What triggers phenological events in plants under seasonal environments? A study with phylogenetically related plant species in sympatry

Abstract In this study, we evaluated variation in vegetative and reproductive phenological events of four phylogenetically related plant species subjected to a seasonal environment. To this aim, we sampled 15 individuals of each plant species every fortnight for one year, between January and December of 2017. To assess when a given phenophase occurred more intensely in the population, the Fournier intensity index was used and the synchrony of individuals of the sample in a given phenological event was estimated using the activity index. The Rayleigh (Z) test was used to determine whether the phenological events have seasonal distribution. The relationship of abiotic factors (photoperiod, precipitation, relative humidity and temperature) with the intensity of phenophases was evaluated for each plant species using generalized linear models (GLMs). The phenophases of all plants showed a seasonal distribution pattern, as well as variation in synchrony of phenophases and specific sets of abiotic factors significantly influenced their phenophases. New leaves, for example, were produced throughout the seasons, with intense leaf fall in the dry season. Flowering periods, on the other hand, did not overlap. Indeed, species exhibited sequential flowering and asynchronous flowering among individuals. Our results suggest that the phenological patterns of four sympatric plant species are directly linked to climatic variables, but different abiotic factors affected different phenophases.

Resumo Neste estudo, avaliamos a variação nos eventos fenológicos vegetativos e reprodutivos de quatro espécies de plantas filogeneticamente relacionadas submetidas a um ambiente sazonal. Para isso, amostramos 15 indivíduos de cada espécie quinzenalmente, pelo período de um ano, entre janeiro e dezembro de 2017. Para avaliar quando uma determinada fenofase ocorreu de modo mais intenso na população foi utilizado o índice de intensidade de Fournier e a sincronia dos indivíduos da amostra em determinado evento fenológico foi estimada utilizando-se o índice de atividade. O teste de Rayleigh (Z) foi usado para determinar se os eventos fenológicos têm distribuição sazonal. A relação dos fatores abióticos (fotoperíodo, precipitação, temperatura e umidade relativa) com a intensidade das fenofases foi avaliada para cada espécie de planta usando modelos lineares generalizados (GLMs). As fenofases de todas as plantas apresentaram um padrão de distribuição sazonal, bem como variação na sincronia das fenofases e conjuntos específicos de fatores abióticos influenciaram significativamente suas fenofases. Folhas novas, por exemplo, foram produzidas ao longo das estações, com intensa queda de folhas na estação seca. Os períodos de floração, por outro lado, não se sobrepuseram. De fato, as espécies exibiram floração sequencial e floração assíncrona entre os indivíduos. Nossos resultados sugerem que os padrões fenológicos das quatro espécies de plantas simpátricas estão diretamente ligados às variáveis climáticas, mas diferentes fatores abióticos afetaram diferentes fenofases.

What triggers phenological events in plants under seasonal environments? A study with phylogenetically related plant species in sympatry / O que desencadeia eventos fenológicos em plantas em ambientes sazonais? Um estudo com espécies de plantas filogeneticamente relacionadas e em simpatria

In this study, we evaluated variation in vegetative and reproductive phenological events of four phylogenetically related plant species subjected to a seasonal environment. To this aim, we sampled 15 individuals of each plant species every fortnight for one year, between January and December of 2017. To assess when a given phenophase occurred more intensely in the population, the Fournier intensity index was used and the synchrony of individuals of the sample in a given phenological event was estimated using the activity index. The Rayleigh (Z) test was used to determine whether the phenological events have seasonal distribution. The relationship of abiotic factors (photoperiod, precipitation, relative humidity and temperature) with the intensity of phenophases was evaluated for each plant species using generalized linear models (GLMs). The phenophases of all plants showed a seasonal distribution pattern, as well as variation in synchrony of phenophases and specific sets of abiotic factors significantly influenced their phenophases. New leaves, for example, were produced throughout the seasons, with intense leaf fall in the dry season. Flowering periods, on the other hand, did not overlap. Indeed, species exhibited sequential flowering and asynchronous flowering among individuals. Our results suggest that the phenological patterns of four sympatric plant species are directly linked to climatic variables, but different abiotic factors affected different phenophases.

Neste estudo, avaliamos a variação nos eventos fenológicos vegetativos e reprodutivos de quatro espécies de plantas filogeneticamente relacionadas submetidas a um ambiente sazonal. Para isso, amostramos 15 indivíduos de cada espécie quinzenalmente, pelo período de um ano, entre janeiro e dezembro de 2017. Para avaliar quando uma determinada fenofase ocorreu de modo mais intenso na população foi utilizado o índice de intensidade de Fournier e a sincronia dos indivíduos da amostra em determinado evento fenológico foi estimada utilizando-se o índice de atividade. O teste de Rayleigh (Z) foi usado para determinar se os eventos fenológicos têm distribuição sazonal. A relação dos fatores abióticos (fotoperíodo, precipitação, temperatura e umidade relativa) com a intensidade das fenofases foi avaliada para cada espécie de planta usando modelos lineares generalizados (GLMs). As fenofases de todas as plantas apresentaram um padrão de distribuição sazonal, bem como variação na sincronia das fenofases e conjuntos específicos de fatores abióticos influenciaram significativamente suas fenofases. Folhas novas, por exemplo, foram produzidas ao longo das estações, com intensa queda de folhas na estação seca. Os períodos de floração, por outro lado, não se sobrepuseram. De fato, as espécies exibiram floração sequencial e floração assíncrona entre os indivíduos. Nossos resultados sugerem que os padrões fenológicos das quatro espécies de plantas simpátricas estão diretamente ligados às variáveis climáticas, mas diferentes fatores abióticos afetaram diferentes fenofases.

