme et al., 2008), we included one species of Eduandrea and three of Aechmea [A. mollis L. B. Sm., r e l a t i o n s h i p w i t h N e o re g e l i a s u b g . ...
The ‘Nidularioid Complex’ is a group within the Bromelioideae usually characterized by inflorescences nested within water-impounding foliar rosettes. Currently, it comprises six genera: Canistrum, Canistropsis, Edmundoa, Neoregelia, Nidularium, and Wittrockia. While most of these genera occur in eastern Brazil, the distribution of Neoregelia is disjunct between the Atlantic Rainforest and Amazonia. Previous phylogenetic studies have not addressed the monophyly of and relationships among these genera; therefore, we undertook a phylogenetic study of the Nidularioid Complex with emphasis on the genus Neoregelia and its subgenera. A parsimony-based phylogenetic analysis with 101 morphological characters retrieved the Nidularioid Complex as non-monophyletic. Nidularium and Edmundoa were monophyletic. Neoregelia was recovered as non-monophyletic due to the inclusion within it of the Amazonian subgenus Hylaeaicum. These results highlight the need for revision of the generic classification of Bromelioideae, pending increased sampling of taxa and characters.
... In the red tist of endangered species of Minas Gerais' flora facing extinction (Mendonça and Lins 2000), Eduandreaselloanais cited by its synonymy, Nidulariumselloanum (Baker) E. Pereira & Leme, as endangered. (Brown and Leme 2005) consider it dose to extinction. ...
... Eduandrea selloanacan be characterized by its terrestrial habit, thick rhizome with an abundant root system; abaxial surface of the leaf, scape and primary bract covered with a thick layer of white tricome in contrast with the glabrous adaxial surface, green leaves and blades with entire foliar margins; well elevated inflorescence above the foliar rosette, with narrow primary bracts that don't form outlying or central tanks and without the capacity of storing water; rachis partially visible in the basal portion of the inflorescence because of the fascicule separation; biporate and supoliporate pollen with reticule exine, and subglobose fruit bigger than the ovaries, green with persistent chalice (Brown & Leme, 2005) (Figure 2). ...
... Eduandreaselloanais an endemic species of Espinhaço range, growing exclusively in riparian forest of the state of :Minas Gerais (figure 1). Despite the citations of occurrence in Ouro Preto and Antônio Pereira, even recently, its populations could only be found in Parque Nacional da Serra do Cipó (National Parque of Serra do Cipó), in populations considered residual, according to (Leme 1998 (Brown & Leme, 2005). The APE-Mutuca, is located in the municipal district of Nova Lima, MG, in the coordinates 43°58' W and 20°02' 30"S, with maximum altitude of 1,448m and an area of 1,250 ha. ...
... Within the subfamily, species are assigned to genera mostly on the basis of floral characteristics, with a major division made between those species that have symmetric versus asymmetric sepals (Smith, 1988). Recent re-examinations of some Bromelioideae species have made progress in defining monophyletic lineages, such as the identification of a Nidularioide lineage (Leme, 2000a), the confirmation of the monophyly of the genus Lymania based on a morphological and chloroplast gene region analyses (Faria et al., 2004b;de Oliveira et al., 2007;de Sousa and Wendt, 2008), and the re-establishment of the genus Andrea (Brown and Leme, 2005). However, phylogenies resulting from the analysis of chloroplast gene regions, AFLPs, and morphological characters for a more extensive sampling within Bromelioideae (Faria et al., 2004b;Schulte et al., 2005;Horres et al., 2007) have low resolution and/or lack support for the monophyly of many recognized genera. ...
... The current study focuses within the core Bromelioideae on the genus Aechmea, including those genera that have shown affinity or close relationship to Aechmea in previous morphological, chloroplast or AFLP based studies (Izquierdo and Pinero, 1998;Duval et al., 2003;Faria et al., 2004b;Brown and Leme, 2005;Schulte et al., 2005Schulte et al., , 2009de Oliveira et al., 2007;Horres et al., 2007). Because chloroplast gene regions have demonstrated a lack of sufficient phylogenetic signal to resolve relationships among fastevolving plant lineages in general (Doyle, 1992;Sang, 2002;Steele et al., 2008) and among Bromelioideae specifically (Horres et al., 2000(Horres et al., , 2007Schulte et al., 2005), several additional markers that are phylogenetically informative in various groups of angiosperms (ETS (Baldwin and Markos, 1998), rpb2 (Denton et al., 1998), and g3pdh (Olsen and Schaal, 1999)) were used along with the chloroplast intron of the trnL gene and the intergenic spacer between trnL and trnF (trnLF). ...
