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Sobre la Posición sistemática de Brunellia Ruiz & Pavon
Abstract
Resumen Bajo el concepto de que Brunellia es un táxon de Cunoniaceae y no una familia monogenérica Brunelliaceae, el presente trabajo examina la posición sistemática de Brunellia analizando las relaciones filogenéticas de este taxón con once taxones que representan a Cunoniaceae, con 20 caracteres morfológicos. Un total de doce taxones hacen parte del grupo interno. El programa Hennig86 basado en el método de simplicidad (parsimony) fue usado para la elección de las hipótesis de relaciones. Los datos del gru-po interno fueron analizados primero con cuatro grupos externos: Connarus, Fothergilla, Davidsonia y Quercus,-con tres excluyendo a Quercusy con cada uno de ellos por sepa-rado; para este último caso los caracteres fueron polarizados. Los resultados muestran a Brunellia como taxón diferente de Cunoniaceae y a Cunoniaceae como grupo no monofilético. Se propone en este trabajo la inclusión de Brunellia, Spiraeanthemum-Acsmithia bajo una familia diferente de Cunoniaceae. Mayor investigación es necesaria para corroborar la hipótesis de relaciones de estos taxones. Se cuestiona también la monofilia de Cunoniaceae por la presencia de Davidsonia dentro del grupo interno. Se confirma la relación estrecha entre las Rosidae basales con las Hamameliidae inferio-res. La apomortla y pleslomorfta de algunos caracteres es discutida. La monofilia de nin-guna de las tribus de Engler es confirmada. Abstract Under the concept that Brunellia belongs to the Cunoniaceae rather thant being separated in Brunelliaceae, this paper examines the systematic position of Brunellia by studying the phylogentic relationships of this taxon with eleven ingrow taxa representing Cunoniaceae and using twenty characters. A total of twelve taxa represent the ingroup. The Hennig86 program based on parsimony was used to choose the hypotheses about relationships. The data of the internal group was analyzed first with four outgroups: Connarus, Davidsonia, Fothergil/a y Quercus, then excluding Quercus and last with each of the outgroups independently. Data were polarized for the last analysis. Results showed that Brunellia does not belong to Cunoniaceae and Cunoniaceae is not a monophyletic group. I propose that Brunellia, and Spiraeanthemun-Acsmithia under a different family from Cunoniaceae. More research is necessary to corroborate the relationships of these taxa. The monophylyot the Cunoniaceae is also doubtful because some of its genera are nested with Davidsonia. The results also confirmed the relationships between the Rosidae and the lower Hamameliidae. Discussion about plesiomorphic and apomorphic states conditions are given for the Rosi~ae. The monophyly of none of Engler's tribes is confirmed.
... Con el fin de polarizar los estados de los caracteres, se efectuó una comparación mediante el uso de grupos externos (Humphries & Funk 1984, Maddison et al. 1984 ). Este tipo de comparaciones permite reconocer los estados plesiomórficos y apomórficos (González 1997, Orozco 1997). Según González (1997) y Weller & Sakai (1999, la inclusión de diferentes grupos externos permite considerar una mayor variación de las características contempladas y a la vez, aporta elementos de juicio reales (taxones) en lugar de grupos hipotéticos; esto último garantiza que los resultados puedan estar sujetos a prueba (Ax 1987, citado por González 1997). ...
... Una razón adicional en este caso consiste en el pobre conocimiento disponible de la filogenia del género Axonopus, por lo que se procedió a considerar varios grupos externos en un intento por aportar un mayor conocimiento del género en todas sus divisiones, seleccionando para ello representantes que abarcaran la máxima variabilidad morfológica, así como cubrir el área de distribución geográfica del género en el continente americano. Además, la consideración de dos o más grupos externos reduce la posibilidad de que éstos formen parte del grupo interno (González 1997, Orozco 1997). Como grupos externos se consideraron varias especies de todas las categorías infragenéricas propuestas para el género. ...
... In Brunelliaceae tanniferous tissue is absent in flowers (except for the unicellular hairs). Brunellia was sometimes included in Cunoniaceae but appears now as a separate family (Orozco, 1997;Savolainen et al., 2000). In Cephalotaceae, flowers exhibit several probable autapomorphies. ...
