Herbivory and leaf expansion of Cyathea phalerata Mart (Cyatheaceae) in subtropical Atlantic Forest, southern Brazil / Herbivoria e expansão foliar de Cyathea phalerata Mart. (Cyatheaceae) em Floresta Atlântica subtropical no sul do Brasil

Herbivory is an interaction with great impact on plant communities since relationships between herbivores and plants are fundamental to the distribution and abundance of species over time and space. The aim of this study was to monitor the rate of leaf expansion in the tree fern Cyathea phalerata and evaluate the damage caused by herbivores to leaves of different ages and whether such damage is related to temperature and precipitation. The study was performed in a subtropical Atlantic Forest fragment located in the municipality of Caraá, in the northeast hillside of Rio Grande do Sul state, in southern Brazil. We monitored 24 mature individuals of C. phalerata with croziers in a population of approximately 50 plants. Leaf expansion rate, percentage of damaged leaves and leaf blade consumption rate by herbivory were calculated. Monthly means for temperature and accumulated rainfall were calculated from daily data. Croziers of C. phalerata were found to expand rapidly during the first and second months after emergence (3.98 cm day-¹; 2.91 cm day-¹, respectively). Damage caused by herbivory was observed in all of the monitored leaves, but none of the plants experienced complete defoliation. The highest percentage (57%) of damaged leaves was recorded at 60 days of monitoring, and also the highest monthly consumption rate of the blade (6.04%) occurred with young, newly-expanded leaves, while this rate remained between 1.50 and 2.21% for mature leaves. Rates of monthly leaf consumption and damaged leaves showed positive and strong relationship with each other and with temperature. The rapid leaf expansion observed for C. phalerata can be considered a phenological strategy to reduce damage to young leaves by shortening the developmental period and accelerating the increase of defenses in mature leaves.

A herbivoria é uma interação de grande impacto sobre as comunidades de plantas, uma vez que as relações entre herbívoros e plantas são fundamentais para a distribuição e abundância das espécies ao longo do tempo e do espaço. O objetivo deste estudo foi monitorar a taxa de expansão foliar da samambaia arborescente Cyathea phalerata e avaliar o dano causado por herbívoros a folhas de diferentes idades, assim como verificar se este dano está relacionado à temperatura e à precipitação. O estudo foi realizado em um fragmento de Floresta Atlântica subtropical localizado no município de Caraá, na encosta nordeste do estado do Rio Grande do Sul, no sul do Brasil. Nós monitoramos 24 indivíduos maduros de C. phalerata com báculos em uma população de aproximadamente 50 plantas. A taxa de expansão foliar, a porcentagem de folhas danificadas e a taxa de consumo da lâmina foliar por herbivoria foram calculados. Médias mensais de temperatura e precipitação acumulada foram calculadas a partir de dados diários. Báculos de C. phalerata expandiram rapidamente durante o primeiro e o segundo mês após emergência (3,98 cm dia-¹; 2,91 cm dia-¹, respectivamente). O dano causado por herbivoria foi observado em todas as folhas monitoradas, mas nenhuma das plantas sofreu desfolhação completa. A maior porcentagem (57%) de folhas danificadas foi registrada aos 60 dias de monitoramento, e também a maior taxa de consumo mensal (6,04%) ocorreu em folhas jovens, recém expandidas, enquanto esta taxa permaneceu entre 1,50 e 2,21% em folhas maduras. As taxas mensais de consumo da lâmina foliar e de folhas danificadas mostraram relação positiva e forte entre si e com a temperatura. A rápida expansão foliar observada em C. phalerata pode ser considerada uma estratégia fenológica para reduzir o dano a folhas jovens, abreviando o período de desenvolvimento e acelerando o aumento das defesas em folhas maduras.

