FIGURE 9 - uploaded by Rafael Batista Louzada
Content may be subject to copyright.
A-F. Orthophytum navioides (L.B.Sm.) L.B.Sm.-A. Habit.-B. Section from the base of flower showing the epigenous tube.-C. Sepal and free stamen.-D. Sepal abaxial surface.-E. Petal showing an adnate stamen, petals appendages and the petals collosities.-F. Style and stigma (Jardim et al. 729).
Source publication
Orthophytum is a bromeliad genus restricted to Brazil with records for the states of Paraíba, Pernambuco, Alagoas, Sergipe, Bahia, Espírito Santo and Minas Gerais. The genus is usually divided in two informal groups based on the presence or absence of a peduncle. This paper presents a taxonomic revision of the 12 species in the group with sessile i...
Similar publications
p> Background : Bromeliaceae family in Mexico has been the object of interest by botanists since 1789; their systematic study was approached from the 1970s onwards, and now there are significant advances in its taxonomic-floristic knowledge.
Question: How many and which species of Bromeliaceae occur in Mexico? How they are distributed, and how ma...
Citations
... & A.A. Conc.) LOUZADA & WAND, is a vulnerable bromeliad, limited solely to the Chapada Diamantina of the Mucugê municipality, Brazil (LOUZADA & WANDERLEY, 2010). The few number of plants acquired via natural propagation related to the predatory collection of the species are the reason for the development of research with the goal of propagating and conserving this genetic resource that has great potential for the ornamental plant market. ...
Sincoraea mucugensis (Wand. & A.A. Conc.) LOUZADA & WAND, an endangered bromeliad, is confined to the central region of the Chapada Diamantina, in the municipality of Mucugê, Brazil. From various researches, it is evident that for the propagation of this species, the in vitro technique is a feasible option. However, due to the low multiplication rates reported in various papers, this study aimed to establish a micropropagation protocol of direct organogenesis for S. mucugensis. First, the inoculation of the stem explants was done in MS ½ culture medium which contained different levels of BAP (0.00; 6.66; 8.88; 11.10; 13.20 µM) and NAA (0.00; 2.60; 5.20 µM). These shoots were then subjected to a couple of distinct rooting periods (of 30- and 60-day duration) using activated charcoal; finally, these microplants were transferred to a greenhouse for acclimatization, and covered with transparent plastic cups, as a water loss prevention test method. All the data were submitted to the analysis of variance (ANOVA), and the means were subjected to regression analysis or compared using the Tukey test. The findings revealed that the S. mucugensis stem explants raised in the NAA-rich medium (6.42 to 7.43 shoots/explants) showed high multiplication rates; the shoot rooting was done for 30 days using activated charcoal with the medium. Acclimatization, which was performed by directly exposing the microplants to the ex vitro environment, showed 95% survival rate.
... Orthophytum has 58 species, all endemic to the Eastern Brazil, from RN to Minas Gerais. This genus has notably two centers of diversity: the rocky outcrops of the Espinhaço Range (BA and MG) and inselbergs in the Atlantic Forest of MG and ES (Wanderley 1990, Louzada & Wanderley 2010, Gouda & Butcher 2018, Flora do Brasil 2020. Based on morphologic and molecular phylogenic studies, current classification supports the subdivision of Orthophytum in five subgenera (Leme et al. 2017), as well as the segregation of several species to the re-established genus Sincoraea (Louzada & Wanderley 2016). ...
... Comments:-Orthophytum disjunctum was part of the traditional "pedunculate inflorescence group" (Louzada & Wanderley 2010, Louzada et al. 2014 and is now placed under Orthophytum subg. Orthophytum (Leme et al. 2017). ...