We developed a phylogeny of the core Bromelioideae including Aechmea and related genera, with the specific goals of investigating the monophyly of Aechmea and its allied genera, redefining monophyletic lineages for taxonomic revision, and investigating the biogeographic history of the group. Chloroplast, nuclear ribosomal, and low copy nuclear DNA sequences from 150 species within the Bromelioideae were used to develop the phylogeny. Phylogenies constructed with the combined four gene dataset provided sufficient resolution for investigating evolutionary relationships among species. Many genera are nested within Aechmea, or are rendered para- or polyphyletic by inclusion of Aechmea species. Several genera and subgenera of Aechmea with species in disjunct geographic locations are found to be polyphyletic, divided into separate clades that reflect geographic distribution rather than morphological similarity. This suggests that certain morphological characteristics thought to be indicative of common ancestry have instead evolved multiple times in parallel (i.e. ecological conservatism), possibly indicative of local adaptations to an epiphytic habit across the range of the Bromelioideae. These apparently homoplastic morphological characters used to assign species to genera or subgenera may be useful taxonomically when geography is also taken into account.
... Besides Lapanthus, other examples are Canistropsis Leme (1998: 20), Edmundoa Leme (1997: 42), and Lymania Read (1984: 201). On the other hand, the transfer of species from Canistrum Morren (1873: 257) to Wittrockia Lindman (1891: 20;Leme 1997, Leme & Luther 2010, Aechmea Ruiz & Pavon (1794: 47) to Hohenbergia Schultes & Schultes f. (1830: 1251Leme 2010b), Hohenbergia to Aechmea (Leme & Siqueira-Filho 2006b), Ronnbergia É. Morren & André (André 1874: 120) to Aechmea (Leme & Kollmann 2011), Wittrockia to Nidularium Lemaire (1854: 60;Leme 2000a), and the re-establishment of Eduandrea (Baker) Leme et al. (2008: 62;Brown & Leme 2005), demonstrate the limited reliability of the traditionally morphology based taxonomy in Bromeliaceae. ...
The authors provide a new circumscription for genera of the “Cryptanthoid complex” of Bromeliaceae subfam. Bromelioideae, originally composed of Cryptanthus, Lapanthus, Orthophytum, and Sincoraea, on the basis of new or reevaluated ecological, geographical and morphological evidence, as well as molecular phylogenies. A new generic status is proposed for Cryptanthus subg. Hoplocryptanthus and two new genera, Forzzaea, and Rokautskyia, as well as four new subgenera in Orthophytum (Capixabanthus, Clavanthus, Krenakanthus, and Orthocryptanthus) are described to render taxonomic units monophyletic. The recognized taxa are well circumscribed by the combination of geographical range, ecology and morphological characters (sex distribution, leaf succulence, sepal and petal size and connation, petal appendages, pollen and stigma morphology, fruit size, calyx persistency, seed size and number per fruit). Field collected living specimens of 78 of the 81 species of Cryptanthus s.l., all species of Lapanthus, 58 of the 59 species of Orthophytum, and all species of Sincoraea were analysed in habitat and/or in cultivation, allowing the documentation and illustration of new and underutilized characters. The molecular analysis incorporated 91 accessions representing 33 species of Cryptanthus, all species (3) of Lapanthus, 42 species of Orthophytum, and 9 species of Sincoraea, including the type species for the first three genera and four outgroup taxa. The results suggest, that some morphological characters generally considered homoplasious for Bromelioideae, for the “Cryptanthoid complex”, are not homoplasious at least within the obtained, biogeographycally well delimited clades and their taxonomical utility is redeemed.