Herbivory and leaf expansion of Cyathea phalerata Mart. (Cyatheaceae) in subtropical Atlantic Forest, southern Brazil

Abstract Herbivory is an interaction with great impact on plant communities since relationships between herbivores and plants are fundamental to the distribution and abundance of species over time and space. The aim of this study was to monitor the rate of leaf expansion in the tree fern Cyathea phalerata and evaluate the damage caused by herbivores to leaves of different ages and whether such damage is related to temperature and precipitation. The study was performed in a subtropical Atlantic Forest fragment located in the municipality of Caraá, in the northeast hillside of Rio Grande do Sul state, in southern Brazil. We monitored 24 mature individuals of C. phalerata with croziers in a population of approximately 50 plants. Leaf expansion rate, percentage of damaged leaves and leaf blade consumption rate by herbivory were calculated. Monthly means for temperature and accumulated rainfall were calculated from daily data. Croziers of C. phalerata were found to expand rapidly during the first and second months after emergence (3.98 cm day-1; 2.91 cm day-1, respectively). Damage caused by herbivory was observed in all of the monitored leaves, but none of the plants experienced complete defoliation. The highest percentage (57%) of damaged leaves was recorded at 60 days of monitoring, and also the highest monthly consumption rate of the blade (6.04%) occurred with young, newly-expanded leaves, while this rate remained between 1.50 and 2.21% for mature leaves. Rates of monthly leaf consumption and damaged leaves showed positive and strong relationship with each other and with temperature. The rapid leaf expansion observed for C. phalerata can be considered a phenological strategy to reduce damage to young leaves by shortening the developmental period and accelerating the increase of defenses in mature leaves.

Resumo A herbivoria é uma interação de grande impacto sobre as comunidades de plantas, uma vez que as relações entre herbívoros e plantas são fundamentais para a distribuição e abundância das espécies ao longo do tempo e do espaço. O objetivo deste estudo foi monitorar a taxa de expansão foliar da samambaia arborescente Cyathea phalerata e avaliar o dano causado por herbívoros a folhas de diferentes idades, assim como verificar se este dano está relacionado à temperatura e à precipitação. O estudo foi realizado em um fragmento de Floresta Atlântica subtropical localizado no município de Caraá, na encosta nordeste do estado do Rio Grande do Sul, no sul do Brasil. Nós monitoramos 24 indivíduos maduros de C. phalerata com báculos em uma população de aproximadamente 50 plantas. A taxa de expansão foliar, a porcentagem de folhas danificadas e a taxa de consumo da lâmina foliar por herbivoria foram calculados. Médias mensais de temperatura e precipitação acumulada foram calculadas a partir de dados diários. Báculos de C. phalerata expandiram rapidamente durante o primeiro e o segundo mês após emergência (3,98 cm dia-1; 2,91 cm dia-1, respectivamente). O dano causado por herbivoria foi observado em todas as folhas monitoradas, mas nenhuma das plantas sofreu desfolhação completa. A maior porcentagem (57%) de folhas danificadas foi registrada aos 60 dias de monitoramento, e também a maior taxa de consumo mensal (6,04%) ocorreu em folhas jovens, recém expandidas, enquanto esta taxa permaneceu entre 1,50 e 2,21% em folhas maduras. As taxas mensais de consumo da lâmina foliar e de folhas danificadas mostraram relação positiva e forte entre si e com a temperatura. A rápida expansão foliar observada em C. phalerata pode ser considerada uma estratégia fenológica para reduzir o dano a folhas jovens, abreviando o período de desenvolvimento e acelerando o aumento das defesas em folhas maduras.