The Bromeliaceae Flora for the state of Rio Grande do Norte, Northeastern Brazil, is presented, based on extensive fieldwork, morphological analyses using herbarium and freshly collected material, and specialized literature. Twenty-six species of bromeliads were recorded in Rio Grande do Norte, distributed in ten genera and in three subfamilies. Bromelioideae was the richest subfamily (eight genera/14 species), followed by Tillandsioideae (one genus/12 species), and Pitcairnioideae (one genus/one species). Aechmea mertensii, Hohenbergia horrida and Tillandsia tenuifolia are new records for Rio Grande do Norte. Eight species (31%) are restricted to the Eastern portion of the state, in the Atlantic Forest. Caatinga dry woodlands harbor 18 species, with remarkable presence of Bromelia laciniosa, Encholirium spectabile, Tillandsia recurvata and T. streptocarpa, the four most widely distributed taxa. We discuss problems related to unclear taxonomic circumscriptions of species or diverging information between authors, more expressively in Hohenbergia, but also in Aechmea, Cryptanthus and Tillandsia. The data presented here might contribute to better understand the morphological variation of these taxa and suggest additional research on their taxonomy. Morphological descriptions, general comments, a map, photo plates and an identification key for all taxa are provided.
... In total, there are 58 genera and 3352 species of Bromeliaceae (Luther, 2012), 132 and 911 species of which occur in the Caatinga domain and in the Atlantic Forest domain, respectively . The role of the sugar loafs in the Atlantic Forest is prominent; they seem to be important for the speciation of certain bromeliad genera, especially those forming mats (Leme, 2004;Versieux & Wendt, 2006;Louzada & Wanderley, 2010;Versieux & Wanderley, 2015). The high beta-diversity and microendemism in Bromeliaceae on inselbergs is outstanding (Benzing, 2000;Martinelli et al., 2008) and is usually attributed to the existence of phytogeographical links to other rock outcrop types and a high degree of demarcation against the surrounding matrix (Porembski, 2007;Saraiva, Mantovani & Forzza, 2015). ...
Inselbergs sensu lato are isolated, mainly granitic and gneissic, rock outcrops, forming old landscape elements on crystalline continental shields on all continents. Mats consisting mainly of monocots occur on open rocky slopes, forming a conspicuous habitat on these outcrops. Brazilian lowland inselbergs consist of species-rich mats with large numbers of endemics and Bromeliaceae standing out as characteristic elements in these communities. For the first time, we provide a checklist of mat-forming bromeliads on lowland inselbergs in Brazil and investigate whether the regional species pool influences the species diversity of inselberg mats. Furthermore, through modelling analysis, we looked for the potential distribution of bromeliads endemic to inselbergs, expecting that they would show a narrower niche width than non-endemic species. Our data indicate the existence of a particular phytogeographical region in the Atlantic Forest in south-eastern Brazil, comprising inselbergs harbouring a highly diverse flora of Bromeliaceae, which we call Sugar Loaf Land. In addition, our projections reveal smaller predicted areas for bromeliads endemic to inselbergs compared with non-endemic species, the former apparently being mainly controlled by climatic factors. In addition to climate and regional species pools, the island-like character of inselbergs seems to play an important role in the speciation of rupicolous Bromeliaceae.
... Recent revisions of exclusively Brazilian genera of Bromelioideae have been carried out: Canistrum, Canistropsis, Cryptanthus, Edmundoa, Eduandrea, Nidularium, Lymania, Portea, Lapanthus, Orthophytum and Wittrockia (Coffani-Nunes 2004;Leme 1997, Louzada 2012Louzada & Wanderley 2010;Ramirez 1996Ramirez , 2000Sousa & Wendt 2008). Species-groups within Aechmea have also been revised (Abondanza 2012;Faria et al. 2010;Siqueira Filho & Leme 2006;Sousa 2004). ...
