The legacy of biogeographic history on the composition and structure of Andean forests.

The biogeographic origin of species may help to explain differences in average tree height and aboveground biomass (AGB) of tropical mountain forests. After the Andean uplift, small-statured trees should have been among the initial colonizers of the highlands (new cold environment) from the lowland tropics, since these species are pre-adapted to cold conditions with narrow vessels that are relatively resistant to freezing. If the descendants of these small-statured clades continue to dominate tropical highland forests, there will be a high co-occurrence of close relatives at high elevations. In other words, this scenario predicts a systematic decline in tree size, AGB, and phylogenetic diversity with elevation. In contrast, the colonization of Andean forests by some large-statured clades that originated in temperate regions may modify this expectation and promote a mixing of tropical and temperate clades, thereby increasing the phylogenetic diversity in tropical highland forests. This latter scenario predicts an increase or no change of tree size, AGB, and phylogenetic diversity with elevation. We assessed how the historical immigration of large-statured temperate-affiliated tree lineages adapted to cold conditions may have influenced the composition and structure of Andean forests. Specifically, we used 92 0.25-ha forest inventory plots distributed in the tropical Andes Mountains of Colombia to assess the relationship between the phylogenetic diversity and AGB along elevational gradients. We classified tree species as being either "tropical affiliated" or "temperate affiliated" and estimated their independent contribution to forest AGB. We used structural equation modeling to separate the direct and indirect effect of elevation on AGB. We found a hump-shaped relationship of phylogenetic diversity, AGB, and tree size with elevation. The high phylogenetic diversity found between 1,800-2,200 m above sea level (asl) was due to the mixing of highland floras containing many temperate-affiliated species, and lowland floras containing mostly tropical-affiliated species. The high AGB in highland forests, which contrasted with the expected decline of AGB with elevation, was likely due to the significant contribution of temperate-affiliated species. Our findings highlight the lasting importance of biogeographic history on the composition and structure of Andean mountain forests.

Biomasa y carbono total almacenado en robledales del sur de los Andes Colombianos: aportes para el enfoque REDD+ a escala de proyectos / Biomass and total carbon in oak forests of Southern Colombian Andes: contributions to the REDD+ project-wide approach

ResumenLas estimaciones de carbono en bosques tropicales, es muy importante para entender el papel de estos ecosistemas en el ciclo de este elemento. Este conocimiento permite apoyar y soportar las decisiones y la formulación de estrategias de mitigación y adaptación para reducir las emisiones de gases de efecto invernadero (GEI). Sin embargo, cuantificaciones detalladas de las existencias totales de carbono en los bosques montanos tropicales son limitadas, a pesar de su importancia científica y para el manejo de los ecosistemas (e.g. REDD+). Cuarenta y cuatro parcelas se establecieron en un paisaje heterogéneo compuesto por bosques maduros ubicados en el sur de los Andes colombianos. El objetivo fue identificar el papel de estos ecosistemas en el almacenamiento de carbono, evaluar la contribución de los reservorios analizados (biomasa aérea, biomasa subterránea y necromasa), y realizar contribuciones al enfoque de REDD+ en la escala de proyecto. En cada parcela se midieron todos los árboles, palmas y helechos con diámetro (D) ≥ 15 cm. En el caso de las palmas, se midió la altura de 40 % de los individuos, siguiendo el Protocolo Nacional de Colombia para estimar la biomasa y el carbono en los bosques naturales. Ecuaciones alométricas nacionales fueron utilizadas para estimar la biomasa aérea, y una ecuación general propuesta por el IPCC, fue utilizada para la estimación de la biomasa subterránea. La biomasa aérea de las palmas se estimó mediante un modelo local desarrollado para bosques montanos y premontanos. La necromasa se estimó para los árboles muertos en pie y los detritos gruesos. En este último caso, se midió la longitud y los diámetros de los extremos de las piezas. Las muestras para las estimaciones de densidad de madera, se recogieron en el campo y se analizaron en el laboratorio. El carbono promedio total se estimó en 545.9 ± 84.1 Mg/ ha (± SE). La biomasa aérea contribuyó con 72.5 %, la biomasa subterránea con 13.6 %, y la necromasa con 13.9 %. La principal conclusión de este estudio, es que los bosques montanos tropicales almacenan grandes cantidades de carbono, similares a las almacenadas por los bosques tropicales de tierras bajas. Además, se observó que la inclusión de otros compartimientos podría contribuir en más del 20 % de total del almacenamiento de carbono, lo que indica que las estimaciones que sólo incluyen la biomasa aérea, subestiman en gran medida las reservas de carbono en los ecosistemas forestales. Por tal razón, se sugiere incluir otros compartimientos en las estimaciones de carbono para iniciativas o proyectos REDD+.

