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Abstract Montane forests are one of the most decimated of Neotropical biomes even though they provide a suite of valuable ecosystem services such as provision of water to lowland settlements and prevention of erosion and mudslides. In some instances, to restore these and other ecosystem services, degraded montane forests are replaced by exotic tree plantations, which cover sizeable areas in several countries. Despite their importance for assessment of ecological services and for intrinsic ecological value, comparative studies of paired native montane forest and conifer plantation are scarce along the Northern Andean Cordillera. Additionally, extrapolations are challenging because each pair of communities is highly site specific due to environmental setting, age and density of plantation, and reforestation species. Here, we assess and compare structure, biogeochemistry and ecosystem services provided by closely positioned native forest and pine plantation from a protected montane area in Venezuela. Soil nutrients and soil carbon content were 60 and 54 % respectively higher in the forest. As consequence of pine' growth form and leaf biochemistry, aboveground biomass and litter mass, as well as nutrient content and carbon stocks, were higher in the plantation. This results in the plantation storing 30 % more MgC ha-1 than the nearby forest. Canopy structure and litter properties influence the hydrology of both ecosystems through differences in rain throughfall. Most of the ecosystem services itemized are superior in the native forest, with exception that the younger plantation sequesters more carbon. An additional service provided by plantations might be that of ecological corridors that connect fragmented native forests. Our study, a specific case of nutrient and carbon cycling dynamics in paired montane forests and pine plantations, provides another set of data for the design of policy and management of considerable areas in the Neotropics with established conservation plantations.(AU)

Resumen Los bosques montanos son uno de los biomas más devastados del Neotrópico aunque proporcionan valiosos servicios ecológicos al suministrar agua a los asentamientos humanos y proteger contra la erosión y los deslaves. En algunas instancias, para mantener esos servicios ecológicos, los bosques destruidos son reemplazados por plantaciones forestales en áreas considerables de varios países. Aunque existen unos pocos estudios comparativos en a lo largo del norte de la Cordillera Andina, las extrapolaciones son difíciles debido a que estos son altamente específicos debido al entorno ambiental, edad de las plantaciones, y a la especie de conífera dominante. Nosotros analizamos y comparamos la diversidad y estructura de la vegetación, la biogeoquímica y los servicios ecológicos proporcionados por un bosque montano y una plantación de pino adyacente en un área protegida de Venezuela. Los suelos del bosque nativo contienen 60 % más de nitrógeno y 54 % más de carbono que los de la plantación. Como consecuencia de la forma de vida de los pinos y de su composición foliar, la biomasa aérea y el contenido de nutrientes en la hojarasca seca son mayores en la plantación dando como resultado que la plantación contiene 30 % MgC ha-1 más que el bosque. La estructura del dosel influencia la hidrología de ambos ecosistemas mediante diferencias en la penetración del agua de lluvia. La mayoría de los servicios son de menor valor ecológico en la plantación de pino. Sin embargo, esta ofrece servicios que no existirían en su ausencia. En regiones montañosas, las plantaciones de coníferas fueron establecidas para mejorar la provisión de agua y estabilizar los suelos. Sin embargo, nuestros resultados sugieren que las plantaciones pueden proporcionar corredores ecológicos que conecten bosques nativos fragmentados. Nuestro estudio, al proveer un ejemplo adicional de dinámica ecológica comparada, expone las diferencias en servicios ecológicos proporcionados por un bosque montano y una plantación de pino y proporciona información para establecer políticas de conservación y manejo de recursos naturales en extensas áreas neotropicales.(AU)