... While in the past there have been only few changes in generic concepts for early-diverging Bromelioideae, generic delimitation in the tank-epiphytic clade, also known as the ''Core Bromelioideae" (sensu Sass and Specht, 2010), has proven highly problematic and resulted in frequent taxonomic changes over the past decades (e.g. Brown and Leme, 2005;Leme and Kollmann, 2011;Smith and Kress, 1989;Smith and Spencer, 1992). ...
The tank-epiphytic clade of berry-fruited bromeliads, also known as the Core Bromelioideae, represents a remarkable event of adaptive radiation within the Bromeliaceae; however, the details of this radiation have been difficult to study because this lineage is plagued with generic delimitation problems. In this study, we used a phylogenetic approach to investigate a well supported, albeit poorly understood, lineage nested within the Core Bromelioideae, here called the "Ronnbergia Alliance." In order to assess the monophyly and phylogenetic relationships of this group, we used three plastid and three nuclear DNA sequence markers combined with a broad sampling across three taxonomic groups and allied species of Aechmea expected to comprise the Ronnbergia Alliance. We combined the datasets to produce a well-supported and resolved phylogenetic hypothesis. Our main results indicated that the Ronnbergia Alliance was a well-supported monophyletic group, sister to the remaining Core Bromelioideae, and it was composed by species of the polyphyletic genera Aechmea, Hohenbergia and Ronnbergia. We identified two major internal lineages with high geographic structure within the Ronnbergia Alliance. The first of these lineages, called the Pacific Clade, contained species of Aechmea and Ronnbergia that occur exclusively from southern Central America to northwestern South America. The second clade, called the Atlantic Clade, contained species of Aechmea, Hohenbergia and Ronnbergia mostly limited to the Atlantic Forest and the Caribbean. We also explored the diagnostic and evolutionary importance of 13 selected characters using ancestral character reconstructions on the phylogenetic hypothesis. We found that the combination of tubular corollas apically spreading and unappendaged ovules had diagnostic value for the Ronnbergia Alliance, whereas flower size, length of the corolla tube, and petal pigmentation and apex were important characters to differentiate the Pacific and Atlantic clades. This study opens new perspectives for future taxonomic reorganizations and provides a framework for evolutionary and biogeographic studies.
... ). Mez (1896) dividió formalmente por primera vez al género en seis subgéneros y actualmente se reconocen ocho (Smith yDowns, 1979), mismos que Smith yKress (1989Kress ( , 1990Wendt, 1997;Izquierdo y Piñero, 1998;Horres et al., 2000;Faria et al., 2004;Brown y Leme, 2005;Betancur y Salinas, 2006;Horres et al., 2007;Aguirre-Santoro y Betancur, 2008;Schulte y Zizka, 2008;Faria et al., 2010;Sass y Specht, 2010) que no es un género monofilético y que lo mismo ocurre con sus relaciones infragenéricas. Sin embargo, no se cuenta hasta el momento con una clasificación alternativa a la propuesta de Smith yDowns (1979), por lo que ésta sigue siendo una referencia obligada para los distintos trabajos sobre el género. ...
Aechmea aenigmatica López-Ferrari, Espejo, Ceja et A. Mendoza, a new species from Oaxaca, Mexico is described and illustrated. The new taxon is compared with A. lueddemanniana (K. Koch) Mez and A. mexicana Baker, species with similar characteristics, from which it differs by the presence of symmetrical sepals and petals without basal appendages. Identification keys for all species of Aechmea present in Oaxaca, as well as for all species of subgenus Podaechmea are included.
... In consequence, several studies proposed taxonomic changes to reflect de-emphasis of these two characters (e.g. Read 1984, Smith & Kress 1989, Smith & Spencer 1992, Brown & Leme 2005, whereas other studies used phylogenetic approaches to discover objective phylogenetic classifications and then explore for informative morphological synapomorphies (e.g. Faria et al. 2004, Schulte et al. 2005, 2009, Sousa et al. 2007, Sass & Specht 2010, Silvestro et al. 2014, Evans et al. 2015). ...
