Reproductive Biology of Varronia curassavica Jacq. (Boraginaceae)

ABSTRACT Varronia curassavica, a subshrubby medicinal species associated with restinga in the Atlantic Forest, has been exploited by local people and the pharmaceutical industry. Indeed, restingas have experienced a continuous process of degradation, and thus, with species and ecosystem both at risk, efforts to support conservation actions are required. The present study aimed to evaluate aspects of V. curassavica reproductive biology. To accomplish this, morphological characterization was performed by monitoring flowering events. The availability of nectar and pollen, as well as the frequency and behavior of floral visitors and dispersers, was also evaluated. This species exhibits both heterostyly and protogyny. Anthesis is diurnal, and flowers last less than a day. The high number of flower and fruit abortions suggests that mechanisms, such as self-incompatibility intra-morphs and easily detached flowers, contribute to reduced fruit production. The high diversity of floral visitors indicate a generalist pollination syndrome. Diptera, Hymenoptera and Lepidoptera were the main pollinators, and nectar was the main resource sought by these insects. Fruits were dispersed by birds and ants. It can be concluded that the interaction of V. curassavica with several species is a key factor in its own survival and for maintaining the biological diversity of restinga.

High species richness of native pollinators in Brazilian tomato crops / Alta riqueza de espécies de polinizadores nativos em plantações Brasileiras de tomateiro

Abstract Pollinators provide an essential service to natural ecosystems and agriculture. In tomatoes flowers, anthers are poricidal, pollen may drop from their pore when flowers are shaken by the wind. However, bees that vibrate these anthers increase pollen load on the stigma and in fruit production. The present study aimed to identify the pollinator richness of tomato flowers and investigate their morphological and functional traits related to the plant-pollinator interaction in plantations of Central Brazil. The time of anthesis, flower duration, and the number and viability of pollen grains and ovules were recorded. Floral visitors were observed and collected. Flower buds opened around 6h30 and closed around 18h00. They reopened on the following day at the same time in the morning, lasting on average 48 hours. The highest pollen availability occurred during the first hours of anthesis. Afterwards, the number of pollen grains declined, especially between 10h00 to 12h00, which is consistent with the pollinator visitation pattern. Forty bee species were found in the tomato fields, 30 of which were considered pollinators. We found that during the flowering period, plants offered an enormous amount of pollen to their visitors. These may explain the high richness and amount of bees that visit the tomato flowers in the study areas. The period of pollen availability and depletion throughout the day overlapped with the bees foraging period, suggesting that bees are highly effective in removing pollen grains from anthers. Many of these grains probably land on the stigma of the same flower, leading to self-pollination and subsequent fruit development. Native bees (Exomalopsis spp.) are effective pollinators of tomato flowers and are likely to contribute to increasing crop productivity. On the other hand, here tomato flowers offer large amounts of pollen resource to a high richness and amount of bees, showing a strong plant-pollinator interaction in the study agroecosystem.

Resumo Polinizadores fornecer um serviço essencial para os ecossistemas naturais e para agricultura. Em tomateiros, as anteras são poricidas e o pólen pode sair a partir dos poros quando as flores são agitadas pelo vento. No entanto, as abelhas que vibram as anteras aumentam a carga de pólen no estigma e na produção de frutos. O presente estudo teve como objetivo identificar a riqueza dos polinizadores das flores de tomate e investigar suas características morfológicas e funcionais relacionadas com a interação planta-polinizador em plantações do Brasil central. Foram registrados o tempo de antese, duração flor, bem como o número e viabilidade de grãos de pólen e óvulos. Os visitantes florais foram observados e coletados. Os botões florais abriram-se em torno 06h30 e fechou em torno de 18h00. As flores reabrem no dia seguinte ao mesmo tempo na parte da manhã, com longevidade média de 48 horas. A maior disponibilidade de pólen ocorreu durante as primeiras horas da antese. Depois disso, o número de grãos de pólen diminuiu, especialmente entre as 10h00 às 12h00, o que é consistente com os padrões de visitação de polinizadores. Quarenta espécies de abelhas foram encontradas nos campos de tomate, 30 das quais foram consideradas polinizadores. Durante o período de floração, as plantas oferecem enorme quantidade de pólen para os seus visitantes. Isto pode explicar a alta riqueza e quantidade de abelhas que visitam as flores de tomate nas áreas de estudo. O período de disponibilidade e redução de pólen durante todo o período do dia é sobreposto com o período de alimentação das abelhas, o que sugere que elas são altamente eficazes na remoção de grãos de pólen das anteras. Muitos desses grãos provavelmente são depositados no estigma da mesma flor, levando à auto-polinização e o desenvolvimento de frutos. Abelhas nativas (Exomalopsis spp.) são polinizadores efetivos de flores de tomate, podendo contribuir para o aumento da produtividade das culturas. Por outro lado, as flores de tomate oferecem grandes quantidades de pólen de recursos para uma alta riqueza e quantidade de abelhas, que mostram um estudo forte interação planta-polinizador nos agroecossistemas.

Anatomía Floral de Peristethium leptostachyum (Loranthaceae) / Floral anatomy of Peristethium leptostachyum (Loranthaceae)