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AbstractPine plantations in the tropics are often employed to recondition eroded slopes from mudslides, as the Pinus caribaea plantation that shields the Universidad Simón Bolívar campus in Caracas (Venezuela). However, mismanagement of this plantation has led to its rapid degradation. The best option to maintain the protective service is to restore the plantation and direct its successional trajectory towards the neighbouring montane forest. Through experimental manipulation, we aimed to determine which factors block secondary succession and to investigate their effects. Within the experimental constraints imposed by the plantation small area, we analysed the effects of light and fertility limitation, litter accumulation and access to seed on plantation restoration. Light availability was manipulated by clearing and thinning three 800 m2 main plots. Fertilization and litter removal was applied to sub-plots within the light plots. Soils were analysed, microclimate was monitored and, for four years, stem density, species richness and basal area were tallied. Our results showed that light accessibility was the main factor deterring the successional trajectory of the plots, with varying grades of interaction with the sub-treatments. By the end of the fourth year, the cleared plot showed the largest responses in all traits (triplicating stem density and basal area and >20 times higher species richness). The main colonizers were Croton megalodendron, Ocoteafendleri, and Clusia spp. all dominant trees in the nearby native forest. We concluded that the results of this pioneer study, showed that small clearings, repeated in 3-4 year cycles are appropriate for similar restoration schemes. This procedure would create a mosaic of vegetation patches at different successional stages while protecting the slopes from erosion and increasing local biodiversity. Rev. Biol. Trop. 64 (2): 461-471. Epub 2016 June 01.

ResumenEn los trópicos, las plantaciones de pino se emplean comúnmente para proteger laderas erosionadas. Este es el caso de la plantación de Pinus caribaea que resguarda el campus de la Universidad Simón Bolívar en Caracas (Venezuela) de los deslaves de lodo. Sin embargo, el inadecuado manejo de la plantación está conduciendo a su rápido deterioro. La opción más adecuada para mantener el papel protector de la plantación es la restauración dirigida hacia una sucesión similar a la del bosque montano vecino. Mediante manipulaciones experimentales nos proponemos determinar cuáles son los factores que bloquean la sucesión secundaria e investigar sus efectos específicos. Dentro de las limitaciones impuestas por el reducido tamaño de la plantación y su rol protector, aquí analizamos los efectos de la reducida radiación solar y fertilidad del suelo, la acumulación de acículas de pino y el acceso a la fuente de semillas para la restauración. La disponibilidad de radiación solar se varió cortando y entresacando los pinos de parcelas de 800 m2. Los tratamientos de fertilidad y la remoción de acículas (por fuego y manualmente) se realizaron en subparcelas replicadas dentro de las parcelas principales. Los suelos se analizaron fisicoquímicamente, se monitoreo el microclima y, durante 4 años, se censó la densidad, el área basal y la riqueza de especies de los elementos leñosos en las parcelas. Los resultados muestran que el acceso a la radiación solar fue el factor principal que influyó sobre la colonización y crecimiento de nuevos individuos en las parcelas con diversos grados de interacción con los sub-tratamientos. Al finalizar el cuarto año, la parcela totalmente deforestada mostró las respuestas más elevadas (tres veces superior en cuanto a densidad y área basal y hasta veinte veces mayor en cuanto a riqueza de especies) comparada con las parcelas control y parcialmente deforestada. Las principales especies leñosas colonizadoras fueron: Croton megalodendron, Ocotea fendleri y Clusia spp., todas ellas dominantes en el bosque montano vecino. Concluimos que este estudio pionero muestra que el aclareo total de parcelas pequeñas, repetido en ciclos de 3-4 años es apropiado para proyectos de restauración similares. Así se obtendría un mosaico de vegetación en diferentes estadios sucesionales que mantienen el papel protector de la cubierta vegetal e incrementando la biodiversidad local.

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Pine plantations in the tropics are often employed to recondition eroded slopes from mudslides, as the Pinus caribaea plantation that shields the Universidad Simón Bolívar campus in Caracas (Venezuela). However, mismanagement of this plantation has led to its rapid degradation. The best option to maintain the protective service is to restore the plantation and direct its successional trajectory towards the neighbouring montane forest. Through experimental manipulation, we aimed to determine which factors block secondary succession and to investigate their effects. Within the experimental constraints imposed by the plantation small area, we analysed the effects of light and fertility limitation, litter accumulation and access to seed on plantation restoration. Light availability was manipulated by clearing and thinning three 800 m2 main plots. Fertilization and litter removal was applied to sub-plots within the light plots. Soils were analysed, microclimate was monitored and, for four years, stem density, species richness and basal area were tallied. Our results showed that light accessibility was the main factor deterring the successional trajectory of the plots, with varying grades of interaction with the sub-treatments. By the end of the fourth year, the cleared plot showed the largest responses in all traits (triplicating stem density and basal area and >20 times higher species richness). The main colonizers were Croton megalodendron, Ocotea fendleri, and Clusia spp. all dominant trees in the nearby native forest. We concluded that the results of this pioneer study, showed that small clearings, repeated in 3-4 year cycles are appropriate for similar restoration schemes. This procedure would create a mosaic of vegetation patches at different successional stages while protecting the slopes from erosion and increasing local biodiversity.

