Leaf morphological strategies of seedlings and saplings of Rhizophora mangle (Rhizophoraceae), Laguncularia racemosa (Combretaceae) and Avicennia schaueriana (Acanthaceae) from Southern Brazil

AbstractThe initial phase of a plant life cycle is a short and critical period, when individuals are more vulnerable to environmental factors. The morphological and anatomical study of seedlings and saplings leaf type enables the understanding of species strategies of fundamental importance in their establishment and survival. The objective of this study was to analyze the structure of seedlings and saplings leaf types of three mangrove species, Avicennia schaueriana, Laguncularia racemosa, Rhizophora mangle, to understand their early life adaptive strategies to the environment. A total of 30 fully expanded cotyledons (A. schaueriana and L. racemosa), 30 leaves of seedlings, and 30 leaves of saplings of each species were collected from a mangrove area in Guaratuba Bay, Paraná State, Brazil. Following standard methods, samples were prepared for morphological (leaf dry mass, density, thickness) and anatomical analysis (epidermis and sub-epidermal layers, stomata types, density of salt secretion glands, palisade and spongy parenchyma thickness). To compare leaf types among species one-way ANOVA and Principal Component Analysis were used, while Cluster Analysis evaluated differences between the species. We observed significant structural differences among species leaf types. A. schaueriana showed the thickest cotyledons, while L. racemosa presented a dorsiventral structure. Higher values of the specific leaf area were observed for seedlings leaves of A. schaueriana, cotyledons of L. racemosa and saplings leaves of A. schaueriana and R. mangle. Leaf density was similar to cotyledons and seedlings leaves in A. schaueriana and L. racemosa, while R. mangle had seedlings leaves denser than saplings. A. schaueriana and R. mangle showed hypostomatic leaves, while L. racemosa amphistomatic; besides, A. chaueriana showed diacytic stomata, while L. racemosa anomocytic, and R. mangle ciclocytic. Seedling leaves were thicker in R. mangle (535 µm) and L.racemosa (520 µm) than in A. schaueriana (470.3 µm); while saplings leaves were thicker in L. racemosa (568.3 µm) than in A. schaueriana seedlings (512.4 µm) and R. mangle (514.6 µm). Besides, seedlings leaves palisade parenchyma showed increasing thickness in L. racemosa (119.2 µm) <A. schaueriana (155.5 µm) <R. mangle (175.4 µm); while in saplings leaves as follows R. mangle (128.4 µm) <A. schaueriana (183.4 µm) <L. racemosa (193.9 µm). Similarly, seedlings leaves spongy parenchyma thickness values were as follows A. schaueriana (182.6 µm) = R. mangle (192.8 µm) <L. racemosa (354.4 µm); while in saplings were A. schaueriana (182.6 µm) = R. mangle (187.3 µm) <L. racemosa (331.3 µm). The analyzed traits, in different combinations, represent morphological adjustments of leaf types to reduce water loss, eliminate salt excess, increase the absorption of light, allowing a higher efficiency on the maintenance of physiological processes in this initial growth stage. Rev. Biol. Trop. 64 (1): 305-317. Epub 2016 March 01.

ResumenLa fase inicial del ciclo de vida de una planta es un período corto y crítico, cuando los individuos son más vulnerables a factores ambientales. El estudio morfológico y anatómico del tipo de hojas de las plántulas y árboles pequeños, permite la comprensión de las estrategias de las especies, que es de importancia fundamental para su establecimiento y supervivencia. El objetivo de este estudio fue analizar la estructura de los tipos de hojas de las plántulas y árboles pequeños de tres especies de mangle: Avicennia schaueriana, Laguncularia racemosa y Rhizophora mangle, para entender sus estrategias de vida tempranas de adaptación al ambiente. Un total de 30 cotiledones completamente abiertos (A. schaueriana y L. racemosa), 30 hojas de plántulas, y 30 hojas de árboles pequeños de cada especie se recolectaron en una área de manglar en Bahía Guaratuba, Estado de Paraná, Brasil. Siguiendo los métodos estándares, se prepararon muestras para análisis morfológicos (biomasa de hoja seca, densidad y espesor) y anatómicos (epidermis y capas sub-epidérmicas, tipos de estomas, densidad de glándulas secretoras de sal y grosor del parénquima empalizado y del esponjoso). Para comparar los tipos de hojas entre las especies se utilizaron un modelo lineal y Análisis de Componentes Principales, mientras que un análisis de conglomerados evaluó las diferencias entre las especies. Observamos diferencias estructurales significativas entre tipos de hoja en las especies. A.schaueriana mostró cotiledones más gruesos, mientras que L. racemosa presenta una estructura dorsiventral. Se observaron valores más altos del área foliar específica para las hojas de las plántulas de A. schaueriana, cotiledones de L. racemosa y hojas de árboles pequeños de A. schaueriana y R. mangle. La densidad de la hoja fue similar a la de los cotiledones y hojas de plántulas de A. schaueriana y L. racemosa, mientras que R. mangle tenía las hojas de las plántulas más gruesas que los árboles pequeños. A. schaueriana y R. mangle mostraron hojas hipostomáticas; L. racemosa anfiestomáticas; por otro lado A. chaueriana mostró estomas diacíticos, L. racemosa anomocíticos y R. mangle ciclocíticos. Las hojas de las plántulas eran más gruesas en R. mangle (535 micras) y L. racemosa (520 micras) que en A. schaueriana (470.3 m); mientras que las hojas de las plántulas eran más gruesas en L. racemosa (568.3 m) que en A. schaueriana (512.4 micras) y R. mangle (514.6 m). Además el parénquima empalizado de las plántulas mostró un aumento de espesor en L. racemosa (119.2 m) <A. schaueriana (155.5 m) <R. mangle (175.4 m); mientras que en las hojas de los árboles pequeños fue de siguiente manera: R. mangle (128.4 m) <A. schaueriana (183.4 m) <L. racemosa (193.9 m). Del mismo modo, en las hojas de las plántulas los valores del espesor del parénquima esponjoso fueron: A. schaueriana (182.6 m) = R. mangle (192.8 m) <L. racemosa (354.4 m); mientras que en los árboles pequeños: A. schaueriana (182.6 m) = R. mangle (187.3 m) <L. racemosa (331.3 m). Los rasgos analizados, en diferentes combinaciones, representan ajustes morfológicos de tipos de hojas para reducir la pérdida de agua, eliminar el exceso de sal, aumentar la absorción de la luz, lo que permite una mayor eficiencia en el mantenimiento de los procesos fisiológicos en esta etapa de crecimiento inicial.