ResumenPeristethium leptostachyum es una especie hemiparásita de la familia Loranthaceae, distribuida en Colombia, Costa Rica, Ecuador, Perú, Venezuela y Panamá. Previamente tratada como Struthanthus leptostachyus, la especie fuerecientemente fue reubicada en Peristethium junto con otras que previamente estaban en los géneros Cladocolea y Struthanthus. La decisión de reconocer a Peristethium como género es controversial y fue tomada con base en caracteres de la inflorescencia y de la flor; en tanto que la monofilia de los tres géneros nombrados es incierta. En esta investigación se estudió la morfoanatomía de flores e inflorescencias de Peristethium leptostachyum, detallando la estructura del androceo y gineceo, así como los procesos de microgametogénesis y megagametogénesis; adicionalmente se realizaron comparaciones con especies afines y precisiones en relación con las diagnosis previas. Se recolectaron flores en diversas fases de desarrollo en Santa María (Boyacá-Colombia), se prepararon y analizaron bajo microscopio secciones histológicas teñidas con astrabluefucsina, además de disecciones bajo estereomicroscopio. Los resultados mostraron que P. leptostachyum comparte caracteres inflorenciales con Cladocolea (inflorescencia determinada, flor terminal ebracteada), pero también con Struthanthus (pares de tríadas a lo largo del eje, brácteas caducas y flores actinomorfas). Las flores de P. Leptostachyum de Santa María son claramente hermafroditas, con androceos y gineceos totalmente desarrollados; lo cual contradice la descripción hecha por Kuijt que reporta una condición dioica para esta especie. El androceo resultó afín al de Struthanthus vulgaris, con tapetum glandular y microsporogénesis simultánea; en contraste, Cladocolea loniceroides presenta tapetum periplasmodial y microesporogénesis sucesiva. El gineceo de P. leptostachyum, al igual que en Cladocolea, Struthanthus y Phthirusa, es unilocular con mamelón y tejido arquesporial orientado hacia el estilo, el cual es sólido y con tejido amilífero. P. leptostachyum es afín a Cladocolea loniceroides y difiere de Struthanthus vulgaris por presentar varios sacos embrionarios y pelvis (hipostasa) no lignificada. La presencia de un canal estilar sólido se propone como sinapomorfía de la tribu Psittacanthinae. Dado que P. Leptostachyum comparte caracteres anatómicos florales tanto con Cladocolea como con Strutanthus, la relación entre estos tres géneros no queda resuelta, se requieren estudios filogenéticos para establecer esta relación y poner a prueba las hipótesis de monofilia de cada uno de ellos.

AbstractPeristethium leptostachyum is a hemiparasite species of the family Loranthaceae, distributed in Colombia, Costa Rica, Ecuador, Peru, Venezuela and Panama. Previously treated as Struthanthus leptostachyus, the species was recently transferred to Peristethium together with other species of Cladocolea and Struthanthus. The present research describes the inflorescence and floral morphoanatomy of Peristethium leptostachyum, detailing the structure of the androecium and gynoecium and the processes of microgametogenesis and megagametogenesis, thus allowing comparison with Struthanthus and Cladocolea. Flowering material was collected in February and August 2012, in Santa María, Boyacá, Colombia. Histological sections were prepared and stained with astrablue-fuchsin and floral dissections were performed under a stereomicroscope. Peristethium leptostachyum shares inflorescence characters with Cladocolea (determinate inflorescence, ebracteate terminal flower), but also with Struthanthus (pairs of triads along the axis, deciduous bracts and actinomorphic flowers). The flowers of P. leptostachyum from Santa María are clearly hermaphrodites with androecium and gynoecium fully developed. This observation contradicts the description by Kuijt who reported this species to be dioecious. The androecium was observed to be similar to that of Struthanthus vulgaris, with a glandular tapetum and simultaneous microsporogenesis; in contrast, Cladocolea loniceroides has a periplasmodial tapetum and successive microsporogenesis. The gynoecium of P. leptostachyum, like that of Cladocolea, Struthanthus and Phthirusa, has a unilocular ovary with a mamelon and arquesporial tissue isoriented towards the style, which in turn is solid and amyliferous. Peristethium leptostachyum is similar to Cladocolea loniceroides and differs from Strutanthus vulgaris in presenting multiple embryo sacs and an unlignified pelvis (hipostase). The presence of a solid stylar canal is proposed as a synapomorphy of the tribe Psittacanthinae. Given that P. leptostachyum shares characters with both Cladocolea and Struthanthus generic placement cannot be clearly determined on the basis of anatomical evidence. Phylogenetic studies that include representative species of all three genera are desirable to test hypotheses of monophyly. The sexual system observed here in P. leptostachyum is different from that reported by Kuijt and more studies are needed to identify the factors (geographic, ecological, etc.) that influence this variation. Rev. Biol. Trop. 64 (1): 341-352. Epub 2016 March 01.

Salinity-induced changes in the morphology and major mineral nutrient composition of purslane (Portulaca oleracea L) accessions

This study was undertaken to determine the effects of varied salinity regimes on the morphological traits (plant height, number of leaves, number of flowers, fresh and dry weight) and major mineral composition of 13 selected purslane accessions. Most of the morphological traits measured were reduced at varied salinity levels (0.0, 8, 16, 24 and 32 dS m-1), but plant height was found to increase in Ac1 at 16 dS m-1 salinity, and Ac13 was the most affected accession. The highest reductions in the number of leaves and number of flowers were recorded in Ac13 at 32 dS m-1 salinity compared to the control. The highest fresh and dry weight reductions were noted in Ac8 and Ac6, respectively, at 32 dS m-1 salinity, whereas the highest increase in both fresh and dry weight was recorded in Ac9 at 24 dS m-1 salinity compared to the control. In contrast, at lower salinity levels, all of the measured mineral levels were found to increase and later decrease with increasing salinity, but the performance of different accessions was different depending on the salinity level. A dendrogram was also constructed by UPGMA based on the morphological traits and mineral compositions, in which the 13 accessions were grouped into 5 clusters, indicating greater diversity among them. A three-dimensional principal component analysis also confirmed the output of grouping from cluster analysis.

Biologia floral e sistema reprodutivo da erva-baleeira (Varronia curassavica Jacq. ) / Flowering phenology and reproductive system of the Cordia (Varronia curassavica Jacq. )

