Abstract Cypselae anatomical studies have helped to understand the evolution and classification of some groups within Asteraceae. In Eupatorieae, there are many uncertainties about the Campuloclinium circumscription. There are currently two classifications for the genus, and still no consensus for their delimitation. Since structural studies have contributed to the delimitation of groups in Asteraceae, we studied the cypselae of Campuloclinium, searching how the pericarpial taxonomic features could enlighten the genus classification. We studied the fruits of eleven species of this genus through morphological and anatomical observation. Our results showed relevant features to the classification of Campuloclinium and its closely related groups. The stipitate cypsela together with other diagnostic characters are relevant to delimitation of this genus within of Eupatorieae. The trichomes present in cypselae have taxonomic proved to be a possible diagnostic character for the genus, and the six-celled trichomes are essential to distinguish C. campuloclinioides and C. hirsutum. The combination of phylogenetic and structural studies may lead to future research on the delimitation of Campuloclinium and its clades and understand how the stipitate cypselae and the phytomelanin layer evolve in Eupatorieae.

Abstract Phytomelanin is a mechanically hard, blackish, and inert substance rarely found in plants. In Asteraceae, this substance was historically associated with the Heliantheae alliance, but recent studies have observed it in unrelated groups as Heterocoma and Wunderlichia. During a taxonomic investigation, we found phytomelanin in cypselae of Lychnophora salicifolia an unusual feature in Vernonieae previously found only in Heterocoma. Furthermore, phytomelanin fills the intercellular spaces of the sclerenchymatic outer mesocarp in L. salicifolia. Our results doubt the (syn)apomorphy status in Heterocoma, suggest the phytomelanin may have not the same evolutionary significance in Lychnophorinae as in other tribes and proposes new perspectives for evolutionary studies in Asteraceae.

ABSTRACT Lepidaploa is a taxonomically complex genus of Vernonieae, which is difficult to delimit morphologically due to vegetative and reproductive characters that overlap with Chrysolaena and Lessingianthus. Anatomical studies of cypselae are taxonomically useful for delimiting subtribes, genera and species of Asteraceae, and especially within Astereae and Eupatorieae. Given the importance of cypselae to the taxonomy of tribes of Asteraceae, we searched for morphological patterns among the species of Lepidaploa. Using light microscopy, we analyzed fruits of 21 species of Lepidaploa to evaluate the taxonomic position of the genus and species of questionable placement in the group. Our results showed that the morphologies of the cypselae of species of Lepidaploa are homogeneous and similar to other species of Vernonieae. However, pappus vascularization and the number of mesocarp layers could be useful for differentiating the sister groups Chrysolaena, Lepidaploa and Lessingianthus, which present similar macro- and micro-morphological, palynological and chromosomal characters. Also, the presence of glandular trichomes and idioblasts in the cypsela, and lignified cells in the carpopodium exocarp, can be used to separate closely related species.

Ageratum has a complex circumscription, and recent studies have indicated its polyphyletism. The genus has been placed in the tribe Eupatorieae whose embryology is not fully known. Embryological data are conservative and important indicators of phylogenetic relationships and can improve family relationships. This study presents, for the first time in Eupatorieae, embryological data for Ageratum conyzoides and A. fastigiatum. Both species have common features of the family such as a unitegmic anatropous ovule, basal placentation, secretory tapetum, Polygonum megagametophyte, and Asterad embryogenesis. The data obtained reinforce the heterogeneity of the family embryology and show, for the first time, the anther wall development of the monocotyledonous type for Asteraceae. The species studied show also differences between themselves. A. conyzoides has bisporangiated and introrse anthers, conspicuous pappus, and cypselae with trichomes on the ribs, whereas A. fastigiatum has tetrasporangiate and latrorse anthers, pappus absent at maturity, and glabrous cypselae. The data presented support recent phylogenetic molecular studies, suggesting the replacement of A. fastigiatum to another genus along with Gyptidinae.