Effect of seasonality on the leaf phenology and physiology of Byrsonima species (Malpighiaceae)

Introduction: Defined seasonality in savanna species can stimulate physiological responses that maximize photosynthetic metabolism and productivity. However, those physiological responses are also linked to the phenological status of the whole plant, including leaf phenophases. Objective: To study how physiological traits influence phenophase timing among congeneric and co-occurring savanna species. Methods: We evaluated the leaf phenology and physiological traits of populations of Byrsonima intermedia, B. coccolobifolia, and B. verbascifolia. Physiological measurements were performed at the onset of the dry and rainy seasons and again late in the season. Results: B. intermedia and B. coccolobifolia were classified as brevideciduous and B. verbascifolia as evergreen. The maximum quantum yield for B. intermedia and B. coccolobifolia were lowest during the dry season. At the onset of the dry period, the highest chloroplastidic pigment levels were observed, which decreased as the season advanced, total chlorophyll/carotenoid ratios were lowest, and carotenoid contents were highest. We detected low starch content values at the start of the rainy season, coinciding with the resumption of plant growth. Two months into this season, the leaves were at their peak structural and functional maturity, with high water-soluble polysaccharide values and photosynthetic rates, and were storing large amounts of starch. Conclusions: Physiological and leaf phenological strategies of the Byrsonima species were related to drought resistance and acclimatization to the seasonality of savanna water resources. The oscillations of the parameters quantified during the year indicated a strong relationship with water seasonality and with the phenological status of the leaves.

Introducción: La marcada estacionalidad en las especies de sabana puede estimular respuestas fisiológicas que maximicen el metabolismo fotosintético y la productividad. Sin embargo, esas respuestas fisiológicas están vinculadas al estado fenológico de toda la planta, incluidas las fenofases de las hojas. Objetivo: Estudiar cómo los rasgos fisiológicos influyen en el tiempo de la fenofase entre especies de sabana congenéricas y concurrentes. Métodos: Evaluamos la fenología y características fisiológicas de poblaciones de Byrsonima intermedia, B. coccolobifolia y B. verbascifolia. Las mediciones fisiológicas se realizaron al inicio de la estación seca y lluviosa, y de nuevo al final de la estación. Resultados: B. intermedia y B. coccolobifolia se clasificaron como brevicaducifolias y B. verbascifolia como perennifolias. El rendimiento cuántico máximo para B. intermedia y B. coccolobifolia fueron más bajos durante la época seca. Al inicio del período seco, se observaron niveles de pigmentos cloroplastídicos más altos, aunque los niveles de clorofila disminuyeron a medida que avanzaba la estación seca, las proporciones clorofila/carotenoides totales fueron más bajas y los contenidos de carotenoides más altos. Detectamos valores bajos de contenido de almidón al inicio de la época lluviosa, que coincide con la reanudación del crecimiento de la planta. A dos meses de esta época, las hojas estaban en su máxima madurez estructural y funcional, con altos valores de polisacáridos solubles en agua y tasas fotosintéticas, y almacenaban grandes cantidades de almidón. Conclusiones: Las estrategias fisiológicas y fenológicas de las hojas de las especies de Byrsonima estaban relacionadas con la resistencia a la sequía y la aclimatación a la estacionalidad de los recursos hídricos de la sabana. Las oscilaciones de los parámetros cuantificados durante el año indicaron una fuerte relación con la estacionalidad hídrica y con los estados fenológicos de las hojas.

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.

Basic ecology of ant gardens in a dry-premontane transitional forest / Ecología básica de jardines de hormigas en un bosque transicional seco-premontano

Abstract Introduction: "Ant gardens'' are ant nests located at different heights on trees on which vascular epiphytic plants that have been transported and sown by ants have germinated. Although this mutualistic relationship has been studied in humid tropical ecosystems, information on other tropical and Colombian ecosystems is scarce. Objective: To characterize the distribution, building, diversity, demography and phenology of ant gardens in dry tropical forest. Methods: In January and February, 2018, we identified 170 ant gardens on a 100x5 m transect on the banks of the Quesada River, Colombia, and in adjacent secondary dry premontane transitional forest; we monitored changes, for gardens and plants, in 28 of them, every two weeks (March 2018-February 2019). Results: The gardens, built by Azteca ulei, were aggregated near water bodies; had 10 species of epiphytes and were on 13 tree species. Larger gardens had more epiphyte species. Some epiphytes had a bimodal phenological pattern. Less seedlings become established in the dry season, and less adults remain in the gardens. Garden characteristics benefit both epiphytes and ants. Conclusions: The gardens built by A. ulei have ecological characteristics that favor the germination, establishment, and reproduction of diverse epiphytes in this dry tropical ecosystem, including aggregation near water flows.

Resumen Introducción: Los "jardines de hormigas'' son nidos de hormigas en árboles, que se localizan a diferentes alturas, sobre los que germinan plantas epífitas vasculares que han sido transportadas y sembradas previamente por hormigas. Aunque esta relación mutualista ha sido estudiada en ecosistemas húmedos tropicales, la información en otros ecosistemas tropicales y en Colombia, es escasa. Objetivo: Caracterizar la distribución, construcción, diversidad, demografía y fenología de los hormigueros en bosque tropical seco. Métodos: En enero y febrero de 2018, identificamos 170 hormigueros en un transecto de 100x5 m a orillas del río Quesada, Colombia, y en el bosque secundario premontano seco de transición adyacente; monitoreamos cambios, para jardines y plantas, en 28 de ellos, cada dos semanas (marzo 2018-febrero 2019). Resultados: Los jardines, construidos por Azteca ulei, se agregaron cerca de cuerpos de agua; tenían 10 especies de epífitas y estaban sobre 13 especies arbóreas. Los jardines más grandes tenían más especies epífitas. Algunas epífitas tuvieron un patrón fenológico bimodal. Se establecen menos plántulas en la estación seca y quedan menos adultos en los jardines. Las características del jardín benefician tanto a las epífitas como a las hormigas. Conclusiones: Los jardines construidos por A. ulei tienen características ecológicas que favorecen la germinación, establecimiento y reproducción de diversas epífitas en este ecosistema tropical seco, incluyendo la agregación cerca de cursos de agua.