Poales represents a major part of Angiosperm and Monocot diversity. The families encompass ca. 20,000 species which is about 7% of the Angiosperms and 33% of the Monocots. Bromeliaceae, Cyperaceae, Erio-caulaceae, Juncaceae, Mayacaceae, Poaceae, Rapateaceae, Thurniaceae, Typhaceae, and Xyridaceae are the families represented in the Neotropics. In general terms some areas in the Neotropics could be considered hotspots for Poales with a high number of species in several genera and several centers of endemism – the Guayana Shield, Espinhaço Range and Atlantic Forest are highly diverse in Poales. In terms of the Brazilian flora, the order is well represented in the entire country with almost 4,400 species. It represents more than 50% of the total number of species of Monocotyledons in Brazil. The main goal here is to summarize the available information and provide an overview of the Poales in the Neotropics. The state of knowledge for each family, focused on the Brazilian flora, is provided and reinforces the importance of new studies in key-groups looking beyond the understanding of their diversity on the continent but also the conservation of the species.
... Orthophytum has been divided into two informal groups based on the inflorescence being either scapose or sessile (Leme 2004;Louzada and Wanderley 2010;Louzada and Versieux 2010 Sass and Specht (2010), Orthphytum supthutii, now Lapanthus duartei, has been included in all of the Bromelioideae-focused molecular studies. However, in Schulte et al. (2005), Horres et al. (2007), and Schulte and Zizka (2008), this was the only included Orthophytum. ...
Abstract—
Of the eight subfamilies currently recognized in Bromeliaceae, Bromelioideae is perhaps the most poorly understood. Generic circumscriptions are unclear, and an exceptionally diverse morphology coupled with an unusually low rate of sequence divergence within
Bromeliaceae has made it difficult to resolve phylogenetic relationships within the subfamily. Although recent molecular studies have begun elucidating relationships among species in Bromeliaceae, most have not sampled deeply and/or broadly across Bromelioideae. The purpose of this study was
to conduct a phylogenetic analysis within subfamily Bromelioideae using three chloroplast DNA regions (matK, psbA-trnH, and trnL-trnF), with the inclusion of multiple species from a broad sampling of bromelioid genera. Ochagavia, Deinacanthon, Fascicularia,
Bromelia and Fernseea diverged relatively early in the history of the subfamily, with the remaining taxa being placed in a large and poorly resolved eubromelioid clade. Bromelia and Cryptanthus were found to be monophyletic, while 13 other genera were polyphyletic.
Aechmea, the most morphologically diverse genus within the subfamily, was highly polyphyletic, with species distributed among 12 different lineages, with little support for subgeneric circumscriptions.
... Currently, there are records for the states of Ceará, Paraíba, Pernambuco, Alagoas, Sergipe, Bahia, Minas Gerais, Espírito Santo (Forzza et al. 2012). All species are heliophytes and the majority grows in rocky environments, such as outcrops or inselbergs (Louzada and Wanderley 2010). The diversity centers of Orthophytum are the states of the Bahia and Minas Gerais, and found growing along the Espinhaço mountain range's rocky fields, and on inselbergs from northeastern Minas Gerais to central-north Espírito Santo The second new occurrence belongs to Tillandsia, which is the largest genus within Bromeliaceae, with 627 known species (Luther 2010). ...
In the present work we describe the first occurrence of the genus Orthophytum (Bromeliaceae, Bromelioideae) and of the species Tillandsia paraibensis within the state of Rio Grande do Norte, northeastern Brazil. Both taxa were recorded on inselbergs in areas of caatinga. These findings are important, improving the knowledge of the Flora of Rio Grande do Norte and giving a better understand of the biogeography of Brazilian bromeliads.
... There are two centers of diversity for the genus: one in the Espinhaço Range, and the other one in the Atlantic Rainforest areas (Louzada & Wanderley 2010). The Orthophytum species are generally saxicolous, but few species can be terrestrial. ...
... Two morphological groups can be distinguished: the sessile inflorescence group and the pedunculate inflorescence group (Wanderley & Conceição 2006, Louzada & Wanderley 2010, although both are clearly not monophyletic groups (Louzada et al. in prep). Based on morphological studies of dry and living material in collections and field studies, we describe a new species with a pedunculate inflorescence, from campos rupestres of Chapada Diamantina, Bahia, morphologically similar to Orthophytum toscanoi Leme (2003: 23 Distribution and habitat:-Orthophytum argentum is known from Rio de Contas municipality in Bahia. ...
