INCREASED WATER STORAGE CAPACITY IN CACTUS WOOD: A STUDY IN THE TRIBE CEREEAE (CACTOIDEAE, CACTACEAE)

Secondary xylem of fibrous cactus wood is characterized by short narrow vessel elements with both simple perforation plates and large intervessel pits, libriform septate fibers, and large rays. These are present in basal cactus taxa, as well as in many other groups of the family. In Cactoideae, the most diversified and most derived subfamily, there are remarkable variations found in the secondary xylem, with the more highly derived taxa containing the greatest water storage capacity. Unlignified parenchyma is one specialization found in the fibrous wood of cacti. We observed this tissue in the secondary xylem at the base of the sterns of several Brazilian endemic species of Arrojadoa, Melocactus, and Stephanocereus, all members of the tribe Cereeae. In Arrojadoa and Melocactus the unlignified parenchyma occurs in lines and bands amongst the axial and radial xylem elements, while in Stephanocereus it is mainly restricted to the rays and does not form bands. We address the adaptive importance of the unlignified parenchyma in the fibrous wood in tribe Cereeae and the family Cactaceae as a whole.

A macro- and micromorphological survey of floral and extrafloral nectaries in the epiphytic cactus Rhipsalis teres (Cactoideae: Rhipsalideae)

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)

A New Subgeneric Classification of Rhipsalis (Cactoideae, Cactaceae)

Most Cactaceae have succulent stems and inhabit dry or arid areas, but some are epiphytes of humid regions. Rhipsalis is the largest genus of epiphytic cacti. Species of Rhipsalis are notoriously difficult to identify, and the subgeneric classification of the genus has remained controversial. Between 1837 and 1995, eight different subgeneric classifications have been proposed for Rhipsalis. The most comprehensive taxonomic treatment of the genus recognized five subgenera, Phyllarthrorhipsalis, Rhipsalis, Epallagogonium, Calamorhipsalis, and Erythrorhipsalis, characterized mainly by stem morphology. Here, molecular phylogenetic information combined with morphological data is used to re-evaluate the former subgeneric classifications proposed for the genus. Three monophyletic subgenera are recognized, Rhipsalis, Calamorhipsalis and Erythrorhipsalis, which are mainly characterized by floral traits. The changes proposed include expanding the circumscription of Rhipsalis by the inclusion of species previously included in Phyllarthrorhipsalis and Epallagogoniwn and recognizing a broader Calamorhipsalis, also including species from subgenus Epallagogonium. The circumscription of Erythrorhipsalis remains unchanged. For each subgenus a list of synonyms, a brief description and a list of species included are presented. A key for the identification of subgenera is also provided.

DNA barcodes for Mexican Cactaceae, plants under pressure from wild collecting

DNA barcodes could be a useful tool for plant conservation. Of particular importance is the ability to identify unknown plant material, such as from customs seizures of illegally collected specimens. Mexican cacti are an example of a threatened group, under pressure because of wild collection for the xeriscaping trade and private collectors. Mexican cacti also provide a taxonomically and geographically coherent group with which to test DNA barcodes. Here, we sample the matK barcode for 528 species of Cactaceae including approximately 75% of Mexican species and test the utility of the matK region for species-level identification. We find that the matK DNA barcode can be used to identify uniquely 77% of species sampled, and 79-87% of species of particular conservation importance. However, this is far below the desired rate of 95% and there are significant issues for PCR amplification because of the variability of primer sites. Additionally, we test the nuclear ITS regions for the cactus subfamily Opuntioideae and for the genus Ariocarpus (subfamily Cactoideae). We observed higher rates of variation for ITS (86% unique for Opuntioideae sampled) but a much lower PCR success, encountering significant intra-individual polymorphism in Ariocarpus precluding the use of this marker in this taxon. We conclude that the matK region should provide useful information as a DNA barcode for Cactaceae if the problems with primers can be addressed, but matK alone is not sufficiently variable to achieve species-level identification. Additional complementary regions should be investigated as ITS is shown to be unsuitable

Molecular systematics of the Cactaceae

Bayesian, maximum-likelihood, and maximum-parsimony phylogenies, constructed using nucleotide sequences from the plastid gene region trnK-matK, are employed to investigate relationships within the Cactaceae. These phylogenies sample 666 plants representing 532 of the 1438 species recognized in the family. All four subfamilies, all nine tribes, and 69% of currently recognized genera of Cactaceae are sampled. We found strong support for three of the four currently recognized subfamilies, although relationships between subfamilies were not well defined. Major clades recovered within the largest subfamilies, Opuntioideae and Cactoideae, are reviewed; only three of the nine currently accepted tribes delimited within these subfamilies, the Cacteae, Rhipsalideae, and Opuntieae, are monophyletic, although the Opuntieae were recovered in only the Bayesian and maximum-likelihood analyses, not in the maximum-parsimony analysis, and more data are needed to reveal the status of the Cylindropuntieae, which may yet be monophyletic. Of the 42 genera with more than one exemplar in our study, only 17 were monophyletic; 14 of these genera were from subfamily Cactoideae and three from subfamily Opuntioideae. We present a synopsis of the status of the currently recognized genera

Molecular phylogeny of tribe Rhipsalideae (Cactaceae) and taxonomic implications for Schlumbergera and Hatiora