Disponibilidad y utilización de los recursos poliníferos por Apis mellifera (Hymenoptera: Apidae) en el este de las Yungas de Jujuy (Argentina) / Availability and use of polliniferous resources by Apis mellifera (Hymenoptera: Apidae) in the Eastern Yungas of Jujuy (Argentina)

Resumen Introducción: El conocimiento de la riqueza vegetal y la estacionalidad alrededor de los apiarios de Apis mellifera es una herramienta de planificación indispensable para los apicultores. Debe incluir la disponibilidad de recursos, las preferencias alimenticias y el comportamiento de búsqueda de alimento. Dicha información no está disponible para las Yungas argentinas, uno de los ecosistemas forestales más estacionales de América del Sur. Objetivo: Evaluar la disponibilidad de recursos tróficos a través de un calendario de floración y su relación con las cargas de polen de A. mellifera en las Yungas. Métodos: En El Fuerte, Jujuy, recolectamos muestras mensuales de septiembre a marzo (2014-2015 y 2015-2016) utilizando trampas de polen. Utilizamos técnicas estandarizadas para los análisis palinológicos e índices de asociación para el uso de recursos. Las fenofases fueron Inicio de floración, Plena floración y Fin de floración. Resultados: Se identificaron 47 especímenes botánicos a nivel de especie y 9 a nivel de género. En ambos períodos hubo una oferta moderada de flores al inicio de la primavera, representada igualmente por plantas arbustivas y herbáceas, con un pico de floración en noviembre. Posteriormente, hubo una caída en la disponibilidad, con un pico de floración nuevamente al final de la temporada. En cinco especies de plantas hubo una asociación de media a alta entre la especie vegetal disponible y la presencia de ésta en el espectro polínico de la muestra de polen corbicular recolectada (Vachellia aroma, Blepharocalyx salicifolius, Cantinoa sp., Vernonanthura sp. y Zanthoxylum coco). Conclusión: En esta región hay una oferta moderada de flores de plantas arbustivas y herbáceas a principios de la primavera, con un pico de floración en noviembre y al final de la temporada. Solo cinco, de casi 50 especies de plantas, muestran una asociación de disponibilidad y uso por parte de las abejas.

Abstract Introduction: Knowledge of vegetation richness and seasonality around Apis mellifera apiaries is an indispensable planning tool for beekeepers. It must include resource availability, food preferences and foraging behaviour. Such information is unavailable for the Argentinian Yungas, one of the most seasonal forest ecosystems in South America. Objective: To assess the availability of trophic resources through a flowering calendar and its relationship with A. mellifera pollen loads in the Yungas. Methods: In El Fuerte, Jujuy, we collected monthly samples from September to March (2014-2015 and 2015-2016) using pollen traps. We used standardized techniques for palynological analyses, and association indices for resource use. The phenophases were Beginning of flowering, Full flowering, and End of flowering. Results: We identified 47 botanical specimens to species level and 9 only to genus. In both periods there was a moderate supply of flowers at the beginning of spring, represented equally by shrub and herbaceous plants, with peak flowering in November. Subsequently, there was a drop in availability, with peak flowering again at the end of the season. In five plant species, there was a medium to high association between the plant species available and their presence in the pollen spectrum of the corbicular pollen samples collected (Vachellia aroma, Blepharocalyx salicifolius, Cantinoa sp., Vernonanthura sp. And Zanthoxylum coco). Conclusion: In this region, there is a moderate supply of shrub and herbaceous plant flowers at the beginning of spring, with peak flowering in November and at the end of the season. Only five, out of nearly 50 plant species, show an association of availability and use by bees.

Distribution, diversity, endemism, and ecology of Nymphalid butterflies (Lepidoptera: Nymphalidae) in the Loxicha Region, Oaxaca, Mexico / Distribución, diversidad, endemismo y ecología de las mariposas Nymphalidae (Lepidoptera: Nymphalidae) en la Región Loxicha, Oaxaca, México

Abstract Introduction: The Loxicha Region of Oaxaca, Mexico, has been historically important for the study of Nymphalidae, second in the Papilionoidea for species richness. Describing the diversity patterns of this butterfly clade in Loxicha can improve our understanding of the evolutionary history of the Sierra Madre del Sur, the Mexican Pacific slope, and Mexico in general. Objective: To describe the temporal and spatial patterns of Nymphalidae diversity along an elevational gradient (80-2 600 m), and to compare Loxicha's fauna with other regions in Mexico. Methods: We obtained 28 756 records from 21 sites in the Loxicha Region, representing seven years of sampling. We estimate and analyze the diversity, endemism, and distributional patterns for three elevational levels and five vegetation types. We estimated species composition and similarity with other regions of the Pacific and Atlantic slopes. Results: We identified 189 taxa, including species and subspecies, from 85 genera and ten subfamilies of Nymphalidae. Loxicha contains 46 % of the species in the family recognized for Mexico, including ten endemic species and 56 endemic subspecies. Cloud forest and low elevations were the most diverse habitats for this family. There is a clear divergence between the Atlantic and Pacific faunas, and the Sierra Madre del Sur has two faunal components. High-elevation sites in Oaxaca, and in the neighboring state of Guerrero, have a distinctive fauna, apparently isolated from low-elevation sites, revealing an archipelagic distribution for cloud forest Nymphalidae. Conclusions: The Loxicha Region is one of the richest areas for Nymphalidae in Mexico. Distribution on the Pacific slope is determined by geographical history and ecological conditions, including elevation. Nymphalidae can be used to test hypotheses of biogeographic regionalization in Mexico.

Resumen Introducción: La Región Loxicha de Oaxaca, México, ha sido históricamente importante para el estudio de Nymphalidae, segunda en riqueza de especies en Papilionoidea. Describir los patrones de diversidad de este taxón de mariposas en Loxicha puede mejorar nuestra comprensión de la historia evolutiva de la Sierra Madre del Sur, la vertiente del Pacífico mexicano y México en general. Objetivo: Describir los patrones temporales y espaciales de la diversidad de Nymphalidae a lo largo de un gradiente altitudinal (80-2 600 m), y comparar la fauna de Loxicha con otras regiones de México. Métodos: Obtuvimos 28 756 registros de 21 sitios de la Región Loxicha, que representan siete años de muestreo. Estimamos y analizamos la diversidad, el endemismo y los patrones de distribución para tres niveles altitudinales y cinco tipos de vegetación. Estimamos la composición de especies y la similitud con otras regiones de las vertientes del Pacífico y Atlántico de México. Resultados: Identificamos 189 taxones, incluyendo especies y subespecies, de 85 géneros y diez subfamilias de Nymphalidae. Loxicha contiene 46 % de las especies de la familia reconocidas para México, incluidas diez especies endémicas y 56 subespecies endémicas. El bosque mesófilo y las elevaciones bajas fueron los hábitats más diversos para esta familia. Existe una clara divergencia entre las faunas del Atlántico y del Pacífico, y la Sierra Madre del Sur tiene dos componentes faunísticos. Los sitios de elevaciones altas en Oaxaca, y en el estado vecino de Guerrero, tienen una fauna distintiva, aparentemente aislada de los sitios de elevaciones bajas, lo que revela una distribución archipelágica para los Nymphalidae del bosque mesófilo. Conclusiones: La Región Loxicha es una de las zonas más diversas para Nymphalidae en México. La distribución en la vertiente del Pacífico está determinada por la historia geográfica y las condiciones ecológicas, incluida la elevación. Nymphalidae puede usarse para probar hipótesis de regionalización biogeográfica en México.