A ontogênese das cipselas e de suas partes acessórias em Campuloclinium macrocephalum, Chromolaena stachyophylla, Mikania micrantha, Praxelis pauciflora, Symphyopappus reticulatus e Vittetia orbiculata, parte delas segregadas do gênero Eupatorium, foi estudada em microscopia de luz e eletrônica de varredura. A camada de fitomelanina presente no fruto aparentemente é secretada pelo mesocarpo externo e possui espes-sura variavel entre as especies, depositando-se em espaco esquizógeno entre o mesocarpo externo e interno. As cerdas dos pápus são vascularizadas, exceto em . M. micrantha, e possuem células projetadas superficialmente, dispostas acropetamente; em S. reticulatus, algumas projeções são retrorsas e a presença de linha de fratura sob o disco floral, observada apenas nesta espécie, pode indicar processo duplo de dispersão. As numerosas diferencas registradas entre as cipselas estudadas reforcam se-gregações anteriores do gênero Eupatorium sensu lato.

The ontogeny of cypselae and their accessory parts were examined using light and scanning electron microscopy for the species Campuloclinium macrocephalum, Chromolaena stachyophylla, Mikania micrantha, Praxelis pauciflora, Symphyopappus reticulatus, and Vittetia orbiculata, some of these being segregated from the genus Eupatorium. A layer of phytomelanin observed in the fruit appears to be secreted by the outer mesocarp into the schizogenous spaces between the outer and inner mesocarp; its thickness was observed to vary among the different species examined. The bristles of the pappus are vascularized, except in M micrantha, and have cells that are superficially projected and arranged acropetally; in S. reticulatus some of the projections are retrorse and a fracture line on the floral disk that is only seen in this species may indicate a double dispersal process. Numerous differences observed among the cypselae examined here reinforce earlier segregations of the genus Eupatorium sensu lato.

Aspectos morfo-anatômicos dos frutos e sementes em desenvolvimento de Clusia parviflora são apresentados e discutidos, visando dar continuidade aos estudos com estes órgãos em Clusiaceae. O fruto é cápsula septífraga; o exocarpo suberificado deriva da epiderme externa do ovário. O mesocarpo, originado do mesofilo ovariano, permanece parenquimático. O endocarpo é derivado da epiderme interna do ovário e de três a quatro camadas subepidérmicas, cujas células tangencialmente alongadas tornam-se lignificadas, contribuindo para a deiscência do fruto. Os óvulos são anátropos, bitegumentados, com endotélio, e originam sementes também anátropas, bitegumentadas e exalbuminosas. A exotesta apresenta células de conteúdo fenólico. O exotégmen consta inteiramente de esclereídes com paredes anticlinais onduladas. O restante do tégmen torna-se colapsado. O embrião é levemente curvado e apresenta um eixo hipocótilo-radicular cilíndrico e muito desenvolvido, com dois cotilédones muito pequenos. Parece haver uniformidade em Clusiacom relação ao número de camadas no tegumento seminal maduro, mas o número de camadas no tegumento ovulífero pode ser um caráter diagnóstico em nível específico.

Aspects of the morpho-anatomy of developing fruits and seeds of Clusia parviflora are presented and discussed as a continuation of the study of these organs in Clusiaceae. The fruit is a septifrage capsule; the suberized exocarp is derived from the external epidermis of the ovary. The mesocarp originates from the ovarian mesophyll and remains parenchymal in nature. The endocarp is derived from the internal epidermis of the ovary and the endocarp is derived from the inner ovary epidermis as well as from three to four adjacent subepidermal layers, with tangentially elongated cells which become lignified and contribute to fruit dehiscence. The ovules are anatropous, bitegmic, with an endothelium, and give rise to equally anatropous seeds. The exotesta has cells containing phenolic compounds. The exotegmen consists entirely of sclerids with anticlinal and undulating cell walls, while the rest of the tegmen collapses during maturation. The embryo is slightly curved and the hypocotyl-radicle axis is well developed, with two very small cotyledons. There seems to be uniformity in the genus Clusia as regards the final number of layers in the mature seed coat, being evident the continuous lignified exotegmen and the hypocotylar embryo. It should be pointed out that the number of layers in the ovule integument can be used for diagnosis at the species level