Tribe Rhipsalideae is composed of unusual epiphytic or lithophytic cacti that inhabit humid tropical and subtropical forests. Members of this tribe present a reduced vegetative body, a specialized adventitious root system, usually spineless areoles and flowers and fruits reduced in size. Despite the debate surrounding the classification of Rhipsalideae, no studies have ever attempted to reconstruct phylogenetic relationships among its members or to test the monophyly of its genera using DNA sequence data; all classifications formerly proposed for this tribe have only employed morphological data. In this study, we reconstruct the phylogeny of Rhipsalideae using plastid (trnQ-rps16, rpl32-trnL, psbA-trnH) and nuclear (ITS) markers to evaluate the classifications previously proposed for the group. We also examine morphological features traditionally used to delimit genera within Rhipsalideae in light of the resulting phylogenetic trees. In total new sequences for 35 species of Rhipsalideae were produced (out of 55: 63%). The molecular phylogeny obtained comprises four main clades supporting the recognition of genera Lepismium, Rhipsalis, Hatiora and Schlumbergera. The evidence gathered indicate that a broader genus Schlumbergera, including Hatiora subg. Rhipsalidopsis, should be recognized. Consistent morphological characters rather than homoplastic features are used in order to establish a more coherent and practical classification for the group. Nomenclatural changes and a key for the identification of the genera currently included in Rhipsalideae are provided. (C) 2011 Elsevier Inc. All rights reserved.

Wide-band tracheids (WBTs) of photosynthetic and non-photosynthetic stems in species of Cactaceae

ARRUDA, E. AND G. F. A. MELO-DE-PINNA (Departamento de Botanica, Instituto de Biociencias. Universidade de Sao Paulo, Rua do Matao, travessa 14, Cidade Universitaria, Butanta, Caixa Postal 11461, 05422-970. Sao Paulo, SP, Brasil). Wide-band tracheids (WBTs) of the photosynthetic and non-photosynthetic stems in species of Cactaceae. J. Torrey Bat. Soc. 137: 16-29. 2010.-The absence of WBTs and wood polymorphisms in some species of the Caryophyllales may be related to the particular area of plant analyzed. The present research has the objective of studying the photosynthetic and non-photosynthetic stems of different species and stages of differentiation to register wood polymorphisms and to understand the distribution and occurrence of WBTs. Wood polymorphism was observed in the non-photosynthetic stern of young and adult plants of Opuntioideae and Cactoideae and is also found in the photosynthetic stem of young plants of some species of Cactoideae. Cactoideae present WBT/fibrous dimorphic wood that can be related to cambial variation associated with growth habits and plant development. As expected, in the photosynthetic stem of the adult columnar cacti the wood is monomorphic fibrous in which WBTs were not found. This wood contains a great amount of fibers due to necessity of the mechanical support. In contrast, the globular species do not possess fibers in this area of the stem in either adult or young plants. Opuntia monacantha Haw. had non-fibrous wood in which WBTs were observed in the axial system and in the inner parts of the rays. Fiber clusters were present in the axial system. This wood represents a variation in the wood types described for Opuntioideae. Also, in O. monacantha, cells similar to the WBTs were observed in the pith, which can be interpreted as variation in the morphogenic processes during the ontogeny of the plant, probably a case of homeosis. Monomorphic fibrous wood without WBTs was found along the entire stem of Pereskia bahiensis Gurke. This feature has been observed in other pereskias, and in addition to the others, indicates its proximity to the ancestral cacti.

The systematic significance of floral morphology, nectaries, and nectar concentration in epiphytic cacti of tribes Hylocereeae and Rhipsalideae (Cactaceae)

A long-standing interest in cactus taxonomy has existed since the Linnaean generation, but an appreciation of the reproductive biology of cacti started early in the 1900s. Numerous studies indicate that plant reproductive traits provide valuable systematic information. Despite the extensive reproductive versatility and specializations in breeding systems coupled with the striking floral shapes, the reproductive biology of the Cactaceae has been investigated in approximately 10% of its species. Hence, the systematic value of architectural design and organization of internal floral parts has remained virtually unexplored in the family. This study represents the most extensive survey of flower and nectary morphology in the Cactaceae focusing on tribes Hylocereeae and Rhipsalideae (subfamily Cactoideae). Our objectives were (1) to conduct comparative morphological analyses of flowers and floral nectaries and (2) to compare nectar solute concentration in these two tribes consisting of holo- and semi-epiphytic species. Flower morphology, nectary types, and sugar concentration of nectar have strong taxonomic implications at the tribal, generic and specific levels. Foremost, three types of nectaries were found, namely chamber nectary (with the open and diffuse subtypes), furrow nectary (including the holder nectary subtype), and annular nectary. All Hylocereeae species possess chamber nectaries, in which the nectarial tissue has both trichomes and stomata. The Rhipsalideae are distinguished by two kinds of floral nectaries: furrow and annular, both nectary types with stomata only. The annular nectary type characterizes the genus Rhipsalis. Nectar concentration is another significant taxonomic indicator separating the Hylocereeae and Rhipsalideae and establishing trends linked to nectar sugar concentration and amount of nectar production in relation to flower size. There is an inverse relationship between flower size and amount of nectar production in the smaller Rhipsalideae flowers, in which nectar concentration is more than two-fold higher despite the smaller volume of nectar produced when compared to the large Hylocereeae flowers. Variability of nectary morphology and nectar concentration was also evaluated as potential synapomorphic characters in recent phylogenies of these tribes. In conclusion, our data provide strong evidence of the systematic value of floral nectaries and nectar sugar concentration in the Cactaceae, particularly at different taxonomic levels in the Hylocereeae and Rhipsalideae. © 2013 Perspectives in Plant Ecology, Evolution and Systematics.