In this paper we describe and illustrate Orthophytum argentum as a new species from campos rupestres of Espinhaco Range, morphologically similar to Orthophytum toscanoi.
A phylogenomic analysis of the so far phylogenetically unresolved subfamily Bromelioideae (Bromeliaceae) was performed to infer species relationships as the basis for future taxonomic treatment, stabilization of generic concept, and further analyses of evolution and biogeography of the subfamily. A target-enrichment approach was chosen, using the Angiosperms353 v.4 kit RNA-baits and including 86 Bromelioideae species representing previously identified major evolutionary lineages. Phylogenetic analyses were based on 125 target nuclear loci, assembled off-target plastome as well as mitogenome reads. A Bromelioideae phylogeny with a mostly well-resolved backbone is provided based on nuclear (194 kbp), plastome (109 kbp), and mitogenome data (34 kbp). For the nuclear markers, a coalescent-based analysis of single-locus gene trees was performed as well as a supermatrix analysis of concatenated gene alignments. Nuclear and plastome datasets provide well-resolved trees, which showed only minor topological in-congruences. The mitogenome tree is not sufficiently resolved. A total of 26 well-supported clades were identified. The genera Aechmea, Canistrum, Hohenbergia, Neoregelia, and Quesnelia were revealed polyphyletic. In core Bromelioideae, Acanthostachys is sister to the remainder. Among the 26 recognized clades, 12 correspond with currently employed taxonomic concepts. Hence, the presented phylogenetic framework will serve as an important basis for future taxonomic revisions as well as to better understand the evolutionary drivers and processes in this exciting subfamily.
Phylogenetic trouble unleashed
The first part of my thesis deals with a comprehensive phylogeny of the Bromelioideae subfamily. The family Bromeliaceae is subdivided into eight subfamilies, one of them is the Bromelioideae. Phylogenetic relationships among the Bromelioideae are still poorly understood and many of the extant genera are suspected to be not monophyletic. Especially Aechmea, the largest and most polymorphic genus constitutes many questions and the genus was used as a depot for taxonomically problematic species. The phylogenetic study presented here is the most comprehensive one so far, covering about half of the known species (434 of 965, Table 1) of Bromelioideae. The phylogeny was generated using plastid (atpB-rbcL, matK, rps16, ycf1_1, ycf1_6) and nuclear (AGT1_exon, ETS, G3PDH, PHYC, RPB2) genetic markers. The markers were analysed individually as well as combined using maximum likelihood and Bayesian analysis. The comparison of plastid vs. nuclear data revealed significant differences which were discussed in detail and hypothesised to indicate hybridisation in certain lineages. Nevertheless, the combination of both datasets increased the overall resolution of the phylogeny and was used to discuss the results in the light of previous studies. The entire phylogeny was divided into 32 groups for discussion. These groups represent potential genera or starting points for further studies in order to reorganise the polyphyletic genera of Bromelioideae into monophyletic lineages. Many extant genera of the eu-Bromelioideae were found to be not monophyletic. Monophyly was observed for the genera Acanthostachys, Billbergia, Cryptanthus, Disteganthus, Hoplocrypanthus, Lapanthus, Orthocryptanthus, Orthophytum, Rokautskyia, Ronnbergia, Sincoraea, Wittmackia and the monotypic ones (Deinacanthon, Eduandrea, Fascicularia, Hohenbergiopsis, Pseudananas). The genus concept proposed by Smith and Downs (1979) is therefore rejected, as well as the taxonomic utility of petal appendages, which were mainly used to delimit genera. In summary, this study and recent studies highlighted other morphological characters (e.g. pollen morphology, stigma type) as much more informative. However, no single character should be used to delimit genera and combinations of relevant characters are required. Even the petal appendages can pose a taxonomical important character at certain taxonomic level.