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Trachypogon spicatus, formerly known as Trachypogon plumosus, is a dominant grass in some savannas of Northern South America. Its dispersal unit, like many other species of the Andropogoneae tribe, bears a hygroscopic awn which facilitates its establishment in favorable microsites. Some authors have previously proposed that there is a positive correlation between awn length and dispersal unit burial, and that this relationship increases the probability of seed survival in the event of a fire, since soil acts as insulator. In this study we experimentally tested this relationship for T. spicatus. A total of 192 diaspores were placed in randomized blocks, in aluminum trays filled with soil under greenhouse conditions. Diaspores were sprayed with water daily for a month to guarantee awn movement; on the last day of the experiment, they were sprayed with red aerosol paint to determine burial depth. The effects of awn length, presence of caryopses, and presence of a pivot for the passive segment of the awn on diaspore burial were evaluated. Germination viability was tested using a tetrazolium salt test for 35 caryopses. No significant differences in diaspore burial were observed between diaspores with and without caryopses (F(2,126) = 0.034, p=0.853). A positive correlation between awn length and diaspore burial was observed only if the passive awn lacked a pivot (r(66)=0.394, p<0.05). Diaspores whose awns had a pivot point achieved significantly deeper burial distances than their counterparts (F(2,126)=7.063, p=0.005). Viability test found that 0% of caryopses tested were able to germinate; this is possibly due to the time difference between sampling and testing. We considered the presence or absence of caryopsis as an important factor, since previous studies have not yet considered it and the high production of sterile diaspores in grasses. These results suggest that the physical mechanism behind T. spicatus diaspore burial is awn torque. This would explain why our results partially support the hypothesis previously proposed by other authors regarding the effect of awn length upon dispersal unit burial, but would also account for the differences observed in burial depth between diaspores with an available pivot point for the passive awn and diaspores lacking thereof.

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Trachypogon spicatus, formerly known as Trachypogon plumosus, is a dominant grass in some savannas of Northern South America. Its dispersal unit, like many other species of the Andropogoneae tribe, bears a hygroscopic awn which facilitates its establishment in favorable microsites. Some authors have previously proposed that there is a positive correlation between awn length and dispersal unit burial, and that this relationship increases the probability of seed survival in the event of a fire, since soil acts as insulator. In this study we experimentally tested this relationship for T. spicatus. A total of 192 diaspores were placed in randomized blocks, in aluminum trays filled with soil under greenhouse conditions. Diaspores were sprayed with water daily for a month to guarantee awn movement; on the last day of the experiment, they were sprayed with red aerosol paint to determine burial depth. The effects of awn length, presence of caryopses, and presence of a pivot for the passive segment of the awn on diaspore burial were evaluated. Germination viability was tested using a tetrazolium salt test for 35 caryopses. No significant differences in diaspore burial were observed between diaspores with and without caryopses (F(2,126)= 0.034, p=0.853). A positive correlation between awn length and diaspore burial was observed only if the passive awn lacked a pivot (r(66)=0.394, p<0.05). Diaspores whose awns had a pivot point achieved significantly deeper burial distances than their counterparts (F(2,126)=7.063, p=0.005). Viability test found that 0% of caryopses tested were able to germinate; this is possibly due to the time difference between sampling and testing. We considered the presence or absence of caryopsis as an important factor, since previous studies have not yet considered it and the high production of sterile diaspores in grasses. These results suggest that the physical mechanism behind T. spicatus diaspore burial is awn torque. This would explain why our results partially support the hypothesis previously proposed by other authors regarding the effect of awn length upon dispersal unit burial, but would also account for the differences observed in burial depth between diaspores with an available pivot point for the passive awn and diaspores lacking thereof. Rev. Biol. Trop. 62 (1): 321-326. Epub 2014 March 01.