A phylogenetic study testing the monophyly of the geographically disjunct genus Ronnbergia (Bromeliaceae, Bromelioideae) is presented. The phylogenetic analyses were based on taxon sampling that included all but one species of Ronnbergia, and representative lineages across the subfamily Bromelioideae. Three chloroplast DNA sequence markers (matK, psbA-trnH, and trnL-trnF) and morphological data were used for the phylogenetic reconstruction. Both the molecular and morphological datasets supported the polyphyly of Ronnbergia, either independently or in combination. These findings suggest that the geographic disjunction of this genus is most likely a product of taxonomic misinterpretation. The results also indicate that the species currently circumscribed in Ronnbergia are closely related to species in the genus Aechmea with similar geographic ranges. The datasets do not have enough resolution power to reconstruct a deep phylogenetic history that involves all the species of Ronnbergia. Nevertheless, this study provides clues for future approaches that should focus on a larger species sampling and the use of multi-locus DNA sequence data.
... Basiónimo:Barfuss et al. 2005). Especialmente, dentro de la subfamilia Bromelioideae la circunscripción de los géneros ha sido bastante problemática a través del tiempo (de Faria et al. 2004, Brown & Leme 2005 Aguirre-Santoro & Betancur 2006), lo que se ve confirmado por la afirmación que Smith & Downs (1979) hacen en su monografía: " ...sus géneros están tan pobremente definidos que aún con información completa es difícil asignar alguna especie a género " . De hecho, cuando se analizan detenidamente las características morfológicas que separan los géneros dentro de la subfamilia Bromelioideae se concluye que sus estados de variación, en algunos casos, son ambiguos y, en muchos otros, se sobrelapan o se comparten. ...
The inclusion of the monotypic genus Pseudaechmea into Billbergia (Bromeliaceae: Bromelioideae) is proposed and discussed. Moreover, the respective combination, illustration and morphologic redescription of Billbergia ambigua (L. B. Sm. & R. W. Read) Betancur & N. R. Salinas, species know only from Antioquia Department, Colombia, is presented.
... Numerous generic level changes (e.g. SMITH & KRESS, 1989, 1990SMITH & SPENCER, 1992;READ & BAENSCH, 1994;BROWN & LEME, 2005;BETANCUR & SALINAS, 2006) have been proposed since then, further illustrating the problematic concept of the subfamily. Although urgently needed, an updated generic concept for the subfamily is not in sight mainly due to severe uncertainties concerning the taxonomic value of morphological characters. ...
For the first time a molecular phylogeny based on five plastid markers is presented for subfamily Bromelioideae (Bromeliaceae). The species set includes 40 genera / 81 species of Bromeliaceae representing all subfamilies: Bromehoideae (29 genera / 58 species), Tillandsioideae (6 genera / 8 species) and Pitcairnioideae s.l. (5 genera / 14 species). Basal clades among the Bromehoideae are identified, nevertheless the "Core Bromelioids" comprising the majority of the species display low resolution. The phylogeny obtained makes evident, that the generic concept for Aechmea Ruiz & Pav. and its allied taxa does not describe monophyletic groups. The same holds true for several subgenera of Aechmea. The phylogeny allows the assessment ofthe systematic value of two characters that have been regarded as systematic valuable for generic delimitation in the (sub)family, 1) the petal appendages and 2) the pollen morphology. Basal Bromehoideae are characterized by sulcate pollen, while the more derived Bromehoideae display three different pollen types and several transitions between the character states.
... Generic-level systematics in Bromeliaceae is poorly understood and problematic, especially within the Bromelioideae (Wendt, 1997;Canela, Paz & Wendt, 2003;Faria, Wendt & Brown, 2004). Morphological characters traditionally used to circumscribe genera within the Bromeliaceae often fail to delimit natural groups (Brown & Terry, 1992;Grant, 1993), and the generic limits are frequently undergoing change (for example, Read, 1984;Smith & Spencer, 1992;Brown & Leme, 2005). The genus Lymania (subfamily Bromelioideae) is one example of these changes. ...
A taxonomic revision of Lymania Read is presented, including a key to the species, descriptions, conservation status, notes, and specimens examined, as well as illustrations, photographs, and distribution maps. One species, Aechmea brachycaulis, is resurrected out of synonymy under Lymania corallina, resulting in one new combination: Lymania brachycaulis (E. Morren ex Baker) L.F.Sousa, comb. nov. As recognized here, the genus Lymania comprises nine species, eight of which are endemic to southern Bahia, Brazil. The conservation status of each species is evaluated using World Conservation Union (IUCN) guidelines/criteria, and categories of threat are proposed. Five of the nine species are considered to be ‘Critically Endangered’, three ‘Endangered’, and one ‘Vulnerable’. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157, 47–66.