Feno-morfología de dos especies de páramo de Miconia (Melastomataceae): optimización de la floración y fructificación para mejorar la dispersión / Pheno-morphology of two paramo species of Miconia (Melastomataceae): optimization of flowering and fructification to enhance dispersal

Introducción: El Páramo es uno de los ecosistemas más afectados por actividades humanas, lo que aumenta la necesidad de estudios fenológicos como base para el manejo y la conservación. Objetivo: Describir la feno-morfología de Miconia ligustrina y Miconia elaeoides. Métodos: De septiembre 2019 a febrero 2020, y de abril 2021 hasta agosto del mismo año, se monitoreó la fenología de 12 individuos de cada especie. Adicionalmente, se vincularon datos fenológicos de ejemplares de herbario, para corroborar los resultados obtenidos. En los dos conjuntos de datos, se describió su distribución usando estadística circular, además, se obtuvieron registros de precipitación y temperatura del área. Adicionalmente, se describió la morfometría de cada especie. Resultados: El pico de floración de M. lingustrina es en abril, y la fructificación se concentra entre junio y julio; M. elaoides, florece de forma masiva en enero, con la máxima producción de frutos en mayo. Hubo correlación entre fenología y lluvias: la floración se da en época seca y la fructificación en la época lluviosa. Las especies difieren en morfología, pero, como en otras especies de Miconia, las flores y frutos son pequeños (aprox. 5 mm). Conclusión: La floración se presenta en la época seca y la fructificación en la época lluviosa, cuando las tasas de fotosíntesis son bajas y la frugivoría incrementa. Este patrón fenológico favorece la dispersión óptima de semillas.

Introduction: Paramos are among the ecosystems that are most affected by anthropic activities, increasing the need for phenological studies as a basis for management and conservation. Objective: To describe the pheno-morphology of Miconia ligustrina and Miconia elaeoides. Methods: From September 2019 to February 2020, and from April 2021 to August of the same year, the phenology of 12 individuals of each species was monitored. Additionally, phenological data of herbariums were linked to corroborate obtained results. In the two data sets, distribution of data was described using circular statistics; in addition, we recorded the precipitation and temperature in the area. Additionally, we described the morphometry of each species. Results: The flowering peak of M. ligustrina is in April, and fruiting concentrates between June and July; in M. elaeoides, flowering is massive in January, and the highest fruit production in May. The phenology was correlated with rainfall: flowering takes place in the dry season and fruiting in the rainy season. The species differ morphologically, but, as in other Miconia species, the flowers and fruits are small (approx. 5 mm). Conclusions: Flowering occurs in the dry season and fructification in the rainy season, when the photosynthetic rate is low and frugivory increases. This phenological pattern favors optimal seed dispersal.

Comportamiento fenológico ex situ de Lippia integrifolia "incayuyo" (Verbenaceae) de una población de base genética amplia / Ex situ phenological behavior of Lippia integrifolia "incayuyo" (Verbenaceae) from a population with a broad genetic base

Abstract: Lippia integrifolia "incayuyo" is an aromatic, sub - woody shrub used in popular medicine, aperitit drinks and compound herb s. Its choleretic, antispasmodic, biocidal, antibacterial and larvicidal activity has been proven. The objective of the work was to register the phenology of a sample of 70 genotypes from a population with a broad genetic base. The phenophases studied were : vegetative growth, flower bud, flowering and fruiting fortnightly for two years. The initiation, intensity and prolongation of the phenophases were evaluated. The moment of full bloom occurs during the second half of December. Taking this date as a refer ence, a differentiated beginning of flowering was evidenced. The results of two campaigns were compared, observing that 70% of the specimens had a similar behavor, standing out some genotypes for presenting an early flowering and longer duration. The recor ded variability suggests that much of it could be due to intrinsic factors of the plant, and therefore, feasible to be selected .

Resumen: Lippia integrifolia "incayuyo" es un arbusto aromático, subleñoso empleado en la medicina popular, bebidas aperitivas y yerbas compuestas. Se ha comprobado su actividad colerética, antiespasmódica, biocida, antibacteriana y larvicida. El objetivo del trabajo fue registrar la fenología d e una muestra de 70 genotipos de una población de base genética amplia. Las fenofases estudiadas fueron: crecimiento vegetativo, botón floral, floración y fructificación quincenalmente durante dos años. Se evaluó inicio, intensidad y prolongación de las fe nofases. El momento de plena floración ocurre durante la segunda quincena de diciembre. Tomando esta fecha como referencia, se evidenció un inicio de floración diferenciada. Se compararon los resultados de dos campañas, observando que el 70% de los ejempla res tuvo un compartimiento semejante, destacándose algunos genotipos por presentar una floración temprana y de prolongación superior. La variabilidad registrada sugiere que gran parte de ésta podría deberse a factores intrínsecos de la planta, y por ello, factible de ser seleccionados.