Seed dispersal and predation of Buchenavia tomentosa Eichler (Combretaceae) in a Cerrado sensu stricto, midwest Brazil / Dispersão e predação de sementes de Buchenavia tomentosa Eichler (Combretaceae) em Cerrado sentido restrito, centro-oeste do Brasil

Abstract The ecology of seed dispersal is critical to understand the patterns of distribution and abundance of plant species. We investigated seed dispersal aspects associated with the high abundance of Buchenavia tomentosa in the Serra Azul State Park (PESA). We estimated fruit production and conducted fruit removal experiments. We carried out diurnal and nocturnal observations on frugivory as well as germination tests. Fruiting occurred in the dry season and totaled 1,365,015 ± 762,670 fruits.ha–1. B. tomentosa fruits were utilized by eight animal species. The lowland tapir (Tapirus terrestris) was considered the main seed disperser. Leafcutter ants (Atta laevigata and Atta sexdens) participated in the seed cleaning and occasionally dispersed seeds. The beetle Amblycerus insuturatus, blue-and-yellow macaw (Ara ararauna) and red-and-green macaw (Ara chloropterus) were considered pre-dispersal seed predators. The seeds manually cleaned presented higher germination rate (100%) and speed index (4.2 seeds.d–1) than that of seeds with pulp. Germination of seeds found in tapirs’feces was 40%, while for the seeds without pulp it was 25%. The high abundance of B. tomentosa in the cerrado of PESA may be due to massive fruit production, low rates of seed predation, and efficient seed dispersal by tapirs, occurring before the rains which promote germination and recruitment of this species.

Resumo A ecologia da dispersão de sementes é importante para entender a distribuição e abundância das espécies vegetais. Investigamos os fatores relacionados à elevada abundância de B. tomentosa no Parque Estadual da Serra Azul (PESA) avaliando aspectos da dispersão de sementes. Estimamos a produção de frutos e realizamos experimentos de remoção de frutos. Fizemos observações diurnas e noturnas sobre a frugivoria e testamos a germinação de sementes. A frutificação ocorreu na estação seca totalizando 1.365.015 ± 762.670 frutos.ha–1. Oito espécies de animais utilizaram frutos de B. tomentosa. Anta (Tapirus terrestris) foi considerada o principal dispersor. Saúvas (Atta laevigata e Atta sexdens) participaram na limpeza das sementes e como dispersores ocasionais. O besouro Amblycerus insuturatus, arara-canindé (Ara ararauna) e arara-vermelha (Ara chloropterus) foram considerados predadores pré-dispersão. Sementes despolpadas manualmente tiveram maior percentual (100%) e velocidade de geminação (4,2 sementes.dia–1) que aquelas com polpa. Sementes encontradas nas fezes de antas apresentaram 40,0% de germinação e sementes sem polpa 25,0%. A massiva produção de frutos, baixa taxa de predação de sementes, além da eficiente dispersão por antas, antes da estação chuvosa, favorecem o recrutamento e são fatores que contribuem de forma decisiva para a elevada abundância de B. tomentosa observada no cerrado do PESA.

Estructura y dinámica del manglar del delta del río Ranchería, Caribe colombiano / Structure and dynamics of the mangrove forest in the Rancheria River delta, Colombian Caribbean

We registered seedling survival and biomass increase for Rhizophora mangle L., Avicennia germinans L. and Laguncularia racemosa (L.) Gaertn. f, main mangrove species in the Rancheria River delta, Colombia. Only seedlings of R. mangle were found to survive. We also measured maximum rate of litterfall. We estimated annual litterfall through interpolation within an exponential regression performed with maximum and annual litterfall data published in other sources; the value of annual litterfall for the area was estimated to be 12.9 mgha(-1)y(-1). We found a 7.4 mgha(-1)y(-1)(-1) increase in biomass. Litterfall constitutes the larger fraction of the 20.2 mgha(-1)y(-1) productivity of this mangrove. We believe this is a very high value for a forest under unfavorable natural and human conditions, such as high seasonality and continuous use of the forest to feed goats and sheep. We consider that the high productivity is a response to both natural and anthropogenic stress.