... Frequent changes of generic limits within Bromelioideae reflect the considerable uncertainties concerning the taxo-nomic value of morphological characters (e.g. Kress, 1989, 1990; Read and Baensch, 1994;Brown and Leme, 2005;Betancur and Salinas, 2006;de Sousa and Wendt, 2008). Furthermore, since the last comprehensive monograph of the subfamily (Smith and Downs, 1979) the number of described species has increased by more than one third (Luther, 2006), and a modern taxonomic revision is urgently needed. ...
Phylogenetic relationships within subfamily Bromelioideae (Bromeliaceae, Poales) were inferred using DNA sequence data from the low-copy nuclear gene phosphoribulokinase (PRK) and five plastid loci (matK gene, 3'trnK intron, trnL intron, trnL-trnF spacer, atpB-rbcL spacer). The PRK dataset exhibited a considerably higher proportion of potentially informative characters than the plastid dataset (16.9% vs. 3.1%), leading to a higher resolution and improved nodal support of the resulting phylogenies. Bromelia is resolved as sister to the remainder of the subfamily, albeit this relationship receives only weak nodal support. The basal position of Bromelia, as well as Deinacanthon, Greigia, Ochagavia, Fascicularia and Fernseea within the subfamily is corroborated and the remainder of the subfamily forms a highly supported clade (the eu-bromelioids). By the inclusion of nuclear data the sister group position of Fernseea to the eu-bromelioids is now highly supported. Within the eu-bromelioids the resolution of the clade representing the more advanced core bromelioids has increased and further demonstrates the highly problematic generic concept of Aechmea as well as Quesnelia. Moreover, the data were used to examine the evolution of sepal symmetry and the tank habit. Tracing of character transitions onto the molecular phylogeny implies that both characters have undergone only few transitions within the subfamily and thus are not as homoplasious as previously assumed. The character state reconstruction reveals the great importance of the evolution of the tank habit for the diversification of the core bromelioids.
The Serra do Rola-Moça State Park is located on the central southern portion of the Espinhaço Range, in a region denominated Quadrilátero Ferrífero (Iron Quadrangle). It comprises the municipalities of Belo Horizonte, Brumadinho, Ibirité and Nova Lima. The main vegetation types include the rocky grassland, semideciduous seasonal forest and savanna vegetation. This work aimed to study the floristics of the Bromeliaceae family in Serra do Rola-Moça State Park. Twenty-five species belonging to 11 genera were identified, and Dyckia is the most species-rich genus. Eduandrea selloana, Cryptanthus schwackeanus, Dyckia consimilis, D. densiflora, D. macedoi, D. simulans , D. schwackeana, D. trichostachya, Vriesea longistaminea and V. minarum are cited on the Minas Gerais Endangered Flora and Fauna Species List. Here we re-establish Dyckia oligantha as a species.
Se describe e ilustra Aechmea aenigmatica López-Ferrari, Espejo, Ceja et A.
Mendoza, una nueva especie de Oaxaca, México. El taxon aquí descrito se compara con
A. lueddemanniana (K. Koch) Mez y con A. mexicana Baker, especies con características
similares, de las cuales difiere por la presencia de sépalos simétricos y de pétalos sin
apéndices basales. Se incluyen claves para la identificación de los taxa de Aechmea presentes
en Oaxaca, así como para todos los representantes del subgénero Podaechmea.