Plant age, crop stage and surrounding habitats: their impact on sucking pests and predators complex in cotton (Gossypium hirsutum L) field plots in arid climate at district Layyah, Punjab, Pakistan / Idade da planta, estágio da cultura e habitats circundantes: seu impacto no complexo de pragas sugadoras e predadores em parcelas de algodão (Gossypium hirsutum L) em clima árido no distrito de Layyah, Punjab, Paquistão

Sucking pests are major threat to cotton field crop which cause unbearable losses to the crop yield. Aim of the current study was to record seasonal dynamics of major sucking insect pests including whitefly, jassid, thrips and their natural arthropod predators i.e. green lacewings and spiders in cotton field plots. The effects of surrounding field crops on pests' density and predatory efficiency of predators were also recorded. For sampling and survey of insects, the visual counting was found to be the most efficient method for recording the abundance of insects, trailed by net sweeping and tapping. Whitefly was the most dominant sucking pest found on the vegetative stage of cotton, followed by jassid and thrips. Fluctuated populations of predatory arthropods, spiders and green lacewings were also recorded during whole cropping season however, the densities of pests and predators varied with crop phenology. Spiders' population was encouraging at both vegetative and flowering stage and also the same trend of jassid and whitefly were observed at both stages of the crop. Surrounding habitats showed non-significant effect on population densities of insect pests and predators. For abiotic factors, the spiders showed strong positive correlation with humidity and temperature. However, green lacewing was only positively correlated with humidity. On the other hand, the populations of whitefly, jassid and thrips showed non-significant correlation with both temperature and humidity. Overall densities of sucking insect pests were found above economic threshold level. The plant age, crop stage and surrounding habitats effect on the population fluctuation of pests as well as the predators' abundance. The future studies are also warranted to investigate the altered habitats and multiple trap cropping to find out their impact on unattended insect predators and parasitoids in cotton crop.

As pragas sugadoras são uma grande ameaça para a cultura do algodão, causando perdas insuportáveis no rendimento da cultura. O objetivo do estudo atual foi registrar a dinâmica sazonal das principais pragas de insetos sugadores, incluindo mosca-branca, jassid, tripes e seus artrópodes predadores naturais, ou seja, crisopídeos e aranhas verdes em parcelas de algodão. Os efeitos das plantações circundantes na densidade de pragas e na eficiência predatória de predadores também foram registrados. Para amostragem e pesquisa de insetos, a contagem visual foi considerada o método mais eficiente para registrar a abundância de insetos, seguido por varredura e batida de rede. A mosca-branca foi a praga sugadora mais dominante encontrada na fase vegetativa do algodoeiro, seguida pelo jassid e tripes. Populações flutuantes de artrópodes predadores, aranhas e crisálidas também foram registradas durante toda a safra, no entanto as densidades de pragas e predadores variaram com a fenologia da cultura. A população de aranhas foi encorajadora tanto na fase vegetativa como na floração e também a mesma tendência de jassid e mosca-branca foi observada em ambas as fases da cultura. Os habitats circundantes mostraram efeito não significativo nas densidades populacionais de insetos-praga e predadores. Para os fatores abióticos, as aranhas apresentaram forte correlação positiva com umidade e temperatura. No entanto, lacewing verde foi apenas positivamente correlacionado com a umidade. Por outro lado, as populações de mosca-branca, jassid e tripes apresentaram correlação não significativa com temperatura e umidade. As densidades gerais de pragas sugadoras de insetos foram encontradas acima do nível do limiar econômico. A idade da planta, o estágio da cultura e os habitats circundantes afetam a flutuação populacional de pragas, bem como a abundância de predadores. Os estudos futuros também são necessários para investigar os habitats alterados e cultivo com armadilhas múltiplas [...].

Plant age, crop stage and surrounding habitats: their impact on sucking pests and predators complex in cotton (Gossypium hirsutum L.) field plots in arid climate at district Layyah, Punjab, Pakistan

Abstract Sucking pests are major threat to cotton field crop which cause unbearable losses to the crop yield. Aim of the current study was to record seasonal dynamics of major sucking insect pests including whitefly, jassid, thrips and their natural arthropod predators i.e. green lacewings and spiders in cotton field plots. The effects of surrounding field crops on pests density and predatory efficiency of predators were also recorded. For sampling and survey of insects, the visual counting was found to be the most efficient method for recording the abundance of insects, trailed by net sweeping and tapping. Whitefly was the most dominant sucking pest found on the vegetative stage of cotton, followed by jassid and thrips. Fluctuated populations of predatory arthropods, spiders and green lacewings were also recorded during whole cropping season however, the densities of pests and predators varied with crop phenology. Spiders population was encouraging at both vegetative and flowering stage and also the same trend of jassid and whitefly were observed at both stages of the crop. Surrounding habitats showed non-significant effect on population densities of insect pests and predators. For abiotic factors, the spiders showed strong positive correlation with humidity and temperature. However, green lacewing was only positively correlated with humidity. On the other hand, the populations of whitefly, jassid and thrips showed non-significant correlation with both temperature and humidity. Overall densities of sucking insect pests were found above economic threshold level. The plant age, crop stage and surrounding habitats effect on the population fluctuation of pests as well as the predators abundance. The future studies are also warranted to investigate the altered habitats and multiple trap cropping to find out their impact on unattended insect predators and parasitoids in cotton crop.

Resumo As pragas sugadoras são uma grande ameaça para a cultura do algodão, causando perdas insuportáveis no rendimento da cultura. O objetivo do estudo atual foi registrar a dinâmica sazonal das principais pragas de insetos sugadores, incluindo mosca-branca, jassid, tripes e seus artrópodes predadores naturais, ou seja, crisopídeos e aranhas verdes em parcelas de algodão. Os efeitos das plantações circundantes na densidade de pragas e na eficiência predatória de predadores também foram registrados. Para amostragem e pesquisa de insetos, a contagem visual foi considerada o método mais eficiente para registrar a abundância de insetos, seguido por varredura e batida de rede. A mosca-branca foi a praga sugadora mais dominante encontrada na fase vegetativa do algodoeiro, seguida pelo jassid e tripes. Populações flutuantes de artrópodes predadores, aranhas e crisálidas também foram registradas durante toda a safra, no entanto as densidades de pragas e predadores variaram com a fenologia da cultura. A população de aranhas foi encorajadora tanto na fase vegetativa como na floração e também a mesma tendência de jassid e mosca-branca foi observada em ambas as fases da cultura. Os habitats circundantes mostraram efeito não significativo nas densidades populacionais de insetos-praga e predadores. Para os fatores abióticos, as aranhas apresentaram forte correlação positiva com umidade e temperatura. No entanto, lacewing verde foi apenas positivamente correlacionado com a umidade. Por outro lado, as populações de mosca-branca, jassid e tripes apresentaram correlação não significativa com temperatura e umidade. As densidades gerais de pragas sugadoras de insetos foram encontradas acima do nível do limiar econômico. A idade da planta, o estágio da cultura e os habitats circundantes afetam a flutuação populacional de pragas, bem como a abundância de predadores. Os estudos futuros também são necessários para investigar os habitats alterados e cultivo com armadilhas múltiplas para descobrir seu impacto sobre predadores de insetos e parasitoides desacompanhados na cultura do algodão.