Floral morphology, anatomy and histology were studied in representatives of all families of current Oxalidales, which were recently constituted as a result of molecular systematic studies by other authors, and are composed of families of different positions in traditional classifications (Oxalidaceae, Connaraceae, Brunelliaceae, Cephalotaceae, Cunoniaceae, Elaeocarpaceae, Tremandraceae). Two of the three pairs of sister (or nested) families that come out in molecular analyses are highly supported by floral structure: Oxalidaceae/Connaraceae and Elaeocarpaceae/Tremandraceae, whereas Cephalotaceae/Cunoniaceae are not especially similar at the level of Oxalidales. Oxalidaceae and Connaraceae share petals that are postgenitally united into a basal tube (although they are imbricate in both) but free at the insertion zone, stamens that are congenitally united at the base, uniseriate glandular hairs on the stamen filaments, and ovules that are hemianatropous to almost orthotropous. The sharing of a special type of sieve-tube plastids and of trimorphic heterostyly, studied by other authors, should also be mentioned. With Brunelliaceae, the two families share an androgynophore and nectaries at the base of the stamens in alternisepalous sectors. Elaeocarpaceae and Tremandraceae share buzz-pollinated flowers and a syndrome of features functionally connected with it. In addition, petals are larger than sepals in advanced bud, they are valvate, involute and enwrap part of the adjacent stamens, they have three vascular traces. Lignified hairs are common on the anthers and are found in the ovary locules and on the ovules (not lignified) of representatives of both families. Ovules have a chalazal appendage, and the inner integument is much thicker than the outer. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society, 2002, 140, 321–381.
Evergreen trees, unarmed; twigs with angular internodes alternating with prominent nodes. Leaves simple or pinnate (when pinnate with stipels on the rachis), opposite or ternate, mostly dentate; stipules lateral, sometimes fragmented, free; indumentum of unicellular hairs, ovary and fruit usually bristly. Inflorescences axillary, thyrsopaniculate, provided with small, usually caducous prophylls. Flowers hermaphrodite or through abortion mostly unisexual and dioecious, 4, 5 or 6(-8)-merous; sepals valvate, persistent in fruit; petals 0; nectary disk intrastaminal, adnate to calyx, cupular, 8–10-lobed; stamens free, twice as many as sepals, in 2 whorls, rarely more; filaments inserted in the notches of the nectary disk, those of outer whorl alternate with, of inner opposite to sepals; anthers bithecate, introrse, dehiscing longitudinally, the connective with a small protrusion; disk intrastaminal, thick, flat, concave, with as many indentations as stamens or staminodes, tomentose or hispidulous; carpels free, basally immersed in disk, as many as sepals or fewer and alternating with them; ovaries ovoid or ellipsoid, hairy and mostly hispid, biovulate; stylodia vertical in flower, apically hooked or curled; stigmas linear, sutural-decurrent; ovules 2 per carpel, collateral, bitegmic, anatropous, epitropous, with micropyle directed upwards; obturator +; female flowers with sterile staminodia, male with rudimentary pistil. Fruit polyfollicular; follicles tomentose and mostly additionally hispid; stylo-dia diverging horizontally; the hard endocarp detaching from softer exocarp at maturity and expelling seeds which remain attached to a placentary stalk continuing the funicle; seeds with hard, shiny testa and raised raphe; embryo large, straight, embedded in carnose, mealy, white endosperm. n = 14.
Trees, shrubs, occasionally hemiepiphytic or strangling (some Weinmannia); hairs generally simple, sometimes trichomes stellate and peltate, long, stiff and irritating in Davidsonia; bark usually grey to light brown; fissures longitudinal; stems commonly with lenticels. Leaves evergreen, winter deciduous in Eucryphia glutinosa; opposite-decussate in pairs or sometimes 3 or 4 leaves per node (spiral in Davidsonia); imparipinnate, trifoliolate, palmately compound, or unifoliolate, pinnately veined, stipellate or not, firm, often coriaceous; leaf margins toothed, often glandular-serrate, sometimes entire; venation craspedodromous, semicraspedodromous, occasionally brochidodromous or reticulodromous (some Eucryphia); small tuft or pocket domatia along midvein sometimes present; stipules often conspicuous, often interpetiolar. Inflorescences terminal, axillary, sometimes cauliflorous, paniculate, thrysoid or cymose and with straight peduncle, or capitate, rarely flowers solidarity and axillary; bracts usually stipulate; flowers often with prophylls; pedicels present or sometimes 0.
Thesis (doctoral)--Universiteit van Amsterdam, 2001. Includes bibliographical references.