The combination of biogeography and phylogeny revealed that species of some groups which co- occur in a biome or region are also phylogenetically closely related. These groups were not recognised before because the misinterpretation of homoplastic characters led to wrong taxonomical conclusion. For example, the recent re-organisation of the Cryptanthoid group and the re-establishment of Wittmackia with the former Hohenbergia subgen. Wittmackiopsis species highlighted, among other characters, the importance of biogeography. Another case is the subgenus Neoregelia subgen. Hylaeaicum which is geographically and phylogenetically separated from the Nidularioid group and therefore has to be excluded.
5
The large phylogeny presented here gives evidence for multiple invasions of the Brazilian biomes (Amazon Forest, Atlantic Forest, Cerrado, Caatinga) as well as of Central America and the Greater Antilles. It is important to note that the phylogeny is lacking resolution in the deeper nodes. Confident assumptions are therefore hindered and the historical biogeography of Bromelioideae remains cryptic. Anyway, the Atlantic Forest is nowadays the diversity hotspot of the core Bromelioideae and critically endangered. Extensive conservation efforts are required to protect the diverse flora, including the bromeliads.
The genetic markers used so far in bromeliad phylogenies provided only limited variation resulting in often unresolved complexes. The search for additional suitable genetic markers in bromelioid phylogenies yielded the nuclear marker AGT1. The amplified fragment consists of one well conserved exon region as well as a highly variable intron. The intron was too variable for aligning it across the entire bromelioid set. On the other hand, the intron provides relevant information for inferring phylogenies of closely related species groups (e.g. in Ananas, Cryptanthoid group). Furthermore, AGT1 is proposed as a genetic barcode in Bromelioideae because it poses much more information then the commonly used ones (e.g. matK).
Does size matter?
The second part of this thesis deals with the genome size evolution within the family Bromeliaceae. Samples from seven subfamilies were screened with the emphasis on the subfamily Bromelioideae. The data were combined with data from literature and the observed patterns were discussed in relation to known phenomena (e.g. correlations to environment and life form). In the second sub-chapter I have chosen the species Tillandsia usneoides to study the intraspecific genome size variation in combination with morphology and biogeography. Genome size and base composition were measured using the flow cytometry technique.
Bromeliaceae comprises mostly diploid species with predominantly 50 small chromosomes (2n), small genome sizes (0.59-4.11 pg) and normal GC content (36.46-42.21 %) compared to other families. Polyploidy was observed so far in the subfamilies Bromelioideae, Tillandsioideae and Pitcairnioideae. Triploids, tetraploids and potential hexaploids were identified. The genera show significant differences in holoploid genome size and base composition throughout the entire family. GC content is weakly positively correlated with genome size. Significant intraspecific genome size variation has been observed, including polyploidization, but no endopolyploidy and no variation in dioecious species. Within the subfamily Bromelioideae, the observed genome size between the early diverging lineages and the core Bromelioideae supports this division. The differences are due to a higher proportion of polyploids in the early diverging lineages and a significant higher
6
GC content in the core Bromelioideae. Both groups differ in their life strategies and occupy principally different habitats with corresponding morphological adaptations. Hence, the early diverging lineages are predominantly terrestrial and xeromorphic. In contrast, the prevailing epiphytic core Bromelioideae are characterised by a tank habit and mostly adapted to more humid environments. Across the family and the subfamily Bromelioideae in particular, significant genome size differences between the different life forms have been observed, but no correlation to biomes within Brazil.
Tillandsia usneoides is the most widely distributed species of the family Bromeliaceae. It ranges from the southeastern United States to Argentina and Chile. Tillandsia usneoides grows epiphytic and is dispersed by seeds as well as by fragments of the plant. Within the species striking morphological differences can be observed as far as size characters are concerned. Morphotypes have shown to be stable in cultivation while growing under the same conditions. In order to investigate possible reasons for the variation the relative genome size of 75 specimens covering the whole distribution range was measured and combined with morphological, distribution and climatic data. Significant variation in the relative genome size corresponded to the morphological differences and reflected the north-south distribution gradient. Genome size and morphotypes showed a positive correlation, as well as with the mean temperature of the driest and coldest quarter and the minimal temperature of the coldest month.