RESUMOO conhecimento do sistema reprodutivo é fundamental para a conservação e manejo de uma espécie. O objetivo deste trabalho foi descrever a fenologia da floração, a antese, registrar os insetos visitantes no período de floração, determinar as características morfométricas das flores e o sistema reprodutivo da erva-baleeira, em um ambiente de Cerrado do Norte de Minas Gerais. Entre maio a dezembro de 2012 foi caracterizado o comportamento fenológico da floração. Na análise da fenologia floral foi determinado: o crescimento da inflorescência, o número de flores e frutos por inflorescências. Utilizou-se seis acessos que tiveram dez inflorescências marcadas em cada acesso, totalizando 60 inflorescências. A antese foi determinada utilizando quatro inflorescências em duas plantas. Os visitantes florais foram observados in loco e capturados em três dias consecutivos de coleta. As características morfométricas foram determinadas com paquímetro utilizando 20 flores, sendo cinco flores de quatro acessos. Para determinar o sistema reprodutivo utilizou-se a razão pólen:óvulo (P:O), utilizando 50 flores, sendo 10 flores de cinco acessos em pré-antese. Nas condições de Montes Claros, o crescimento das inflorescências de erva-baleeira ocorreu entre meados de agosto e início de outubro, totalizando 45 dias. O florescimento foi observado entre meados de setembro e final de outubro, enquanto a frutificação ocorreu de meados de outubro a início de dezembro, sendo que ambos ocorreram de forma irregular. A antese floral de erva-baleeira, neste estudo, ocorre entre 7:00 e 11:00 horas. Os insetos visitantes pertencem as ordens Coleoptera, Hemiptera, Diptera e Hymenoptera. As flores apresentaram o diâmetro de 2,13 ± 0,05 (mm), o comprimento de 3,29 ± 0,08 (mm), diâmetro do ovário de 0,70 ± 0,02 (mm), o comprimento do ovário de 2,48 ± 0,12 (mm), o diâmetro da antera de 0,67 ± 0,01(mm) e o comprimento da antera de 0,93 ±0,02 (mm), quatro óvulos e cinco anteras por flor. A razão P:O foi de 576,542, indicando que a espécie é alógama facultativa.

ABSTRACTThe knowledge of the reproductive system is essential for the conservation and management of the species. This study aimed on several procedures, as follows: to describe the phenology of flowering, the anthesis; to record the visiting insects during flowering and to determine the morphometric characteristics of the flowers and the reproductive system of the "erva-baleeira", in an environment of Northern Cerrado in the State of Minas Gerais. From May to December 2012, the flowering phenology of six access was characterized. In the analyzes of floral phenology it were determined the growth of the inflorescence and the number of flowers and fruits per inflorescence by using six access which had ten inflorescences marked in each access l, totalizing sixty inflorescences. The anthensis was assessed using four inflorescences in two plants. The flower`s visitors were observed live and captured in three consecutive days of sampling. The morphometric characteristics were determined with a caliper using 20 flowers, with five flowers from four access. In order to determine the reproductive system it was employed a ratiopollen: ovule (P: O) with 50 flowers and 10 of them belonging to five accesses in the pre-anthesis. Under the conditions of Montes Claros, the growth of inflorescences from Cordia occurred between mid-August and early October, totalizing 45 days. The flowering was observed between mid-September and late October, and the fruiting occurred from mid-October to early December. Both phases happened irregularly. The anthesis of the Cordia, in this study, occurred between 7:00 and 11:00 o`clock.. The visiting insects identified were from the orders of Coleoptera, Hemiptera, Diptera and Hymenoptera. The flowers exhibited a diameter of 2.13 ± 0.05 (mm), length of 3.29 ± 0.08 (mm), diameter of 0.70 ± 0.02 Ovarian (mm), the length of the ovary was 2.48 ± 0.12 (mm), the diameter of the anther 0.67 ± 0.01 (mm) and the length anther was 0.93 ± 0.02 (mm), with five anthers and four ovules per flower. The reason P: O was 576.542, indicating that the species is facultative allogamous.

Estructura floral de dos especies de Trachycarpeae (Arecaceae) / Floral structure of two species of Trachycarpeae (Arecaceae)

Copernicia and Washingtonia are two genera of the Trachycarpeae for which no subtribal classification has been proposed, mainly because of the lack of resolution in phylogenetic studies. Morphology and anatomy of flowers whithin Coryphoideae have proven useful for taxa delimitation and supporting relationships among their members. A description of the morphological and anatomical structure of flowers of C. tectorum and W. filifera is presented in order to explore reproductive characters that may clarify their classification within the subfamily and to contribute with floral biology studies. Flowers of cultivated specimens of both taxa and developing fruits of C. tectorum were fixed in FAA, dissected for morphological analysis, and parafin-embedded flowers and fruits were serially sectioned for obtaining permanent slides, using conventional techniques and safranin-fast green staining. All procedures were carried out in the Laboratory of Morpho-Anatomy, Agronomy Faculty of the Universidad Central de Venezuela (UCV). Both species have hermaphroditic flowers. C. tectorum flowers have a thick and pubescent perianth, six stamens with filaments forming a tube fused to the corolla, with rounded projections and an acute apex where the anthers are inserted. W. filifera flowers have an irregularly dentate calyx, and a shortly acuminate corolla, six stamens united by their filaments to the corolla which at the same time are briefly fused to the gynoecium. Cells with druse crystals in the staminal tube are reported for C. tectorum. Only one of the carpels of the gynoecium of C. tectorum develops at fruit stage, and a layer of abundant raphide cells forming a crustaceous endocarp in mature fruits, was found. W. filifera presents the perianth mesophyll with few layers of thick walled cells and schlerenchymatic tissue, gynoecium with apically fused carpels in the ventral region of ovary, free at the base and the apex of the style, where the ventral sutures are opened. C. tectorum has a ventral hypodermis in the petals made of large and thick walled cells, gynoecium with apically fused carpels in the ovary, free and adpressed basally, style-stigma completely fused, and stylar transmission channel absent distally. Distinct stylar canals in C. tectorum, united distally in W. filifera confirm the close relationship between these species and subtribe Livistoninae. Also, some floral morpho-anatomical similarities (e.g. fleshy calyx base and a hypodermis with thickened cell walls in petals) were found between C. tectorum and Pritchardia, supporting the affinities between both genera. Rev. Biol. Trop. 62 (3): 1137-1146. Epub 2014 September 01.