Leaves of the recently described genus Ticodendron are simple, alternate, stipulate, symmetrical, elliptical with attenuate apices and acute bases, a single order of serrations representing modified rosid teeth, pinnate, craspedodromous principal venation, percurrent tertiaries, well-developed, random areolation, anomocytic stomates, and T-shaped hairs. Phenetic and cladistic analyses of leaf and other characters of Ticodendron were performed in order to determine its affinities and evolutionary relationships. Several new characters, including the number of veins joining the midrib and the venation of the marginal teeth, were added to the suite of standard leaf architectural descriptors because of their demonstrated potential as systematic indicators. After filtering out nonapplicable and symplesiomorphous characters, a total of 70 characters (48 leaf and 22 reproductive and wood) remained. These were organized into two main data sets, with the second further divided to form three subsets, which were then analyzed using a simple similarity comparison and parsimony methods. From the standpoint of its gross phenetic and phylogenetic affinities, Ticodendron belongs to a group of amentiferous taxa, with the extinct genus Fagopsis showing the strongest affinity, followed by Brunellia, Castanopsis, Castanea, Alnus, and Nothofagus. Our cladistic analyses show Ticodendron grouping with the Fagales and occasionally with the Myricaceae in a rosalean clade whose base is embedded in the Cunoniaceae. These data support recognition of the new genus as a member of a distinct family in the order Fagales and with closest affinities to Fagaceae and Betulaceae.
A cladistic analysis was conducted on restriction site variation in the inverted repeat region of the chloroplast DNA of 34 taxa, including 22 genera of Hamamelididae, nine genera of Rosidae, and one genus of Magnoliidae (sensu Cronquist). Parsimony analysis of 45 informative characters resulted in four equally parsimonious cladograms. In all trees, Hamamelididae were polyphyletic, with the rosid taxa nested within "lower" hamamelids and "higher" hamamelids in turn nested within this rosid group. The "higher" Hamamelididae (excluding Leitneria but including Betulaceae, Casuarinaceae, Fagaceae, Juglandaceae, Myricaceae, Nothofagaceae) were monophyletic in all of the most-parsimonious trees, and were always a sister group of Malus (Rosaceae). Within the "higher" hamamelid group, Nothofagus was found to be the sister taxon of the remainder of the taxa, supporting recognition of the family Nothofagaceae. The genera of Fagaceae s. str. formed a monophyletic group and were a sister group of the remainder of the "higher" hamamelids excluding Nothofugus. The sister group of the rosid-"higher" hamamelid clade in all trees was a clade that included Liquidambar, Cercidiphyllum, and representative genera of Hamamelidaceae s. str. The analysis showed Euptelea to be the basal taxon of "lower" hamamelids, whereas Trochodendron was identified as sister taxon to a larger clade comprised of Hamamelidaceae, Cercidiphyllum, rosids, and "higher" hamamelids. Cercidiphyllum was shown to be closely related to Hamamelidaceae s.l., whereas Platanus was placed several nodes away from this clade. This analysis supports recent contentions that Hamamelididae sensu Cronquist is not monophyletic. Our results contradict recent suggestions based on fossil leaves that Fagaceae are derived directly from a platanoid ancestor.
We present a cladistic analysis of genera of Cunoniaceae based on 44 structural characters using Wagner parsimony. A synthetic outgroup based on the attributes of Hamamelidaceae, Rosaceae and Fagaceae was used to polarize character states and root resulting cladograms. In all of the resulting cladograms, a clade consisting of Gillbeea, Brunellia, Spiraeanthemum, and Aistopetalum formed the sister group of the rest of the Cunoniaceae. The continued recognition of Brunellia in a monogeneric family separate from Cunoniaceae was not supported by our results. The hypothesis that apocarpy was secondarily derived within the Cunoniaceae in Brunellia and Spiraeanthemum was supported. The sister group of the above set of taxa shared the derived possession of interpetiolar stipules (although they originated in parallel in the Spiraeanthemum-Aistopetalum group). The group defined by interpetiolar stipules, consisting of the bulk of the Cunoniaceae, includes Eucryphia and Bauera, which have been placed often in monotypic families. Eucryphia is the sister group of a clade defined by bicarpellate gynoecia, which are derived within Cunoniaceae. Bauera is a member of a clade of subshrubs that includes Acrophyllum and Anodopetalum.
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