Trachypogon spicatus es una gramínea dominante de algunas sabanas del norte de Suramérica. Su unidad dispersante presenta una arista higroscópica la cual facilita su inserción en irregularidades del suelo, facilitando su ubicación en microhábitats favorables para su establecimiento. Igualmente, se ha sugerido que existe una relación positiva entre la longitud de la arista y la capacidad de entierro de la unidad dispersante. Esta relación incrementa la probabilidad de supervivencia de la semilla ante el fuego al ser el suelo aislante de las altas temperaturas. Se colocaron 192 diásporas de T. spicatus, en bloques aleatorios, en bandejas de sustrato bajo condiciones de invernadero. Luego, se sometieron a cambios de humedad para garantizar el movimiento de la arista y pintaron con aerosol rojo el último día del experimento para determinar la profundidad de entierro. Se evaluó el efecto de la presencia de cariópsis, la longitud de la arista y la presencia de un punto de apoyo para la arista sobre la profundidad de entierro de la diáspora. Se realizó una prueba de sal de Tetrazolio para probar la viabilidad de germinación de 35 cariópsis. No se observaron diferencias significativas en la profundidad de entierro entre diásporas provistas y desprovistas de cariópsis (F(2,126)=0.034, p=0.853). La longitud de la arista parece guardar una relación positiva con la profundidad de entierro únicamente para aquellas unidades dispersantes cuyas aristas no cuentan con un punto de apoyo (r(66)=0.394, p<0.05). Por otro lado, aquellas diásporas cuyas aristas contaban con un punto de apoyo lograron profundidades de entierro significativamente mayores (F(2,126)=7.063, p=0.005). La prueba de viabilidad determinó que 0% de las cariópsis analizadas eran capaces de germinar, esto posiblemente sea debido a la diferencia entre el tiempo de recolección y de prueba. Consideramos que el factor presencia/ausencia de cariópsis era importante ya que había sido ignorado en estudios previos y por la alta producción de diásporas estériles en las gramíneas. Los resultados sugieren que el mecanismo físico que opera sobre la capacidad de entierro de la unidad dispersante de T. spicatus es el torque. Esto explica por qué solo se apoyaron parcialmente las hipótesis propuestas previamente por otros autores, y justifica las diferencias observadas en cuanto a la profundidad de entierro entre unidades dispersantes cuyas aristas contaban o carecían de un punto de apoyo.

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The soil seed bank is the basis for community establishment and permanence and plays a primary role in natural restoration of degraded or altered ecosystems. As part of a restoration project, this study aimed to quantify the soil seed bank and to evaluate the effect of the needle litter layer on seedling emergence. Soil samples from a pine plantation were collected at random in the field and set to germinate in a greenhouse. Half of them were covered by a 6cm layer of dead pine needles simulating field conditions. In the field, 20 x 20cm plots were established, half were left intact and half were cleaned from the litter needles. All four treatments had 15 replicates and seedling emergence was recorded during six months. Soil seed bank density was 1 222/m2 from 17 morphotypes. In the field, the number of morphotypes and seedlings was only 9% and 6% respectively, of those emerged in the greenhouse, possibly due to watering and lack of predation in the latter. In both cases, herbs and graminoids were the dominant emerging seedlings, making up to 70-90% of the total. The needle layer didn't prevent seeds from reaching the soil but strongly reduced (> 50%) seedling emergence, although high variability within treatments resulted in no statistically significant differences. These results show that the needle layer hinders germination and/or emergence of seedlings from the seed bank. Its removal may be a recommended technique to accelerate natural restoration in pine plantations.