Herbivory and leaf expansion of Cyathea phalerata Mart. (Cyatheaceae) in subtropical Atlantic Forest, southern Brazil / Herbivoria e expansão foliar de Cyathea phalerata Mart. (Cyatheaceae) em Floresta Atlântica subtropical no sul do Brasil

Abstract Herbivory is an interaction with great impact on plant communities since relationships between herbivores and plants are fundamental to the distribution and abundance of species over time and space. The aim of this study was to monitor the rate of leaf expansion in the tree fern Cyathea phalerata and evaluate the damage caused by herbivores to leaves of different ages and whether such damage is related to temperature and precipitation. The study was performed in a subtropical Atlantic Forest fragment located in the municipality of Caraá, in the northeast hillside of Rio Grande do Sul state, in southern Brazil. We monitored 24 mature individuals of C. phalerata with croziers in a population of approximately 50 plants. Leaf expansion rate, percentage of damaged leaves and leaf blade consumption rate by herbivory were calculated. Monthly means for temperature and accumulated rainfall were calculated from daily data. Croziers of C. phalerata were found to expand rapidly during the first and second months after emergence (3.98 cm day-1; 2.91 cm day-1, respectively). Damage caused by herbivory was observed in all of the monitored leaves, but none of the plants experienced complete defoliation. The highest percentage (57%) of damaged leaves was recorded at 60 days of monitoring, and also the highest monthly consumption rate of the blade (6.04%) occurred with young, newly-expanded leaves, while this rate remained between 1.50 and 2.21% for mature leaves. Rates of monthly leaf consumption and damaged leaves showed positive and strong relationship with each other and with temperature. The rapid leaf expansion observed for C. phalerata can be considered a phenological strategy to reduce damage to young leaves by shortening the developmental period and accelerating the increase of defenses in mature leaves.

Resumo A herbivoria é uma interação de grande impacto sobre as comunidades de plantas, uma vez que as relações entre herbívoros e plantas são fundamentais para a distribuição e abundância das espécies ao longo do tempo e do espaço. O objetivo deste estudo foi monitorar a taxa de expansão foliar da samambaia arborescente Cyathea phalerata e avaliar o dano causado por herbívoros a folhas de diferentes idades, assim como verificar se este dano está relacionado à temperatura e à precipitação. O estudo foi realizado em um fragmento de Floresta Atlântica subtropical localizado no município de Caraá, na encosta nordeste do estado do Rio Grande do Sul, no sul do Brasil. Nós monitoramos 24 indivíduos maduros de C. phalerata com báculos em uma população de aproximadamente 50 plantas. A taxa de expansão foliar, a porcentagem de folhas danificadas e a taxa de consumo da lâmina foliar por herbivoria foram calculados. Médias mensais de temperatura e precipitação acumulada foram calculadas a partir de dados diários. Báculos de C. phalerata expandiram rapidamente durante o primeiro e o segundo mês após emergência (3,98 cm dia-1; 2,91 cm dia-1, respectivamente). O dano causado por herbivoria foi observado em todas as folhas monitoradas, mas nenhuma das plantas sofreu desfolhação completa. A maior porcentagem (57%) de folhas danificadas foi registrada aos 60 dias de monitoramento, e também a maior taxa de consumo mensal (6,04%) ocorreu em folhas jovens, recém expandidas, enquanto esta taxa permaneceu entre 1,50 e 2,21% em folhas maduras. As taxas mensais de consumo da lâmina foliar e de folhas danificadas mostraram relação positiva e forte entre si e com a temperatura. A rápida expansão foliar observada em C. phalerata pode ser considerada uma estratégia fenológica para reduzir o dano a folhas jovens, abreviando o período de desenvolvimento e acelerando o aumento das defesas em folhas maduras.

Phenology of the endangered palm Ceroxylon quindiuense (Arecaceae) along an altitudinal gradient in Colombia / Fenología de la palma en peligro Ceroxylon quindiuense (Arecaceae) a lo largo de un gradiente altitudinal en Colombia