Copernicia y Washingtonia son dos de los géneros de Trachycarpeae para los cuales no ha sido propuesta una clasificación subtribal debido a la carencia de resolución en los estudios filogenéticos. La morfología y anatomía floral dentro de Coryphoideae han sido útiles para la delimitación de taxones y apoyo de relaciones. En este trabajo se presenta una descripción de la estructura morfo-anatómica de las flores de C. tectorum y W. filifera, con la finalidad de explorar caracteres reproductivos que puedan aclarar su ubicación dentro de la subfamilia y contribuir con estudios de biología floral. Se fijaron flores de especimenes cultivados de ambos taxones y frutos en desarrollo de C. tectorum, se realizaron disecciones para las descripciones morfológicas y cortes anatómicos seriados para la obtención de láminas permanentes utilizando técnicas convencionales con parafina. Todos los procedimientos fueron llevados a cabo en el Laboratorio de Morfo-Anatomía, Facultad de Agronomía de la Universidad Central de Venezuela (UCV). Ambas especies tienen flores hermafroditas. Las flores de C. tectorum tienen perianto engrosado y pubescente. Las flores de W. filifera presentan un cáliz irregularmente dentado y corola corto-acuminada. Se reportan células con drusas en el tubo estaminal de C. tectorum. Uno solo de los carpelos de C. tectorum se desarrolla en fruto y se encontró una capa de células con rafidios que podrían formar el endocarpo crustáceo en los frutos maduros. W. filifera presenta pocas capas de células de paredes engrosadas y tejido esclerenquimático en el mesofilo del perianto, gineceo con carpelos fusionados en el ápice del ovario (en su porción ventral), libres en la base y en el ápice del estilo, donde presenta suturas ventrales abiertas. Los pétalos de C. tectorum poseen una hipodermis ventral de células con paredes muy engrosadas, gineceo con carpelos fusionados en el ápice del ovario, libres y adpresos en su base, región estilar-estigmática fusionada y canal de transmisión estilar ausente distalmente. La presencia de canales estilares separados en C. tectorum, unidos distalmente en W. filifera confirman la relación cercana de estas especies con la subtribu Livistoninae. Igualmente, se encontraron algunas similitudes entre C. tectorum y Pritchardia (p.e. base carnosa del cáliz y epidermis ventral con células de paredes engrosadas en los pétalos), apoyando ciertas afinidades entre ambos géneros.

Anatomía y desarrollo de estambres y carpelos en Drimys granadensis (Winteraceae) / Anatomy and development of stamens and carpels of Drimys granadensis (Winteraceae)

Winteraceae has long been considered a family with early diversification among angiosperms, with characters such as: flowers with many spirally arranged parts and apocarpic ovary formed by plicated carpels with sessile stigma. In Drimys, the presence or absence of conspicuous glands on the connective of the stamens have been used as a taxonomic character, and it is considered a synapomorphy for the clade including Drimys angustifolia, D. brasiliensis, D. granadensis and D. roraimensis (Northeastern clade); however, the anatomy of stamens and carpels has only been studied in detail for D. winteri (Southwestern clade). In this research, the presence and the structure of glands on the connective of stamens was studied in seven species of the genus from herbarium specimens, and a detailed study of the anatomy and development of stamens and carpels was carried out by scanning electron and optic microscopy in Drimys granadensis. We found similarities between D. granadensis and D. winteri for the following characters: Basic type anther wall formation, secretory tapetum that collapses at maturity, intermediate type microsporogenesis with formation of a transient cell plate in telophase I, ascidiated carpel due to the formation of an adaxial lip during development, stigma closed by interdigitation of epidermal cells. We also determined that the large glands on anther mature connective are originated by an overgrowth of subepidermal oil cells; this character is a Northeastern Drimys clade synapomorphy, while it was absent in both Drimys of Southwestern clade (which includes D. andina, D. confertifolia and D. winteri), and the rest of the Winteraceae. We are proposing the hypothesis that the highly variable enviromental conditions in the tropics where Drimys Northeastern clade is distributed, with a wide range of pollinators, would be associated with the emergence of glandular conspicuously stamens; while, as a prediction to be confirmed, temperated Southwestern clade species, without conspicuous glands, should have a smaller range of visitors and/or pollinators, or even be anemophilous species as Drimys confertifolia. Rev. Biol. Trop. 62 (3): 1147-1159. Epub 2014 September 01.

La familia Winteraceae ha sido tradicionalmente considerada como de diversificación temprana entre las angiospermas por varios caracteres, entre ellos: flores con muchas partes distribuidas en espiral y ovario apocarpico formado por carpelos de tipo plicado con estigma sésil. En el género Drimys, la presencia o ausencia de glándulas conspicuas sobre el conectivo de los estambres ha sido usado como un carácter taxonómico, y su presencia se considera como una sinapomorfía del clado formado por Drimys angustifolia, D. brasiliensis, D. granadensis and D. roraimensis (clado nororiental); sin embargo, la anatomía de estambres o carpelos ha sido solamente estudiada en detalle en D. winteri (clado suroccidental). En esta investigación, la presencia y estructura de las glándulas del conectivo fue estudiada en las siete especies del género Drimys a partir de ejemplares de herbario, además, se realizó un estudio detallado de la anatomía y desarrollo de estambres y carpelos de Drimys granadensis empleando microscopía óptica y microscopía de barrido; y se compararon con los resultados previos en Drimys winteri. Se encontraron similitudes en los siguientes caracteres: formación de la pared de la antera de tipo básico, tapetum secretor que colapsa en la madurez, microsporogénesis de tipo intermedio con formación de una placa celular transitoria en telofase I, carpelo ascidiado debido a la formación de un labio adaxial durante el desarrollo, estigma cerrado por interdigitación de células epidérmicas. Se determinó que las glándulas de gran tamaño sobre el conectivo de la antera madura se originaron por sobrecrecimiento de células oleíferas subepidérmicas, siendo una sinapomorfía del clado nororiental, que esta ausente tanto en el clado suroccidental de Drimys (D. andina, D. confertifolia y D. winteri), como en el resto de la familia. Se propone la hipótesis de que las condiciones altamente variables en las zonas tropicales donde se distribuye el clado nororiental de Drimys, con una amplia gama de polinizadores, estarían asociadas al surgimiento de estambres conspicuamente glandulares; en tanto que proponemos como predicción que las especies circunscritas a la zona templada del Sur de Suramérica (Clado suroccidental), sin glándulas conspicuas, deberían presentar una menor variedad de visitantes y/o polinizadores, o incluso ser especies anemófilas como Drimys confertifolia.