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The soil seed bank is the basis for community establishment and permanence and plays a primary role in natural restoration of degraded or altered ecosystems. As part of a restoration project, this study aimed to quantify the soil seed bank and to evaluate the effect of the needle litter layer on seedling emergence. Soil samples from a pine plantation were collected at random in the field and set to germinate in a greenhouse. Half of them were covered by a 6cm layer of dead pine needles simulating field conditions. in the field, 20x20cm plots were established, half were left intact and half were cleaned from the litter needles. All four treatments had 15 replicates and seedling emergence was recorded during six months. Soil seed bank density was 1 222/m² from 17 morphotypes. in the field, the number of morphotypes and seedlings was only 9% and 6% respectively, of those emerged in the greenhouse, possibly due to watering and lack of predation in the latter. in both cases, herbs and graminoids were the dominant emerging seedlings, making up to 70-90% of the total. The needle layer didn’t prevent seeds from reaching the soil but strongly reduced (>50%) seedling emergence, although high variability within treatments resulted in no statistically significant differences. These results show that the needle layer hinders germination and/or emergence of seedlings from the seed bank. its removal may be a recommended technique to accelerate natural restoration in pine plantations. Rev. Biol. Trop. 59 (3): 1071-1079. Epub 2011 September 01.

El banco de semillas del suelo es la base para el establecimiento y la permanencia de una comunidad y desempeña un papel fundamental en la restauración natural de los ecosistemas degradados o alterados. Como parte de un proyecto de restauración, este estudio tuvo como objetivo cuantificar el banco de semillas del suelo y evaluar el efecto de la capa de hojarasca de pino sobre la emergencia de las plántulas. Las muestras de suelo de la plantación de pinos fueron recolectadas al azar en el campo y se pusieron a germinar en un invernadero. La mitad de ellas estaban cubiertas por una capa de 6cm de hojarasca simulando condiciones de campo. En el campo se establecieron parcelas de 20x20cm, la mitad se dejaron intactas y la otra mitad se limpiaron de la hojarasca. Para los cuatro tratamientos se realizaron 15 repeticiones y la emergencia de plántulas se registró durante seis meses. La densidad del banco de semillas del suelo fue de 1 222/m² de 17 morfotipos. En el campo, el número de morfotipos y plántulas fue sólo de 9% y 6% respectivamente, de los que emergieron en el invernadero, posiblemente debido al riego y la falta de depredación. En ambos casos las plántulas dominantes fueron las hierbas y gramíneas, alcanzando el 70-90% del total. La capa de hojarasca no impidió que las semillas llegaran al suelo, pero se presentó una emergencia de plántulas muy reducida (>50%), debido a la alta variabilidad de los tratamientos no se produjeron diferencias estadísticamente significativas. Estos resultados muestran que la capa de hojarasca impide la germinación y/o la emergencia de las plántulas del banco de semillas del suelo. Su eliminación puede ser una técnica recomendada para acelerar la restauración natural en las plantaciones de pino.

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El género Brachiaria es muy importante económicamente debido a que algunas de sus especies han sido empleadas exitosamente como forraje en las sabanas neotropicales. En este trabajo se compara la respuesta al déficit hídrico del suelo en 5 especies del género. B. humidicola y B. dictyoneura tuvieron una variación relativamente menor en la producción de biomasa, potencial hídrico, contenido relativo de agua y fotosíntesis al ser sometidas a la sequía. B. mutica mostró una marcada respuesta evasiva, con un sistema radical que maximiza la extracción de humedad a lo largo del perfil del suelo. Esto permitiría mantener la turgencia y la actividad fotosintética hasta agotar la humedad disponible. En contraste, B. decumbens y, en menor grado, B. brizantha, mostraron un patrón de extracción de humedad que se restringió fuertemente a los primeros horizontes (0-30cm), respondiendo al déficit hídrico con un rápido cierre estomático y una menor producción de biomasa foliar. Estas especies pudieran adaptarse a períodos de sequía cortos y/o suelos con una alta humedad disponible en los horizontes superficiales. La evasión de B. mutica a lapsos cortos de sequía dependerá de la capacidad de extraer humedad de horizontes más profundos, mientras que B. dictyoneura y B. humidicola pudieran ser más exitosas en ambientes de sabana con suelos profundos y sequías más largas