Abstract Introduction: Understanding the phenology of plant populations is vital for their conservation and management. We studied the vegetative and reproductive phenology of the endangered palm Ceroxylon quindiuense along an altitudinal gradient in the Central Cordillera of Colombia. Objective: We describe the leaf production rate, and flowering and fruiting cycles, and calculate food offer for the fauna, as a tool for the proper management of the palm. Methods: At each sampling site (2 400, 2 600, 2 800, 3 000 m.a.s.l.), we marked 40 adult individuals (20 pistillate, 20 staminate), which we followed bimonthly for 24 months. We studied leaf production by counting fallen leaves. We followed flower and fruit production through observations with binoculars and photographs. Results: Each adult individual produced, on average, one leaf every 69 days. Although isolated individuals flowered throughout the year, most palms flowered synchronously at each elevation in October 2016-August 2017 and in August 2018-February 2019 and had ripe fruits 7-13 months later. Flowering started at 2 600 m, followed by 2 800 and 3 000 m. Palms at 2 400 m, the lower limit of the palm stands in the area, showed a singular behavior, with scarce flower and fruit production, some individuals that changed sex, and a higher proportion of pistillate palms. Each palm produced 1-11 (x̄ = 5.3, SD = 2.2) inflorescences and 1-10 (x̄ = 5.3, SD = 2.2) infructescences. The average number of fruits per infructescence was 4 465 (SD = 1 488). With an estimated population of adult palms between 256 000 and 600 000 and an overall ratio of pistillate: staminate individuals 1:1 or 1:2, total fruit production in the area during each fruiting period is estimated as 2.0-7.1 billion fruits. Conclusions: The huge number of flowers and fruits and their gradual availability along the altitudinal gradient have a major impact on the spatial and temporal distribution of food offer for fauna associated with the palm.

Resumen Introducción: Comprender la fenología de las poblaciones de plantas es vital para su conservación y manejo. Estudiamos la fenología vegetativa y reproductiva de la palma amenazada Ceroxylon quindiuense a lo largo de un gradiente altitudinal en la Cordillera Central de Colombia. Objetivo: Describimos la tasa de producción de hojas, los ciclos de floración y fructificación, y calculamos la oferta alimentaria para la fauna, como una herramienta para el adecuado manejo de la palma. Métodos: En cada sitio de muestreo (2 400, 2 600, 2 800, 3 000 m.s.n.m.), marcamos 40 individuos adultos (20 pistilados, 20 estaminados), que seguimos bimestralmente durante 24 meses. Estudiamos la producción de hojas contando las caídas al suelo. Seguimos la producción de flores y frutos a través de observaciones con binoculares y fotografías. Resultados: Cada individuo adulto produjo, en promedio, una hoja cada 69 días. Aunque los individuos aislados florecieron durante todo el año, la mayoría de las palmas florecieron sincrónicamente en cada elevación entre octubre 2016 y agosto 2017 y de agosto 2018 a febrero 2019 y tuvieron frutos maduros entre 7-13 meses después. La floración comenzó a los 2 600 m, seguida de los 2 800 y los 3 000 m. Las palmas a 2 400 m, límite inferior de los palmares de la zona, mostraron un comportamiento singular, con escasa producción de flores y frutos, varios individuos que cambiaron de sexo y una mayor proporción de palmas pistiladas. Cada palma produjo 1-11 (x̄ = 5.3, SD = 2.2) inflorescencias y 1-10 (x̄ = 5.3, SD = 2.2) infrutescencias. El número promedio de frutos por infrutescencia fue de 4 465. Con una población estimada de palmas adultas entre 256 000 y 600 000 y una proporción total de individuos pistilados: estaminados 1:1 o 1:2, la producción total de frutos en el área durante cada período de fructificación se estima en 2.0-7.1 mil millones de frutos. Conclusiones: La gran cantidad de flores y frutos y su progresiva disponibilidad a lo largo del gradiente tienen un impacto importante en la distribución espacial y temporal de la oferta de alimento para la fauna asociada a la palma.