Plant age, crop stage and surrounding habitats: their impact on sucking pests and predators complex in cotton (Gossypium hirsutum L. ) field plots in arid climate at district Layyah, Punjab, Pakistan / Idade da planta, estágio da cultura e habitats circundantes: seu impacto no complexo de pragas sugadoras e predadores em parcelas de algodão (Gossypium hirsutum L. ) em clima árido no distrito de Layyah, Punjab, Paquistão

Sucking pests are major threat to cotton field crop which cause unbearable losses to the crop yield. Aim of the current study was to record seasonal dynamics of major sucking insect pests including whitefly, jassid, thrips and their natural arthropod predators i.e. green lacewings and spiders in cotton field plots. The effects of surrounding field crops on pests' density and predatory efficiency of predators were also recorded. For sampling and survey of insects, the visual counting was found to be the most efficient method for recording the abundance of insects, trailed by net sweeping and tapping. Whitefly was the most dominant sucking pest found on the vegetative stage of cotton, followed by jassid and thrips. Fluctuated populations of predatory arthropods, spiders and green lacewings were also recorded during whole cropping season however, the densities of pests and predators varied with crop phenology. Spiders' population was encouraging at both vegetative and flowering stage and also the same trend of jassid and whitefly were observed at both stages of the crop. Surrounding habitats showed non-significant effect on population densities of insect pests and predators. For abiotic factors, the spiders showed strong positive correlation with humidity and temperature. However, green lacewing was only positively correlated with humidity. On the other hand, the populations of whitefly, jassid and thrips showed non-significant correlation with both temperature and humidity. Overall densities of sucking insect pests were found above economic threshold level. The plant age, crop stage and surrounding habitats effect on the population fluctuation of pests as well as the predators' abundance. The future studies are also warranted to investigate the altered habitats and multiple trap cropping to find out their impact on unattended insect predators and parasitoids in cotton crop.

As pragas sugadoras são uma grande ameaça para a cultura do algodão, causando perdas insuportáveis no rendimento da cultura. O objetivo do estudo atual foi registrar a dinâmica sazonal das principais pragas de insetos sugadores, incluindo mosca-branca, jassid, tripes e seus artrópodes predadores naturais, ou seja, crisopídeos e aranhas verdes em parcelas de algodão. Os efeitos das plantações circundantes na densidade de pragas e na eficiência predatória de predadores também foram registrados. Para amostragem e pesquisa de insetos, a contagem visual foi considerada o método mais eficiente para registrar a abundância de insetos, seguido por varredura e batida de rede. A mosca-branca foi a praga sugadora mais dominante encontrada na fase vegetativa do algodoeiro, seguida pelo jassid e tripes. Populações flutuantes de artrópodes predadores, aranhas e crisálidas também foram registradas durante toda a safra, no entanto as densidades de pragas e predadores variaram com a fenologia da cultura. A população de aranhas foi encorajadora tanto na fase vegetativa como na floração e também a mesma tendência de jassid e mosca-branca foi observada em ambas as fases da cultura. Os habitats circundantes mostraram efeito não significativo nas densidades populacionais de insetos-praga e predadores. Para os fatores abióticos, as aranhas apresentaram forte correlação positiva com umidade e temperatura. No entanto, lacewing verde foi apenas positivamente correlacionado com a umidade. Por outro lado, as populações de mosca-branca, jassid e tripes apresentaram correlação não significativa com temperatura e umidade. As densidades gerais de pragas sugadoras de insetos foram encontradas acima do nível do limiar econômico. A idade da planta, o estágio da cultura e os habitats circundantes afetam a flutuação populacional de pragas, bem como a abundância de predadores. Os estudos futuros também são necessários para investigar os habitats alterados e cultivo com armadilhas múltiplas para descobrir seu impacto sobre predadores de insetos e parasitoides desacompanhados na cultura do algodão.

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.

Breeding biology in a population of Rufous-collared Sparrow (Zonotrichia capensis, Statius Müller, 1776) at different elevations in the Tropical Andes / Biología reproductiva en una población de Gorrión Copetón (Zonotrichia capensis, Statius Müller, 1776) a diferente elevación en los Andes tropicales

Abstract: Breeding biology in a population of Rufous-collared Sparrow (Zonotrichia capensis, Statius Müller, 1776) at different elevations in the Tropical Andes. Elevation strongly influences the evolution of life history traits associated with bird physiology and reproduction. Since life history traits in birds are modulated by environmental factors that vary with elevation, we expected to find changes in breeding biology traits of Z. capensis at different elevations. In this study, we compared the breeding biology (clutch size, egg volume, and reproductive activity) of a non-migratory population of Z. capensis in two localities at different elevations (low and high elevations, 1 800 m a.s.l., and 3 800 m a.s.l., respectively) and same latitude in the Colombian Tropical Andes for a one-year period. We found no differences in clutch size between the localities; however, egg volume was higher at high elevation. Furthermore, the reproductive activity differed significantly between localities. We propose that clutch size is likely conserved throughout the altitudinal distribution of Z. capensis since the species evolved in Tropical lowlands. On the other hand, the larger egg volume at high elevation could obey local environmental factors that may favor the reproductive success of the population.

Resumen: Biología reproductiva en una población de Gorrión Copetón (Zonotrichia capensis, Statius Müller, 1776) a diferente elevación en los Andes tropicales. La elevación influye fuertemente en la evolución de los rasgos de historia de vida asociados con la fisiología y reproducción de las aves. Dado que los rasgos de historia de vida en las aves están modulados por factores ambientales que varían con la elevación, esperábamos encontrar cambios en los rasgos de la biología reproductiva de Z. capensis a diferentes elevaciones. En este estudio, comparamos la biología reproductiva (tamaño de puesta, volumen del huevo y actividad reproductiva) de una población no migratoria de Z. capensis en dos localidades a diferente elevación (elevación baja 1 800 m snm, y elevación alta 3 800 m snm,) y a la misma latitud en los Andes tropicales colombianos por un período de un año. No encontramos diferencias en el tamaño de puesta entre las localidades; sin embargo, el volumen del huevo fue mayor en elevación alta. Además, la actividad reproductiva difirió entre las localidades. Proponemos que el tamaño de puesta probablemente se conserve en toda la distribución altitudinal de Z. capensis, ya que la especie evolucionó en las tierras bajas tropicales. Por otro lado, el mayor volumen del huevo a elevación alta podría obedecer a factores ambientales locales que pueden favorecer el éxito reproductivo de la población.