A cladistic analysis of Lymania was conducted using morphology and sequences from three chloroplast DNA regions: the matK coding region and the psbA-trnH and trnL-trnF intergenic spacers. The monophyly of the genus and the phylogenetic relationships among related genera were examined. Of the nine Lymania species, eight are endemic to southern Bahia, Brazil. Lymania is the first genus in Bromeliaceae subfamily Bromelioideae to be subjected to a combined morphological and molecular analysis. The genera of Bromelioideae have been particularly difficult to classify and there has been disagreement about their interrelationships and monophyly. Morphological data show better resolution than molecular data alone. The partition homogeneity test supported a combined analysis of the two data sets, yielding a single most parsimonious tree. In the combined analysis, monophyly of Lymania is moderately supported, and the genus is closely related to species of Aechmea subg. Lamprococcus and subg. subg. Ortigiesia. The morphological distinctiveness coupled with low molecular divergence indicates relatively recent and rapid speciation within Lymania. The combined analysis of morphological and molecular data as done in this study provides a framework for future research on other Bromelioideae genera that could foster better taxonomic rearrangements.
The field of plant molecular systematics is expanding rapidly, and with it new and refined methods are coming into use. This paper reviews recent advances in experimental methods and data analysis, as applied to the chloroplast genome. Restriction site mapping of the chloroplast genome has been used widely, but is limited in the range of taxonomic levels to which it can be applied. The upper limits (i.e., greatest divergence) of its application are being explored by mapping of the chloroplast inverted repeat region, where rates of nucleotide substitution are low. The lower limits of divergence amenable to restriction site study are being examined using restriction enzymes with 4-base recognition sites to analyze polymerase chain reaction (PCR)-amplified portions of the chloroplast genome that evolve rapidly. The comparison of DNA sequences is the area of molecular systematics in which the greatest advances are being made. PCR and methods for direct sequencing of PCR products have resulted in a mushrooming of sequence data. In theory, any degree of divergence is amenable to comparative sequencing studies. In practice, plant systematists have focused on two slowly evolving sequences (rbcL and rRNA genes). More rapidly evolving DNA sequences, including rapidly changing chloroplast genes, chloroplast introns, and intergenic spacers, and the noncoding portions of the nuclear ribosomal RNA repeat, also are being investigated for comparative purposes. The relative advantages and disadvantages of comparative restriction site mapping and DNA sequencing are reviewed. For both methods, the analysis of resulting data requires sufficient taxon and character sampling to achieve the best possible estimate of phylogenetic relationships. Parsimony analysis is particularly sensitive to the issue of taxon sampling due to the problem of long branches attracting on a tree. However, data sets with many taxa present serious computational difficulties that may result in the inability to achieve maximum parsimony or to find all shortest trees.
Aechmea (ca. 220 species) is the largest and most diverse genus in Bromelioideae (Bromeliaceae), and several dissimilar generic concepts and infrageneric classifications have been proposed, frequently involving other closely related Bromelioideae. A morphology-based phylogenetic analysis using parsimony was conducted with 86 taxa, including 52 Aechmea (7 of the 8 recognized subgenera represented) and 34 exemplars from 9 closely related genera as the ingroup. Two species of Cryptanthus were included as the outgroup. The main objectives were to assess the validity of the major infrageneric classification systems proposed for Aechmea and to elucidate the phylogenetic position of Aechmea and putatively related genera in subfamily Bromelioideae. The topology of the consensus tree suggests that Acanthostachys, Billbergia, Portea, and non-Brazilian Ronnbergia may be monophyletic. Hohenbergia, Streptocalyx, and Quesnelia are paraphyletic or polyphyletic, as are most subgenera of Aechmea, except for subgenera Chevaliera and Macrochordion, which appear monophyletic. Characters traditionally emphasized in classifications of Bromelioideae displayed high levels of homoplasy, and this may be a reason for the artificiality of the taxonomic systems proposed for these taxa. Due to weak internal support, we refrain from recognizing any new taxonomic rearrangements. These results do provide new insights into the relationships within a number of Bromelioideae genera and suggest directions for future studies.