The stands of Copernicia tectorum (Arecaceae) in the Caribbean lowlands of Colombia: a managed pioneer palm facing river dynamics / Poblaciones de Copernicia tectorum (Arecaceae) en las tierras bajas del Caribe de Colombia: una palma pionera expuesta a la dinámica de un río

Copernicia tectorum is a palm that grows in large populations on seasonally flooded savannas in the Caribbean region of Colombia, where its stems and expanded leaves are used in construction, and its unexpanded leaves are used to make handicrafts. We studied abundance and population structure in 34 plots of 20×10m (0.68 ha) of three localities (Plato, Córdoba and Magangué) at the Mompox Depression, an inner delta formed by the confluence of four large rivers. We recorded growth and mortality of 164 palms of different size classes over 13 months, and additionally we estimated seedling and juvenile mortality in two 10x10m plots within the same sites. Data analysis using Kolgomorov-Smirnov (KS), Kruskal-Wallis and Mann-Whitney tests were made on Statgraphics Plus and SPSS. We related the structure and dynamics of the palm stands to flood intensity. Copernicia tectorum has the fastest leaf production rate recorded for any palm (19-23 leaves/year in subadults and adults), and a short life span of ca. 46 years. The abundance, density and high leaf production rate of this palm offer a great potential for the sustainable use of its unexpanded leaves (especially at Plato, where there are ca. 480ha of palm stands with 300-1 000 individuals/ha), as leaf harvest from subadult and adult individuals does not appear to affect population structure. The palm is a pioneer of the flood plain, and the palm stands at particular sites appear to wax and wane following sedimentation dynamics, in which the species plays in important role. Current population structure and the scarce recruitment of juveniles at the study site suggest that the population is waning in that site. Management actions should be taken to reestablish natural river dynamics at the lagoon complex where the palm grows. Appropriate management of palm stands may contribute to minimize any negative effects of river dynamics.

La palma Copernicia tectorum forma grandes poblaciones en sabanas estacionalmente inundables de la región Caribe de Colombia, donde sus tallos y hojas se utilizan en construcción y sus cogollos para la elaboración de artesanías. Estudiamos la abundancia y la estructura poblacional en 34 parcelas de 20×10m (0.68 ha), en tres localidades (Plato, Córdoba y Magangué) en la Depresión Momposina, un delta interior formado por la confluencia de cuatro grandes ríos. Registramos el crecimiento y la mortalidad de 164 palmas de diferentes clases de tamaño durante 13 meses y estimamos además la mortalidad en plántulas y juveniles en dos parcelas de 10x10m en los mismos sitios. Los análisis se realizaron con las pruebas de Kolgomorov-Smirnov (KS), Kruskal-Wallis y Mann-Whitney en Statgraphics Plus and SPSS. Relacionamos la estructura y dinámica de los palmares con la intensidad de la inundación. Copernicia tectorum tiene la tasa de producción de hojas más rápida registrada hasta ahora para cualquier palma (19-23 hojas por año en subadultos y adultos) y una vida corta de ca. 46 años. La abundancia, densidad y alta tasa de producción de hojas de la palma ofrecen un gran potencial para el uso sostenible de sus cogollos (especialmente en Plato, donde hay unas 480 ha de palmares con 300-1 000 individuos/ha), ya que la cosecha de hojas de palmas adultas y subadultas, parece no afectar la estructura de la población. La palma es una pionera de las llanuras inundables, y los palmares en un sitio particular crecen y se reducen siguiendo la dinámica de sedimentación, en la cual la especie juega un papel importante. La estructura actual de las poblaciones y el escaso reclutamiento en las clases juveniles en el sitio de estudio sugieren que allí la población se está reduciendo. Se deben emprender acciones de manejo que apunten a restablecer la dinámica fluvial natural en los complejos cenagosos en los que crece la palma. A su vez, el manejo apropiado de los palmares ...