Estructura, ontogenia y vascularización de las flores e inflorescencias de Drimys granadensis (Winteraceae) / Structure, ontogeny and vascularization of the flowers and inflorescences of Drimys granadensis (Winteraceae)

Drimys granadensis is a widespread species in montane forests of South and Central America. In this research, the structure, ontogeny, phyllotaxis and vascularization of the flowers and inflorescences of this species was studied in a population from the Eastern hills of Sabana de Bogota, Colombia. The methods used applied both optical microscopy, with astra blue-fuchsin staining, and scanning electron microscopy, using critical point dryed and gold-paladium metallized samples. Besides, results were compared with those of Drimys winteri, a widely studied species distributed in Chile and Argentina. Additionally, we studied the detail of the floral anatomy to determine the bracteal or calicine identity of the caliptra. I confirmed the proliferative status of the monothelic inflorescence, discarding alternative explanations of the terminal flower identity. I found that uniflorescences have an acropetal development until the terminal meristem becomes the terminal flower, then this flower develops rapidly resulting in a determined uniflorescence. I found pseudosyphonosthelic vascularization in peduncles and pedicels. Besides, I discovered some evidence in the vascular and anatomical structures, to consider the caliptra as the fusion product of various structures and therefore of calicine origin. The caliptra showed a whorled phyllotaxis, but the petals, stamens and carpels presented a spiral condition; phyllotaxis change was explained by the long time lapse between the initiation of the calyx and the corolla. I found great similarities among the inflorescences of D. granadensis and D. winteri; they were different in the proliferation start time, and in the frequent presence of nomophylls in D. granadensis, in contrast to the presence of reduced bracts and bracteoles in D. winteri inflorescences.

Drimys granadensis es una especie de amplia distribución en los bosques montanos de Sur y Centro América. En esta investigación se estudiaron, mediante microscopía óptica y electrónica de barrido, la estructura, ontogenia, filotaxis y vascularización de sus flores e inflorescencias, y fueron comparadas con las de Drimys winteri, especie distribuida en Chile y Argentina. Adicionalmente, se buscó evidencia para determinar la identidad bracteal o calicina de la caliptra de sus flores. Se confirmó la condición monotélica proliferante de la inflorescencia, descartando explicaciones alternativas de identidad de la flor terminal. Las uniflorescencias presentan un desarrollo acrópeto, hasta que el meristemo terminal se transforma en flor terminal, entonces esta flor se desarrolla rápidamente dando lugar a una uniflorescencia determinada. La vascularización es pseudosifonostélica para pedúnculos y pedicelos. Se encontró evidencia en la vascularización y estructura anatómica para considerar la caliptra como el producto de fusión de varias estructuras y, por tanto, de origen calicino. La caliptra presentó una filotaxis verticilada, pero los pétalos, estambres y carpelos presentaron una filotaxis espiralada; el cambio se explicó por el tiempo prolongado entre la iniciación de cáliz y corola. Las inflorescencias de D. granadensis y D. winteri son muy similares; siendo diferente el tiempo de inicio de la proliferación y la frecuente presencia de nomófilos en las inflorescencias de D. granadensis, en contraste con la presencia de brácteas y bracteolas reducidas en D. winteri.

Morphoanatomy of the flower of Syagrus inajai (SPRUCE) Becc. (Arecaceae- Arecoideae- Attaleinae), Amazon / Morfoanatomia floral de Syagrus inajai (Spruce) Becc. (Arecaceae- Arecoideae- Attaleinae), Amazônia

The occurrence of Syagrus inajai (Spruce) Becc., popularly known as pupunha palm, among other names, has been registered in the Guianas and in the North of Brazil in areas of terra firme (non-flooding) and gallery forests. In order to characterize the inflorescence and further knowledge of this family, a morphoanatomical study was carried out of the palm S. inajai in a green area of the Campus of the Federal University of Amazonas - UFAM, Manaus, Amazonas. The inflorescences are branched to one order, pedunculate, and interfoliar, measuring 62-82 cm in length, with woody bracts with longitudinal grooves on the external surface, and flowers in triads. The number of flowers to each inflorescence varies from 5,904 to 17,316 for staminate flowers, and from 180 to 3,528 for pistillate flowers. Staminate flowers with six anthers and one vascular bundle each; three-lobed pistillodium, vascularized pistillodium. Its pistillate flowers have six staminodia joined to form a circle, syncarpic, tricarpellary, trilocular gynoecium, one ovule to each locule, synascidiate in the ovary, and plicated above. Tripartite stigma, apical and sessile, with epidermis composed of elongated papillary cells, pattern of epidermis that is maintained throughout the stylar canal. Bitegmented, anatrope, pachychalazal ovule.

Syagrus inajai (Spruce) Becc., popularmente conhecida como pupunha-brava entre outras denominações, teve sua ocorrência registrada para Guianas e no norte do Brasil em áreas de floresta de terra firme e floresta de galeria. Com o intuito de caracterizar a inflorescência ampliando o conhecimento a respeito da família foi realizado o estudo morfoanatômico da palmeira S. inajai, em área verde do Campus da Universidade Federal do Amazonas - UFAM, Manaus, Amazonas. A inflorescência é ramificada em primeira ordem, pedunculada, interfoliar com 62-82 cm de comprimento, bráctea lenhosa com fissuras longitudinais na superfície externa e flores em tríades. O número de flores por inflorescência varia entre 5.904 - 17.316 para flores estaminadas e 180 - 3.528 para as flores pistiladas. Flores estaminadas com seis anteras e um feixe vascular cada; pistilódio trífido e vascularizado. Flores pistiladas com seis estaminódios unidos formando um círculo. Gineceu sincárpico, tricarpelar, trilocular, um óvulo por lóculo, sinascidiado no ovário e plicado acima. Estigma tripartido, apical e séssil, com epiderme composta por células papilosas alongadas, padrão de epiderme que se mantem por todo canal estilar. Óvulo é anátropo, paquicalazal e bitegumentado.