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El norte de Suramérica posee una de las áreas más extensas de sabanas. En Venezuela, éstas ocupan diversos climas y suelos donde predomina la gramínea C4 Trachypogon plumosus. Esta amplia distribución sugiere una apreciable variación interpoblacional. Aquí se reportan y analizan resultados en campo e invernadero sobre la historia de vida y ecofisiología de 9 poblaciones de T. plumosus en Venezuela. Los caracteres difieren entre poblaciones y se relacionan con la geografía y ambiente de los hábitats locales. La biomasa y el tamaño individual se relacionan con la disponibilidad de agua, mientras que el contenido de nutrientes foliares esta vinculado a la fertilidad de los suelos. El área foliar específica es menor en poblaciones de ambientes más secos, lo que contribuye a minimizar los efectos de la sequía. Bajo condiciones de invernadero las diferencias interpoblacionales se mantienen sugiriendo que algunas poblaciones de T. plumosus son ecotipos establecidos genéticamente. La variación genética y fenotípica de las poblaciones son herramientas valiosas para la restauración de sabanas perturbadas, mientras que la conservación de poblaciones locales es importante para el funcionamiento y estabilidad de ecosistemas regionales. Aunque de limitada calidad nutricional, los pastos nativos dominados por Trachypogon spp. son importantes para la ganadería. La selección de ecotipos que respondan a la fertilización aumentando su potencial nutritivo merece ser considerada tanto en Venezuela como en sabanas de Colombia, Brasil y Guyana. La distribución poblacional similar a la de T. plumosus en las especies presentes en las sabanas de Venezuela sugiere que, si la diversidad genética y fenotípica de esas especies es similar a la descrita, la diversidad total asociada a las sabanas debería ser mayor a los valores relativamente bajos señalados en la literatura

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The invasion of African grasses into Neotropical savannas has altered savanna composition, structure and function. The projected increase in atmospheric CO(2) concentration has the potential to further alter the competitive relationship between native and invader grasses. The objective of this study was to quantify the responses of two populations of a widespread native C(4) grass (Trachypogon plumosus) and two African C(4) grass invaders (Hyparrhenia rufa and Melinis minutiflora) to high CO(2) concentration interacting with two primary savanna stressors: drought and herbivory. Elevated CO(2) increased the competitive potential of invader grasses in several ways. Germination and seedling size was promoted in introduced grasses. Under high CO(2), the relative growth rate of young introduced grasses was twice that of native grass (0.58 g g(-1) week(-1) vs 0.25 g g(-1) week(-1)). This initial growth advantage was maintained throughout the course of the study. Well-watered and unstressed African grasses also responded more to high CO(2) than did the native grass (biomass increases of 21-47% compared with decreases of 13-51%). Observed higher water and nitrogen use efficiency of invader grasses may aid their establishment and competitive strength in unfertile sites, specially if the climate becomes drier. In addition, high CO(2) promoted lower leaf N content more in the invader grasses. The more intensive land use, predicted to occur in this region, may interact with high CO(2) to favor the African grasses, as they generally recovered faster after simulated herbivory. The superiority of invader grasses under high CO(2) suggests further increases in their competitive strength and a potential increased rate of displacement of the native savannas in the future by grasslands dominated by introduced African species.

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African grasses, introduced into Neotropical savannas to improve forage quality, have spread successfully and displaced native plants. To understand their competitive relationships, we compared biomass production and allocation, plant architecture and phenology, net photosynthesis (Pn), water relations, and nutrient content under fire and simulated herbivory between two C4 grasses, the native Trachypogon plumosus and the introduced Hyparrhenia rufa from a seasonal savanna in Venezuela. All variables were strongly influenced by the rainfall regime. Hyparrhenia produced bigger plants (in mass and size) with a large proportion of mass (>75%) allocated to leaves and culms. Its biomass production was more affected by fire than by defoliation. In contrast, Trachypogon was more affected by defoliation than by fire which promoted a flush of leaf growth even in the dry season. Fire caused up to 85% mortality in Hyparrhenia but none in Trachypogon where it increased inflorescence production. However, fire promoted abundant seed germination and fast seedling growth in Hyparrhenia, enabling it to colonize new areas. During the growing season Trachypogon had higher Pn and lower leaf water potential (Ψ) than Hyparrhenia but differences among treatments were not significant for either grass. Pn of Trachypogon ceased at a lower Ψ (-3.0 MPa) than in Hyparrhenia (-2.0 MPa), indicating its higher tolerance to water stress. During the dry season, Trachypogon leaves remained alive and retained low Pn. Leaf nutrient content was higher during the rainy season in both species. Differences in Pn could not explain the higher seasonal biomass production of Hyparrhenia. However, its water stress evasion strategy, larger biomass allocated to leaves, abundant germination and fast seedling growth appeared to be responsible for the success of Hyparrhenia as an invader of Neotropical savannas.