The field of plant molecular systematics is expanding rapidly, and with it new and refined methods are coming into use. This paper reviews recent advances in experimental methods and data analysis, as applied to the chloroplast genome. Restriction site mapping of the chloroplast genome has been used widely, but is limited in the range of taxonomic levels to which it can be applied. The upper limits (i.e., greatest divergence) of its application are being explored by mapping of the chloroplast inverted repeat region, where rates of nucleotide substitution are low. The lower limits of divergence amenable to restriction site study are being examined using restriction enzymes with 4-base recognition sites to analyze polymerase chain reaction (PCR)-amplified portions of the chloroplast genome that evolve rapidly. The comparison of DNA sequences is the area of molecular systematics in which the greatest advances are being made. PCR and methods for direct sequencing of PCR products have resulted in a mushrooming of sequence data. In theory, any degree of divergence is amenable to comparative sequencing studies. In practice, plant systematists have focused on two slowly evolving sequences (rbcL and rRNA genes). More rapidly evolving DNA sequences, including rapidly changing chloroplast genes, chloroplast introns, and intergenic spacers, and the noncoding portions of the nuclear ribosomal RNA repeat, also are being investigated for comparative purposes. The relative advantages and disadvantages of comparative restriction site mapping and DNA sequencing are reviewed. For both methods, the analysis of resulting data requires sufficient taxon and character sampling to achieve the best possible estimate of phylogenetic relationships. Parsimony analysis is particularly sensitive to the issue of taxon sampling due to the problem of long branches attracting on a tree. However, data sets with many taxa present serious computational difficulties that may result in the inability to achieve maximum parsimony or to find all shortest trees.
The Bromeliaceae were surveyed for petal appendage morphology using light and scanning electron microscopy. Three general morphological types were recognized, each loosely correlating with one of the three subfamilies. Petal appendage elaboration and variation were greatest in the Bromelioideae, while subfamilies Pitcairnioideae and Tillandsioideae exhibit more simplified, consistent morphologies. All Pitcairnioideae examined with petal appendages possessed a single appendage per petal, as opposed to two appendages per petal for the Bromelioideae and Tillandsioideae. Analyses of floral ontogenetic sequences revealed that, in all cases, petal appendage initiation is a late developmental event with petal appendages being the last formed external multicellular structures. There has been an overemphasis in the use of the presence of petal appendages in the circumscription of generic limits in the Bromeliaceae. and especially in the Tillandsioideae (e.g., Tillandsia vs. Vriesea).
Parsimony analysis of 31 sequences of the chloroplast locus ndhF was used to address questions of subfamilial phylogeny in Bromeliaceae. Results presented here are congruent with those from chloroplast DNA restriction site analysis in recognizing a clade containing Bromelioideae and Pitcairnioideae, and in resolving Tillandsioideae near the base of the family. Placements of several taxonomically difficult genera (e.g., Glomeropitcairnia and Navia) corroborate those of traditional treatments; however, these data suggest that Brocchinia (Pitcairnioideae) is the sister group to the remainder of Bromeliaceae. Further evidence for the paraphyly of Pitcairnioideae includes the resolution of Puya as the sister group to Bromelioideae. Implications for taxonomic realignment at the subfamily level are considered.
1. Introduction 2. Vegetative structure 3. Reproductive structure 4. Carbon and water balance 5. Mineral nutrition 6. Reproduction and life history with H. Luther and B. Bennett 7. Ecology 8. Relationships with fauna 9. History and evolution with G. Brown and R. Terry 10. Neoregelia subgenus Hylaeicum I. Ramirez 11. Cryptanthus I. Ramirez 12. Tillandsia and Racinaea W. Till 13. Tillandsioideae W. Till 14. Ethnobotany of Bromeliaceae B. Bennett 15. Endangered Bromeliacea, M. Dimmitt References Index.
The Bromeliaceae were surveyed for petal appendage morphology using light and scanning electron microscopy. Three general morphological types were recognized, each loosely correlating with one of the three subfamilies. Petal appendage elaboration and variation were greatest in the Bromelioideae, while subfamilies Pitcaimioideae and Tillandsioideae exhibit more simplified, consistent morphologies. All Pitcaimioideae examined with petal appendages possessed a single appendage per petal, as opposed to two appendages per petal for the Bromelioideae and Tillandsioideae. Analyses of floral ontogenetic sequences revealed that, in all cases, petal appendage initiation is a late developmental event with petal appendages being the last formed external multicellular structures. There has been an overemphasis in the use of the presence of petal appendages in the circumscription of generic limits in the Bromeliaceae, and especially in the Tillandsioideae (e.g., Tillandsia vs. Vriesea).