Floral biology of Tropaeolum majus L. (Tropaeolaceae) and its relation with Astylus variegatus activity (Germar 1824) (Coleoptera: Melyridae)

Tropaeolum majus L. (nasturtium) is a culture popularly known by its medicinal, ornamental and culinary utility. This work aimed to evaluate the interaction between Astylus variegatus (Germar 1824) (Coleoptera: Melyridae) and the nasturtium flowers associated to the weeks of flourishing, in order to conceive the floral mechanims used by the species to attract A. variegatus. The insects collection was achieved with an entomologic net by the sweeping method, during the weekly flowering, at two hours, from 7am to 5pm. The studies of floral biology were carried out in twenty flowers by the analysis of measures and the arrangement of the floral parts. The nasturtium flower offers pollen and nectar to the insect as a compensation. However, it was observed that A. variegatus only collects pollen as a nutritious resource. Tropaeolum majus presents several flowers for each individual and, during the visits, A. variegatus walked throughout the interior of the flower, among the stamens and contacting the ventral region of its body, and while passing through the stigma it lodged the pollen. Sometimes, it uses the internal cavity for sheltering and mating. The visits occurred preferably on the second and fifth weeks of flourishing, with a populational fluctuation between 9am and 11am.

Tropaeolum majus L. (capuchinha) é uma cultura muito conhecida pelo seu valor medicinal, ornamental e culinário. O objetivo do trabalho foi avaliar a interação entre Astylus variegatus (Germar 1824) (Melyridae) com as flores de capuchinha, associado às semanas de florescimento, visando entender os mecanismos florais utilizados pela espécie na atração de A. variegatus. A coleta dos insetos foi realizada com rede entomológica pelo método varredura durante a floração, semanalmente, a cada duas horas, no período de 7 às 17 horas. Os estudos da biologia floral foram realizados em vinte flores, analisando as medidas e disposição das peças florais. A flor de capuchinha oferece ao inseto pólen e néctar como recompensa, contudo, foi observado que A. variegatus coleta apenas pólen como recurso alimentar. Tropaeolum majus apresenta várias flores por indivíduo sendo que durante as visitas A. variegatus caminhava por todo o interior da flor, entre os estames contatando a região ventral de seu corpo e, ao passarem pelo estigma, depositava o pólen. Em alguns momentos, utiliza a cavidade interna para abrigo e acasalamento. As visitas ocorreram preferencialmente na segunda e quinta semanas de florescimento com pico populacional entre as 9 e 11:00h.

Contrasting bee pollination in two co-occurring distylic species of Cordia (Cordiaceae, Boraginales) in the Brazilian semi-arid Caatinga: generalist in C. globosa vs. specialist in C. leucocephala

In this study we compare the reproductive biology of Cordia globosa and C. leucocephala (Cordiaceae, Boraginales; formerly referred to Boraginaceae) to understand the functioning of the floral morphs and the relations with their effective pollinators. The species are synchronopatric, distylic, and self-incompatible. Though they share melittophilous traits, the main visitor and pollinator of C. globosa was the generalist and exotic bee Apis mellifera, while the only one of C. leucocephala was the oligoletic bee Ceblurgus longipalpis. These two latter species are restricted to the Caatinga of NE Brazil, contrasting with the wide distribution of Cordia globosa. While the fruit-set for C. globosa was high, independently if the pollen donor/stigma receptor was a pin (long-styled) or thrum (short-styled) individual, in C. leucocephala the fruit-set was low and occurred only when a thrum individual was the pollen donor. This raises the possibility of this species moving towards dioecy. The high natural fruit-set of C. globosa confirms the generalist bee as its effective pollinator. The low fruit-set after manual crosses in C. leucocephala may be due to low pollen viability. Additionally, the low natural fruit-set (two times lower than after crosses) may be related with the foraging behavior of the specialist pollinator.

Neste estudo comparamos a biologia reprodutiva de Cordia globosa e C. leucocephala para entender a função dos orfos florais e as relações com seus polinizadores efetivos. As espécies são sincronopátricas, distílicas e auto-incompatíveis. Embora elas compartilhem atributos melitófilos, o principal visitante e polinizador de C. globosa foi Apis mellifera, abelha generalista e exótica, enquanto o de C. leucocephala foi a abelha oligolética Ceblurgus longipalpis. Essas duas últimas espécies são restritas à Caatinga do Nordeste do Brasil, contrastando com a ampla distribuição de C. globosa. Enquanto a formação de frutos de C. globosa foi alta, independente se o doador ou receptor de pólen era um indivíduo brevi- ou longistilo, em C. leucocephala a formação de frutos foi baixa e ocorreu apenas quando o doador de pólen era brevistilo. Este fato levanta a possibilidade desta espécie estar se movendo em direção à dioicia. A alta formação natural de frutos de C. globosa confirma a abelha generalista como seu polinizador efetivo. A baixa formação de frutos após cruzamentos manuais e C. leucocephala pode ser devida à baixa viabilidade polínica. Adicionalmente, a baixa formação natural de frutos (duas vezes menor que a de cruzamentos) pode estar relacionada com o comportamento de forrageamento do polinizador especialista.

Variación florística de especies arbóreas a escala local en un bosque de tierra firme en la Amazonia colombiana / Floristic variation of canopy tree species at a local scale on tierra firme forests in Colombian Amazonia

El presente estudio se llevó a cabo en cinco hectáreas de una parcela permanente establecida en el Parque Nacional Amacayacu, Amazonia colombiana. En éste, se evaluó el efecto de la variación ambiental y la configuración espacial sobre los patrones florísticos de las especies arbóreas (DAP>10 cm) a escala local en un bosque de tierra firme. Se estudió la variación florística y ambiental en cuadrantes de 20x20 m. Adicionalmente, se consideraron diferentes categorías de abundancia (total, alta, media y baja). Se utilizó el Análisis de Correspondencia Linealizado y el Análisis de Correspondencia Canónica, seguido de una partición de la variación, para cuantificar la magnitud a la cual el ambiente y la limitación en dispersión determinan la variación florística. La fracción espacial, representando procesos de autocorrelación como la limitación en dispersión, se analizó mediante dos métodos: Asumiendo un polinomio de tercer grado y por el método de Coordenadas Principales de Matrices Vecinas (PCNM). La diversidad beta de la parcela fue baja. El PCNM aparece como el método de análisis más apropiado para estudios a esta escala. Las diferencias florísticas explicadas a lo largo de la parcela de 5-ha fueron principalmente asociadas con procesos biológicos como la limitación en dispersión. La mayor parte de la variación florística, no obstante, no fue explicada por las variables ambientales o espaciales consideradas. En conclusión, estos resultados sugieren que procesos aleatorios son determinantes esenciales de la variación espacial de las especies arbóreas a escala local en tierra firme en los bosques en el Parque Nacional Amacayacu.