Searching for synthetic mechanisms on how biological traits mediate species responses to climate change / Em busca de mecanismos sintéticos sobre como os atributos biológicos mediam as respostas das espécies às mudanças climáticas

Abstract: Climate change will likely be the most significant challenge faced by species in this century, and species' ability to cope with climate change depends on their life history and ecological and evolutionary traits. Understanding how these traits mediate species' responses is beneficial for identifying more vulnerable species or prone to extinction risk. Here, we carried out a literature review describing how four traits commonly used in vulnerability assessments (i.e. clutch size, diet breadth, dispersal ability, and climatic tolerance) may determine species vulnerability. We also portray the possible mechanisms that explain how these traits govern species responses to climate change. The literature suggests different mechanisms operating for the evaluated traits. The mechanism of response to climate change differs between species inhabiting tropical and temperate regions: while species from the temperate areas may respond positively to temperature rise, tropical species may be severely affected. Since ectotherms depend on environment temperature, they are more sensitive and present different response mechanisms from endotherms.

Resumo: A mudança climática provavelmente será o maior desafio enfrentado pelas espécies neste século e a capacidade das espécies em lidar com a mudança climática depende de seus próprios atributos de história de vida, ecológicos e evolutivos. Entender como esses atributos mediam as respostas das espécies é extremamente útil para identificar espécies que são mais vulneráveis ou sujeitas ao risco de extinção. Aqui, realizamos uma revisão da literatura com foco na descrição de como quatro atributos comumente usados em avaliações de vulnerabilidade (tamanho da ninhada, amplitude da dieta, capacidade de dispersão e tolerância climática) podem realmente determinar a vulnerabilidade das espécies. Também retratamos os possíveis mecanismos que explicam como esses atributos governam as respostas das espécies à mudança climática. A literatura sugere diferentes mecanismos operando para os atributos avaliados. O mecanismo de resposta à mudança climática difere entre as espécies que habitam as regiões tropicais e temperadas: enquanto as espécies das regiões temperadas podem responder positivamente ao aumento da temperatura, as espécies tropicais podem ser severamente afetadas. Como os ectotérmicos dependem da temperatura ambiente, eles são mais sensíveis e apresentam mecanismos de resposta diferentes dos endotérmicos.

Correlations between morphological fruit types, fruit and seed colors, and functional groups / Correlaciones entre tipos morfológicos de frutos, colores de frutos y semillas y grupos funcionales

Abstract: The associations between morphological fruit types, fruit and seed colors, and functional plant traits: life forms, epiphytism, physiology, nutritional relationships, fruit phenology, and successional stage, were determined for 1,139 plant species from contrasting plant communities. Texture and dehiscence were closely related. Dehiscence is largely associated with dry tissues; indehiscence, however, is an attribute of both dry and fleshy fruits. The number of morphological fruit types was 28 or 55 for Gray's and Spjut's classifications, respectively. Fruits were predominantly dark in color (brown, purple-black, black or green), whilst seeds had both dark and light colors (brown, beige, or black). The most representative associations were mainly found between the more abundant fruit types and the colors most common. Asymmetries in the level of specialization, whereby less common fruit and seed colors tended to be associated with the most common fruit types, were also found. Fleshy fruits showed more variation as regards their coloration, and only drupes and berries showed a tendency towards a specific color: purple-black. The relationships among fruit type and color, seed color, and functional plant traits revealed the following trends: trees produced both fleshy and dry fruits; shrubs produced fleshy fruits; and herbaceous species, dry fruits. Woody species tended to have dark or bright colors, depending on their seed dispersal mechanisms and phylogenetic relations. Epiphytes were associated with dry-dehiscent fruits and brown seeds, and parasitic-hemiparasitic species had predominantly fleshy-indehiscent fruits. Pioneer species were more likely to have dry fruits, whereas fleshy fruits tended to be more frequent in late successional stage species. The C4 species, mostly herbs, had mainly one-seeded dry fruits, but multi-seeded fruits in succulent-CAM species showed morphologically diverse fruit types. Unripe and ripe fruits showed seasonal changes, especially during the rainy-dry transition period for the most abundant morphological fruit types, dry fruits during the dry period and fleshy fruited species was positively associated with the rainy season. All these trends are discussed with regard to their environmental significance and the relationships between fruit morphology, colors and functional groups. .

Resumen: Las asociaciones entre tipos morfológicos de frutos, colores de frutos y semillas y los grupos funcionales de las plantas: formas de vida, epifitismo, fisiología, relaciones nutricionales, fenología de frutos y estado sucesional fueron determinados para 1139 especies de plantas de comunidades contrastantes. La textura y dehiscencia estaban muy relacionadas. Dehiscencia está estrechamente asociada con tejidos secos, pero indehiscencia es un atributo de frutos secos y carnosos. El número de tipos morfológicos de frutos fue 28 y 55 para la clasificación de Gray y Spjut respectivamente. Los frutos fueron predominantemente de colores oscuros (marrón, negro-purpura, negro o verdes), mientras que las semillas tenían colores claros y oscuros (marrón, beige o negro). Las asociaciones más representativas fueron principalmente encontradas entre los tipos de frutos más abundantes y los colores más comunes. También se encontraron asimetrías en los niveles de especialización, donde los colores menos comunes de frutos y semillas estuvieron asociados con los tipos de frutos más comunes. Los frutos carnosos mostraron más variación en su coloración, y solo las drupas y bayas tendieron a colores específicos como negro-purpura. Las relaciones entre tipo de fruto y color, color semilla y rasgos funcionales de las plantas revelaron las siguientes tendencias: arboles producen frutos secos y carnosos; arbustos frutos carnosos y hierbas producen frutos secos. Las especies leñosas tendieron a tener colores oscuros y brillantes, relacionados con su mecanismo de dispersión y filogenia. Epifitas estuvieron asociadas con frutos secos dehiscentes y semillas marrones, y las especies parásitas-hemiparásitas tenían predominantemente frutos carnosos indehiscentes. Las especies pioneras tenían comúnmente frutos secos, mientras que los frutos carnosos tendieron a ser más frecuentes en especies de estados sucesionales tardíos. Las especies C4, principalmente hierbas, tenían frutos secos con una semilla, mientras que frutos polispermos en especies CAM mostraron diversos tipos de frutos. La fenología de frutos maduros e inmaduros mostro cambios estacionales para los más abundantes tipos morfológicos, especialmente durante el periodo de transición lluvia-sequia, frutos secos en el periodo seco y frutos carnosos durante el periodo lluvioso. Todas estas tendencias son discutidas con relación a las características ambientales y de acuerdo con las relaciones entre tipos morfológicos de frutos, colores y grupos funcionales.