The recently-developed statistical method known as the "bootstrap" can be used to place confidence intervals on phylogenies. It involves resampling points from one's own data, with replacement, to create a series of bootstrap samples of the same size as the original data. Each of these is analyzed, and the variation among the resulting estimates taken to indicate the size of the error involved in making estimates from the original data. In the case of phylogenies, it is argued that the proper method of resampling is to keep all of the original species while sampling characters with replacement, under the assumption that the characters have been independently drawn by the systematist and have evolved independently. Majority-rule consensus trees can be used to construct a phylogeny showing all of the inferred monophyletic groups that occurred in a majority of the bootstrap samples. If a group shows up 95% of the time or more, the evidence for it is taken to be statistically significant. Existing computer programs can be used to analyze different bootstrap samples by using weights on the characters, the weight of a character being how many times it was drawn in bootstrap sampling. When all characters are perfectly compatible, as envisioned by Hennig, bootstrap sampling becomes unnecessary; the bootstrap method would show significant evidence for a group if it is defined by three or more characters.
Over 400 taxa in the Bromeliaceae were surveyed for stigma morphology. Five morphological categories (conduplicate-spiral, simple-erect, cupulate, convolute-blade, and coralliform) account for all known variation in bromeliad stigma morphology. The coralliform type is described and illustrated for the first time. The subfamilies Bromelioideae and Pitcairnioideae appear to be nearly homogeneous for the conduplicate-spiral stigma type. All five of the stigma types are found in the third subfamily, Tillandsioideae, where stigma variability promises to be the most useful systematically. There appear to be correlations between the simple-erect stigma type and 1) derived sexual systems (i.e., dioecism, andromonoecism, cleistogamy) within the family, and 2) xericepiphytism in the genus Tillandsia. In conjunction with other floral features, stigma morphology may be useful in refining generic and infrageneric circumscriptions.
The recently-developed statistical method known as the "bootstrap" can be used to place confidence intervals on phylogenies. It involves resampling points from one's own data, with replacement, to create a series of bootstrap samples of the same size as the original data. Each of these is analyzed, and the variation among the resulting estimates taken to indicate the size of the error involved in making estimates from the original data, In the case of phylogenies, it is argued that the proper method of resampling is to keep all of the original species while sampling characters with replacement, under the assumption that the characters have been independently drawn by the systematist and have evolved independently. Majority-rule consensus trees can be used to construct a phylogeny showing all of the inferred monophyletic groups that occurred in a majority of the bootstrap samples. If a group shows up 95% of the time or more, the evidence for it is taken to be statistically significant. Existing computer programs can be used to analyze different bootstrap samples by using weights on the characters, the weight of a character being how many times it was drawn in bootstrap sampling. When all characters are perfectly compatible, as envisioned by Hennig, bootstrap sampling becomes unnecessary; the bootstrap method would show significant evidence for a group if it is defined by three or more characters.
Three distinct types of stigma architecture, designated as “simple-erect,” “conduplicate-spiral”, and “convolute-blade” were
identified for Bromeliaceae. Structure of the conduplicate-spiral and convolute-blade stigma types is highly complex and previously
unreported. Stigma morphology, largely neglected in Tillandsioideae until recently, promises to yield new characters valuable
in interpreting systematic relationships in this subfamily. Data indicate that the putatively plesiomorphic state (simple-erect)
is retained in some members of all three subfamilies. We found convolute-blade-type stigmas only in Tillandsioideae, though
not in all members of the subfamily. Our sample indicates that Tillandsioideae stigmas are more like those of Pitcairnioideae
than of Bromelioideae, which is consistent with Smith’s hypothesis of phylogenetic proximity of Tillandsioideae and Pitcairnioideae.
Stigma architecture is proving to be a rich source of data for phylogenetic reconstruction and for classification at several
infrafamilial levels.
- Baker
Dispersão de sementes em bromélias de frutos carnosos: interpretações incorretas e orientação para estudos mais aprofundados
- Benzing D. H.
Contribuiçâo ao estudo do gênero Nidularium (Bromeliaceae) — Parte I — subgênero Canistropsis
- Pereira E.
Cladistic relationships of Aechmea (Bromeliaceae) and allied genera
- Faria A. P. G.