Dinámica del daño foliar en plántulas de Drimys granadensis (Winteraceae) y Clusia multiflora (Clusiaceae) en el bosque altoandino de la Cordillera Oriental colombiana

Leaf damage dynamics of Drimys granadensis (Winteraceae) and Clusia multiflora (Clusiaceae) seedlings in the High-Andean Forest. Because of their slow growth, shade-tolerant species remain exposed to foliar damage long before reproductive age. Foliar damage can be controlled through synchronous leaf production and specialized phytochemical strategies. in this study, the dynamics of foliar damage in different cohorts of seedlings were evaluated to determine if the High-Andean species, Clusia multiflora (Clusiaceae) and Drimys granadensis (Winteraceae) appeal to the synchronous leaf production for controlling the herbivory and pathogenical damage, and to establish the possible relations between the healthy state, growth and mortality of seedlings. Since a recently-emerged seedling could not supply the physiological costs of specialized strategies, we expected the highest synchronous leaf production in the youngest individuals, and differences between cohorts in foliar damage. Four variables were measured to evaluate the health state along time, in three cohorts of seedlings: Proportion of predated leaves (Pp), proportion of healthy leaves (Ps), proportion of leaves with symptoms of disease or punctual damage (Pe) and growing (% increment of height). Both species showed significant differences between periods in the proportion of healthy leaves, but there was not an effect of time-cohort interaction, therefore synchronic production of leaves was not a strategy more used for any particular cohort. Foliar damage oscillated along time, fact that can be explained by the pulses of leaf production. However, this strategy had little efficiency to control the pathogen attack. in general, the unified behavior of all variables was affected by cohort, time, species an all the different interactions. Association between growing and foliar damage were conditioned by climate. The highest mortality ocurred during the dry season, and one fourth of the D. granadensis deaths were caused by the joint action of herbivory and drought. Results suggest that in spite of the synchronous leaf production in all cohorts, this strategy quickly loses the efficiency for controlling foliar damage, because the means of Ps were below of 0.25 at the end of the monitoring period. The differences between species in the dynamics of health state should explain partially their coexistence in the High-Andean forest. Rev. Biol. Trop. 56 (3): 1087-1100. Epub 2008 September 30.

Además de las estrategias fitoquímicas especializadas, las plantas pueden usar la producción sincrónica de hojas como un método de saciado de patógenos y herbívoros. Con el fin de determinar si las especies del bosque altoandino colombiano Clusia multiflora (Clusiaceae) y Drimys granadensis (Winteraceae) recurren a la producción sincrónica de hojas para controlar los efectos de la herbivoría y el ataque microbiano, y establecer que relación existe entre el estado de salud, el crecimiento y la mortalidad de plántulas, se evaluó la dinámica del daño foliar sobre diferentes cohortes. Dado que una plántula tolerante a la sombra recientemente emergida no puede suplir los costos fisiológicos de una estrategia especializada, se esperaba una alta sincronía en la producción de hojas en las plántulas más jóvenes, y diferencias en el daño foliar entre cohortes. Se midieron cuatro variables que evaluaban el estado de salud a lo largo del tiempo, en tres cohortes de plántulas: Proporción de hojas predadas, proporción de hojas sanas, proporción de hojas enfermas o con daño puntual y crecimiento. Ambas especies mostraron diferencias significativas entre épocas, en la proporción de hojas sanas; pero no hubo un efecto de la interacción tiempo-cohorte, por lo tanto la producción sincrónica de hojas no fue una estrategia más usada por alguna cohorte en particular. El daño foliar osciló a través del tiempo, lo cual puede ser explicado por los pulsos en la producción de hojas. Sin embargo, ésta estrategia tuvo poca eficiencia para controlar el ataque por patógenos. En general, el comportamiento unificado de todas las variables fue afectado por la cohorte, el tiempo, la especie y todas las diferentes interacciones. La relación entre crecimiento y daño foliar fue condicionado por el clima. La mayor mortalidad se dio durante la estación seca, y un cuarto de las muertes en D. granadensis fueron causadas por la acción conjunta de herbivoría y sequía. Los resultados sugieren que a pesar de la producción sincrónica de hojas en todas las cohortes, fue poca la eficiencia para controlar el daño foliar, ya que las medias de Ps estuvieron por debajo de 0.25 al final del seguimiento. Las diferencias en la dinámica del estado de salud entre especies podrían explicar parcialmente la coexistencia en el bosque altoandino.