Diversity of phenological patterns of Handroanthus ochraceus (Bignoniaceae) in Costa Rica

The study of phenological patterns in plant communities is of vital importance for understanding the temporal cycles of ecosystems, but there is little information on the diversity of phenological patterns that can occur at the intraspecific level, as well as the genetic or environmental factors causing this variation. In Handroanthus ochraceus, a deciduous tree species of neotropical dry forests, ecophysiological studies have proposed the release of water stress as the mechanism that triggers flowering and the sprouting of new leaves. During four years, I observed the cycles of leaf fall, flowering, fruiting, and new leaf production in seven Costa Rican sites that differed in their proximity to water courses and in soil moisture. Six were located in dry forest areas, and one in premontane forest. There were two general phenological patterns: the explosive, where trees depend on the first rains for floral anthesis and the expansion of leaf meristems, and the staggered pattern, with unsynchronized flowering at the beginning and middle of the dry season, independently of rainfall. Although this phenological variation has been previously recognized, the occurrence of these two patterns is not determined by the proximity of water courses, contradicting phenological models proposed for this tree species.

El estudio de los patrones fenológicos en comunidades de plantas es de importancia vital para entender los ciclos temporales de los ecosistemas, pero hay poca información sobre la diversidad de patrones fenológicos que pueden ocurrir a nivel intraespecífico, así como sobre los factores genéticos o ambientales que explican esta variación. En Handroanthus ochraceus, una especie arbórea de bosques secos, estudios ecofisiológicos han propuesto la reducción del estrés hídrico como el mecanismo que dispara la floración y la brotación de hojas nuevas. Durante cuatro años se observaron los ciclos de caída de hojas, floración, fructificación y producción de hojas nuevas en esta especie en siete sitios en Costa Rica, que diferían en su proximidad a fuentes de agua y por consiguiente en los niveles de humedad del suelo. Dos patrones fenológicos generales fueron observados: el patrón explosivo, en el cual los árboles dependen de las primeras lluvias para desarrollar la antesis floral y la expansión de los meristemos foliares, y el patrón escalonado, en el cual los árboles pueden florecer en una forma modular durante el comienzo y parte media de la estación seca, en forma independiente de las lluvias. Aunque esta variación en patrones fenológicos ha sido previamente reconocida, en este estudio se demuestra por vez primera que esta variación no está necesariamente asociada a diferentes niveles de humedad del suelo, contradiciendo modelos fenológicos propuestos para esta especie de árbol.

Variación mensual y anual de la riqueza y abundancia de aves en un mosaico agrícola periurbano tropical

Los mosaicos agrícolas han sido estudiados para comprender como contribuyen a mantener la riqueza de las especies según el tipo de matriz. Sin embargo, el estudio de la fenología de las especies dentro de estos ambientes ha sido dejado de lado. Por lo tanto mi objetivo con esta investigación fue analizar los cambios en la riqueza y abundancia de las especies de aves entre meses dentro de un mosaico periurbano agrícola tropical. Adicionalmente analicé si los cambios en la riqueza de las especies varían según su grado de dependencia por el bosque y el gremio alimenticio al que pertenecen. Realice muestreos mensuales del 2002 al 2010 dentro de una matriz de 30 ha dominada por potreros y cafetales, mezcladas con regeneración natural y zonas urbano-rurales. A lo largo de los muestreos anoté todos los individuos visto y escuchados de cada especie a lo largo de un trasecto de 3 km. Encontré que la riqueza de especies varió a lo largo de los meses, siendo mayor de enero a marzo (periodo seco del área de estudio) para todas las categorías de residencia (ej.: residentes, migratorias norteñas, sureñas, y altitudinales). Por tipo de gremio alimenticio la riqueza de especies aumento de enero a febrero para los frugívoros pequeños y en setiembre para los insectívoros; mientras que los otros gremios presentaron una riqueza similar de especies a lo largo del año. Por grado de dependencia de bosque solo las aves con dependencia moderada presentaron un aumento en su riqueza de enero a junio. El estudio reveló la ocurrencia de diferentes patrones de riqueza de aves a lo largo del año según el grado de dependencia del bosque, gremio alimenticio, y residencia en el país. Estas matrices poseen una riqueza alta de especies, pero en constante variación a lo largo del año, por lo que estos cambios en los patrones de riqueza deberían tomarse en cuenta para futuras políticas de conservación de dichos agroecosistemas.

Species richness inside agricultural landscape has been studied intensively and according to the type of landscape structure. However, the species phenology inside the mostly agricultural landscape is poorly understood. Therefore, my objective was to analyze species richness and individual abundance changes between months inside a semiurban agricultural landscape. Additionally, I analyzed if changes in species richness varied with the trophic guild and forest dependence of each species. From 2002 to 2010I I sampled 30 ha of coffee plantations and cattle land with natural successional regenerations and rural development. I recorded every individual listened and seen along a 3 km long transect once a month. The species richness changed between months and was higher between January and March (dry season in the study area) for all residence status (i.e., residents and north, south, and altitudinal migrants). Small frugivore species richness was higher between January and February, insectivore species richness was higher in September. The other food guilds had steady richness and abundance throughout the year. Species with a medium forest dependence showed higher species richness between January and June. Species richness phenology changed along the year depending on residence status, trophic guild, and forest dependence, a finding that should be taken into account in future conservation of this type of habitat.