RESUMO

Introduced African grasses are invading Neotropical savannas and displacing the native herbaceous community. This work, which is part of a program to understand the success of the African grasses, specifically investigates whether introduced and native grasses differ in their water relations. The water relations of the native Trachypogon plumosus and the successful invader Hyparrhenia rufa were studied in the field during two consecutive years in the seasonal savannas of Venezuela. The two C4 grasses differed clearly in their responses to water stress. H. rufa consistently had higher stomatal conductance, transpiration rate, leaf water and osmotic potential and osmotic adjustment than the native T. plumosus. Also, leaf senescence occurred much earlier during the dry season in H. rufa. Both grasses showed a combination of water stress evasion and tolerance mechanisms such as stomatal sensitivity to atmospheric or soil water stress, decreased transpiring area and osmotic adjustment. Evasion mechanisms are more conspicuous in H. rufa whereas T. plumosus is more drought tolerant and uses water more "conservatively". The evasion mechanisms and oportunistic use of water by H. rufa, characteristic of invading species, contribute to, but only partially explain, the success of this grass in the Neotropical savannas where it displaces native plants from sites with better water and nutrient status. Conversely, the higher water stress tolerance of t. plumosus is consistent with its capacity to resist invasion by alien grasses on shallow soils and sites with poorer nutrient and water status.

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The African grass Hyparrhenia rufa has established itself successfully in South American savannas (Llanos) and displaced dominant native grasses such as Trachypogon plumosus from the wetter and more fertile habitats. Several ecophysiological traits have been related to the higher competitive capacity of H. rufa. To further analyze the behavior of both species, their growth, biomass allocation, physiological and architectural responses to defoliation and water stress were compared under controlled conditions. Although total, aerial and underground biomass decreased under defoliation in both grasses, increases in clipped-leaf biomass and area compensated for defoliation in H. rufa but not in T. plumosus. This difference was due mainly to a higher proportion of assimilates being directed to leaf and tiller production and a higher leaf growth rate in the African grass as compared to T. plumosus, which showed incrased senescence under frequent defoliation. In both species, water stress ameliorated the effects of defoliation. The ability to compensate for defoliated biomass in H. rufa is possibly related to its long coevolution with large herbivores in its original African habitat and is apparently one of the causes of its success in Neotropical savannas.

RESUMO

Determinations of energy and ash content were made on the organs of some common plant species from an elevational gradient in the Venezuelan páramos. The results indicate differences in energy content among the life forms sampled. Giant rosette plants show the highest values in the roots while woody plants have the highest energy content in the leaves. The reproductive organs show a higher energy content than the vegetative organs. There is a tendency to increase energy content as elevation and environmental stress increase. The adaptive significance of these patterns is discussed. The high average energy value for páramo plants, 20.18 KJ/g ash free dry weight, is similar to values reported for alpine tundra plants and conifer forests.

RESUMO

We compared adaptive strategies in two plants of Venezuelan páramos (alpine areas): the widely distributed, caulescent, and pubescent Espeletia schultzii Wedd. with the acaulescent, nearly glabrous E. atropurpurea A.C. Smith which is restricted to mesic sites just above treeline. Both species occur together at 3,450 m, near treeline.Physiologically, E. schultzii was more drought resistant than E. atropurpurea, and was better adapted for carbon dioxide fixation under low temperatures. The densely pubescent leaves of E. schultzii are highly reflective; this increases the intensity of light needed for photosynthetic saturation and influences leaf temperature. Leaf pubescence may reduce the level of insect predation.Measurements of leaf productivity indicate higher values for E. atropurpurea during the rainy season and higher values for E. schultzii during the dry season. However, annual values of leaf productivity are similar for both species.Benefits of specialization in E. atropurpurea include reduced costs for stem and leaf hair production, higher growth rates during the rainy season and the ability to grow beneath canopies of some larger arborescent species. Costs of specialization include lower growth rates during the dry season, great susceptibility to insect predation and restriction to low elevation, mesic sites.