This study was carried out in a 5-ha permanent plot established in the Amacayacu National Park, Colombian Amazonia. We assessed the extent at which floristic patterns of tree species were determined by either the environmental variation or the spatial configuration in tierra firme forests. Floristic and environmental variation was assessed on quadrants of 20x20 m. To run the analysis, four different categories of abundance (all, high, medium and low) were considered. We used Detrended Correspondence Analysis and Canonical Correspondence Analysis, followed by a variation partitioning, to analyze the patterns of species distribution in order to quantify the role played by both environmental filtering and dispersal limitation. The spatial template, which represents dispersal limitation, was evaluated in two different ways: using a third degree polynomial and using the Principal Coordinates of Neighbour Matrix (PCNM) method. Beta diversity was relatively low within the plot. PCNM appears as the most appropriate method to analyze the spatial template at this scale. The floristic differences explained along the 5-ha plot were mainly associated to biological processes, such as dispersal limitation. The largest proportion of community variation in our dataset was unexplained by either environmental or spatial data. In conclusion, these results support random processes as the major drivers of the spatial variation of tree species at a local scale on tierra firme forests in the Amacayacu National Park.

In vitro regeneration and genetic fidelity of Tigridia pavonia (L. f.) DC

Plants of Tigridia pavonia (L.f.) DC were regenerated from twin-scaling explants cultured on Murashige and Skoog and N6 basal medium. The highest formation of shoots per responding explant was obtained on N6 medium supplemented with 4.5 uM 2,4-dichlorophenoxy acetic acid in combination with 2.2 uM benzylaminopurine. Shoots rooted readily on N6 basal medium supplemented with 1 g I-1 activated charcoal and 2.6 uM naphtalenacetic acid. The rooted shoots achieved 100 percent survival. Inter Simple Sequence Repeat analysis was carried out to check for possible genetic alterations in plants obtained after two consecutive subcultures. The results revealed that the recovered plants did not exhibit any type of polymorphism.

Anatomy and ontogeny of the pericarp of Pterodon emarginatus Vogel (Fabaceae, Faboideae), with emphasis on secretory ducts

Discrepant and incomplete interpretations of fruits of Pterodon have been published, especially on the structural interpretation of the pericarp portion that remain attached to the seed upon dispersal. The present work clarified these doubts and analyzed ultrastructural aspects of the Pterodon emarginatus diaspores using light and transmission electron microscopes. Cell divisions are prevalent among the initial phases of development, and the subadaxial and adaxial meristems form the fibrous inner mesocarp and the endocarp composed of multi-seriate epidermis, respectively. At the median mesocarp, numerous secretory ducts differentiate between the lateral bundles, by lytic process. After lysis of the central cells and the formation of the lumen, the ducts show unistratified secretory epithelium with dense cells; oil droplets are observed on the secretory epithelium and the subadjacent tissues. At maturity, the uniseriate exocarp and the outer mesocarp slough off in an irregular fashion, leaving the diaspore composed of a papery and brittle wing linked to a seed chamber that includes the median mesocarp composed of lignified cells, bordering vascular bundles and many secretory ducts whose epithelial cells develop large vacuoles that accumulate oleoresins. The Pterodon emarginatus fruit is a cryptosamara.

Interpretações discrepantes e incompletas têm sido conferidas ao fruto de Pterodon, especialmente no que tange à determinação estrutural da porção pericárpica que acompanha a semente na dispersão. Assim, com o objetivo de dirimir tais dúvidas e analisar a organização ultra-estrutural das estruturas secretoras presentes no diásporo de Pterodon emarginatus, realizaram-se estudos convencionais aos microscópios de luz e eletrônico de transmissão. Nas fases iniciais de desenvolvimento do fruto, prevalecem divisões celulares; pela ação do meristema subadaxial e do adaxial, formam-se, respectivamente, o mesocarpo interno fibroso e o endocarpo composto por epiderme multisseriada. No mesocarpo mediano, entre os feixes vasculares laterais, diferenciam-se numerosos ductos secretores lisígenos. Após a lise das células centrais e formação do lume, os ductos apresentam epitélio secretor uniestratificado, com células densas; gotas de óleo são observadas no epitélio secretor e tecido subjacente. Na maturidade, o exocarpo unisseriado e o mesocarpo externo, ambos fenólicos, descamam irregularmente, sendo o diásporo constituído pela ala papirácea e quebradiça, ligada ao núcleo seminífero que abrange o mesocarpo mediano de células lignificadas, margeando feixes vasculares e muitos ductos secretores, que apresentam acúmulo de oleorresina e cujas células epiteliais tornam-se vacuoladas. O fruto de Pterodon emarginatus é, portanto, uma criptossâmara.

Fenología reproductiva y anatomía floral de las plantas Aloe vera y Aloe saponaria (Aloaceae) en Cumaná, Venezuela

Reproductive phenology and flower anatomy of the plants Aloe vera and Aloe saponaria (Aloaceae) in Cumana, Venezuela. The reproductive phenology and the floral anatomy of two species of Aloe kept under nursery conditions were analized in eastern Venezuela from September 2001 to September 2002. A. vera flowered between December and May; A. saponaria from November through August. In both species, the anthesis lasted 48 h., the anther dehiscence coincided with the opening of the perianth, the stigma receptivity was higher on the second day of anthesis and nectar production reached 0.34 ml/flower. In A. vera 228±77 yellow flowers on long scapes of 76±11 cm and with 1 - 3 shafts were observed; A. saponaria had 94 ± 33 orange flowers on more compact scapes with 3-5 shafts. The pollen/ovule ratio was 4 115.2 in A. vera and 3 247.1 in A. saponaria, thus they can be classified as compulsory xenogams. Frequent visits of Apis mellifera, Trigona sp., Poliste sp., Eumenes sp., Vespa sp., Leucippus fallax and Amazilia tobaci were recorded. Although both species showed a high production of pollen and nectar, and the presence of potential pollinators, by the end of the flowering period, fruits were only observed in A. saponaria, with a 12 % reproductive efficiency. Coupled with the stigmatic receptivity results and the preliminary intra and interspecific crossing experiments, this suggests the existence of protandry and self-incompatibility as reproductive barriers reducing endogamy in these species. Rev. Biol. Trop. 56 (3): 1109-1125. Epub 2008 September 30.