A preocupação com o uso indiscriminado dos termos cipsela e aquênio para os frutos de Asteraceae tem sido freqüentemente destacada por especialistas na família. O presente trabalho foi desenvolvido com o intuito de verificar a existência de argumentos para justificar a adoção de um termo em detrimento do outro. Após análise histórica e anatômica, concluiu-se que há fundamento técnico para considerar cipsela e aquênio como diferentes tipos de frutos. Para Asteraceae, o correto é denominar cipsela, já que aquênios são formados somente a partir de ovários súperos, como em Plumbaginaceae.

The worry about the indiscriminate use of the terms cypsela and achene for the fruits of Asteraceae has been frequently detached by specialists in this family. The present work was developed aiming to verify the existence of arguments to justify the adoption of a term against the other. After historical and anatomical analysis, we concluded that there is technical basis to consider cypsela and achene as different types of fruits. For Asteraceae, the correct is to call cypsela; achenes are only derived from superior ovaries, as in Plumbaginaceae.

Descreveu-se a estrutura do fruto e da semente em desenvolvimento de Chorisia speciosa, visando elucidar a natureza dos pêlos que envolvem as sementes e subsidiar futuros estudos taxonômicos e ecológicos do grupo. O fruto é uma cápsula loculicida, elipsóide e apresenta o exocarpo constituído por 7-10 camadas de células com paredes celulares muito espessas com pontoações simples e evidentes. O mesocarpo parenquimático contém numerosas cavidades secretoras de mucilagem e feixes vasculares ramificados, acompanhados por fibras esclerenquimáticas. O endocarpo deriva da epiderme ventral da parede ovariana, cujas células alongam-se gradualmente, tornam-se lignificadas, cosntituindo os tricomas que envolvem as sementes. A deiscência do fruto ocorre por meio de uma sutura, na região mediana da folha carpelar, já evidente na parede ovariana. Os óvulos anátropos, bitegumentados e providos de hipóstase, originam sementes campilótropas e bitegumentadas. A testa é unisseriada. O exotégmen é constituído inteiramente por macroesclereídes; no mesotégmen ocorrem cavidades secretoras de mucilagem; e o endotégmen, que se diferencia no endotélio, torna-se colapsado na semente madura. O embrião plicado, que ocupa praticamente toda a cavidade seminal, encontra-se entre camadas de endosperma, sendo ambos ricos em material lipídico.

The structure of the fruit and seed in development of Chorisia speciosa are described with the main purpose of clarifying the origin and nature of the hairs that cover the seeds and aiding future taxonomical and ecological studies of the group. The fruit is an ellipsoid loculicide capsule and presents the exocarp formed by 7-10 cells layers, with very thick walls and evident simple pits. A great number of mucilage secretory cavities and ramified vascular bundles, accompanied by fibers, occur in the parenchymatic mesocarp. The endocarp derives from the ventral epidermis of the ovary wall, whose cells undergo a gradual elongation, become lignified, and constitute the trichomes which cover the mature seeds. The fruit aperture occurs by means of a suture evident in the ovarian wall in the middle region of the carpel leaf. Anatropous and bitegmic ovules, provided by a hypostase, give rise to campilotropous and bitegmic seeds. The testa is uniseriate, the exotegmen is completely formed by macrosclereids, and mucilage secretory cavities occur in the mesotegmen. The endotegmen, which is differentiated in the endothelium, is crushed in the mature seed. The plicate embryo, which occupies practically the entire seminal cavity, is found between endosperm layers, both being rich in lipids.