Se analizó la fenología reproductiva y aspectos anatómicos florales en dos especies de Aloe conservadas en condiciones de vivero en el oriente de Venezuela, de septiembre 2001 a septiembre 2002. A. vera floreció de diciembre a mayo; A. saponaria de noviembre a agosto. En ambas especies, la antesis duró 48 h, la dehiscencia de la antera coincidió con la apertura del perianto, la receptividad del estigma fue mayor al segundo días de antesis y la producción de néctar alcanzó 0.34 ml/flor. En A. vera se observaron 228±77 flores de color amarillo sobre escapos extensos de 76±11 cm y 1-3 ejes; a diferencia de A. saponaria con 94±33 flores anaranjadas en escapos más compactos de 62±5 cm y 3-5 ejes. La relación polen/óvulo fue de 4 151.2 en A. vera y 3 247.1 en A. saponaria, clasificándose como xenógamas obligatorias. Se registraron visitas frecuentes de Apis mellifera, Trigona sp., Poliste sp., Eumenes sp., Vespa sp., Leucippus fallax y Amazilia tobaci. A pesar de que ambas especies manifestaron una elevada producción de polen, néctar y la presencia de potenciales polinizadores, al término de la floración, sólo se observaron frutos en A. saponaria, con una eficiencia reproductiva de 12%, lo cual, junto a los resultados de la receptividad estigmática y los ensayos preliminares de cruzamientos intra e interespecíficos, sugiere la existencia de protandría y autoincompatibilidad como barreras reproductivas que reducen la endogamia en estas especies.

Nota sobre a ocorrência de pétalas reduzidas em espécies de Polygala L. subgênero Hebeclada (Chodat) Blake (Polygalaceae) da Região Sul do Brasil / A note about the occurrence of reduced petals in species of Polygala L. subgenus Hebeclada (Chodat) Blake (Polygalaceae) from southern Brazil

O número de pétalas é um dos caracteres taxonômicos utilizados na circunscrição dos subgêneros de Polygala. Até o momento, a corola das espécies do subgênero Hebeclada era descrita apresentando apenas três pétalas, uma inferior e central, denominada carena e duas laterais superiores aderidas à bainha estaminal. Dentre os 12 subgêneros de Polygala, a ocorrência de corola trímera era registrada apenas nos subgêneros Polygala e Hebeclada. O presente trabalho trata da ocorrência inédita de corola pentâmera em espécies de Polygala do subgênero Hebeclada, apresentando fotografias e comentários adicionais sobre estas pétalas reduzidas, não referidas para o subgênero até o momento.

The number of petals in the flowers is a taxonomic character used in the circumscription of the subgenera of Polygala. Until now, the corolla of the species of subgenus Hebeclada was described having tree petals, one inferior and central called carina and two lateral adnated to the staminal sheath. Among the 12 subgenera of Polygala, the occurrence of trimerous corolla was only known in subgenera Polygala and Hebeclada. This paper presents the discovery of pentamerous corolla in species of Polygala subgenus Hebeclada, presenting photos and additional comments about this reduced petals unknown for the subgenus until this moment.

Homeotic-like modification of stamens to petals is associated with aberrant mitochondrial gene expression in cytoplasmic male sterile Ogura Brassica juncea.

We have previously reported correction of severe leaf chlorosis in the cytoplasmic male sterile Ogura (also called Ogu) Brassica juncea line carrying Ogura cytoplasm by plastid substitution via protoplast fusion. Two cybrids obtained from the fusion experiment, Og1 and Og2, were green and carried the plastid genome of B. juncea cv. RLM198. While Og1 displayed normal flower morphology comparable to that of its euplasmic B. juncea counterpart except for sterile anthers, Og2 retained homeotic-like floral modification of stamens to petal-like structures and several other floral deformities observed in the chlorotic (Ogu) B. juncea cv. RLM198 (or OgRLM). With respect to the mitochondrial genome, Og1 showed 81% genetic similarity to the fertile cultivar RLM while Og2 showed 93% similarity to OgRLM. In spite of recombination and rearrangements in the mitochondrial genomes in the cybrids, expression patterns of 10 out of 11 mitochondrial genes were similar in all the three CMS lines; the only exception was atp6, whose expression was altered. While Og1 showed normal atp6 transcript similar to that in RLM, in Og2 and OgRLM weak expression of a longer transcript was detected. These results suggest that the homeotic-like changes in floral patterning leading to petaloid stamens in Og2 and OgRLM may be associated with aberrant mitochondrial gene expression.

Effect of nectar secretion rate on pollination success of Passiflora coccinea (Passifloraceae) in the Central Amazon

Estudamos a polinização de Passiflora coccinea por beija-flores Phaethornis superciliosus na Amazônia Central, Brasil. Nossa hipótese é que maiores taxas de secreção de néctar (TSN) aumentam o sucesso da polinização de flores individuais através do comportamento de visitas de Ph. superciliosus. Para flores controladas, a TSN foi uma função positiva do diâmetro da base da flor (DBF). O número total de visitas de Ph. superciliosus por flor foi uma função positiva do DBF. Adicionalmente, a deposição de pólen sobre os estigmas aumentou com o aumento do número acumulado de visitas de Ph. superciliosus. Nossos resultados indicam que flores maiores de P. coccinea secretam néctar em taxas mais altas, são visitadas mais vezes pelos beija-flores, e apresentam maior sucesso de polinização. Este parece ser o primeiro estudo não-manipulativo que descreve este efeito da TSN sobre o sucesso de polinização de flores individuais na natureza.