AbstractCarbon estimations in tropical forests are very important to understand the role of these ecosystems in the carbon cycle, and to support decisions and the formulation of mitigation and adaptive strategies to reduce the greenhouse emission gases (GHG). Nevertheless, detailed ground-based quantifications of total carbon stocks in tropical montane forests are limited, despite their high value in science and ecosystem management (e.g. REDD+). The objective was to identify the role of these ecosystems as carbon stocks, to evaluate the contribution of the pools analyzed (aboveground biomass, belowground biomass and necromass), and to make contributions to the REDD+ approach from the project scale. For this study, we established 44 plots in a heterogeneous landscape composed by old-grown forests located in the Southern Colombian Andes. In each plot, all trees, palms and ferns with diameter (D) ≥ 15 cm were measured. In the case of palms, the height was measured for 40 % of the individuals, following the Colombia National Protocol to estimate biomass and carbon in natural forests. National allometric equations were used to estimate aboveground biomass, and a global equation proposed by IPCC was used for belowground biomass estimation; besides, palms' aboveground biomass was estimated using a local model. The necromass was estimated for dead standing trees and the gross debris. In the latter case, the length and diameters of the extremes in the pieces were measured. Samples for wood density estimations were collected in the field and analyzed in the laboratory. The mean total carbon stock was estimated as 545.9 ± 84.1 Mg/ha (± S.E.). The aboveground biomass contributed with 72.5 %, the belowground biomass with 13.6 %, and the necromass with 13.9 %. The main conclusion is that montane tropical forests store a huge amount of carbon, similar to low land tropical forests. In addition, the study found that the inclusion of other pools could contribute with more than 20 % to total carbon storage, indicating that estimates that only include the aboveground biomass, largely underestimate carbon stocks in tropical forest ecosystems. These results support the importance of including other carbon pools in REDD+ initiatives' estimations. Rev. Biol. Trop. 64 (1): 399-412. Epub 2016 March 01.

Contribución de los bosques tropicales de montaña en el almacenamiento de carbono en Colombia / Contribution of tropical upland forests to carbon storage in Colombia

The tropical montane forests in the Colombian Andean region are located above 1 500m, and have been heavily deforested. Despite the general presumption that productivity and hence carbon stocks in these ecosystems are low, studies in this regard are scarce. This study aimed to i) to estimate Above Ground Biomass (AGB) in forests located in the South of the Colombian Andean region, ii) to identify the carbon storage potential of tropical montane forests dominated by the black oak Colombobalanus excelsa and to identify the relationship between AGB and altitude, and iii) to analyze the role of tropical mountain forests in conservation mechanisms such as Payment for Environmental Services (PES) and Reducing Emissions from Deforestation and Degradation (REDD+). Twenty six 0.25ha plots were randomly distributed in the forests and all trees with D≥10cm were measured. The results provided important elements for understanding the role of tropical montane forests as carbon sinks. The information produced can be used in subnational initiatives, which seek to mitigate or reduce the effects of deforestation through management or conservation of these ecosystems, like REDD+ or PES. The AGB and carbon stocks results obtained were similar to those reported for lowland tropical forests. These could be explained by the dominance and abundance of C. excelsa, which accounted for over 81% of AGB/carbon. The error associated with the estimates of AGB/carbon was 10.58%. We found a negative and significant relationship between AGB and altitude, but the higher AGB values were in middle altitudes (≈1 700-1 800m), where the environmental conditions could be favorable to their growth. The carbon storage potential of these forests was higher. However, if the historical rate of the deforestation in the study area continues, the gross emissions of CO2e to the atmosphere could turn these forests in to an important emissions source. Nowadays, it is clear that tropical montane forests are vulnerable to deforestation, especially black Oak forests due to their commercial value. Given their high carbon storage potential, the presence of endemic species and the strategic functions of these ecosystems, we recommend that they should be considered relevant during REDD+, PES or any other conservation assessment. Rev. Biol. Trop. 63 (1): 69-82. Epub 2015 March 01.

Los bosques tropicales de montaña (BTM) en la región Andina de Colombia se encuentran por encima de 1 500m, y han sido fuertemente deforestados. A pesar de la presunción general de que la productividad y las reservas de carbono en estos ecosistemas son bajas, los estudios al respecto son escasos. Este estudio tuvo por objetivo i) estimar la biomasa aérea (BA) de los bosques localizados en el sur de la región Andina colombiana, ii) identificar el potencial de almacenamiento de carbono en bosques dominados por el roble negro Colombobalanus excelsa e identificar la relación entre la BA y la altitud, y iii) analizar su papel en mecanismos de conservación, como el Pago por Servicios Ambientales (PSA) y la Reducción de Emisiones por deforestación y Degradación (REDD+). Veintiséis parcelas de 0.25ha fueron distribuidas al azar, y se midieron todos los árboles con D≥10cm. Los resultados obtenidos de BA y carbono almacenado fueron similares a los reportados para bosques tropicales de tierras bajas. Esto se puede explicar por la abundancia y dominancia de C. excelsa, que contribuye con más del 81% de la BA/carbono; el error asociado a las estimaciones fue de 10.58%. Se encontró una relación negativa y significativa entre la BA y la altitud, pero los valores más altos de BA estuvieron en las altitudes medias (≈1 700-1 800m), donde las condiciones ambientales podrían ser favorables para su crecimiento. El potencial de almacenamiento de carbono de estos bosques fue alto. Los resultados permiten tener elementos importantes para entender el papel de los BTM como sumideros de carbono. Sin embargo, si la tasa histórica de deforestación en el área de estudio continúa, las emisiones brutas de CO2e a la atmósfera podrían convertirlos en una fuente de emisiones importante. Actualmente, los BTM son vulnerables a la deforestación, especialmente los bosques de roble negro debido a su valor comercial. Teniendo en cuenta el alto potencial de almacenamiento de carbono, la presencia de especies endémicas, y las funciones estratégicas que pueden desempeñar estos ecosistemas, se recomienda que sean considerados relevantes para REDD+, PSA u otra estrategia de conservación.

The effect of highly variable topography on the spatial distribution of Aniba perutilis (Lauraceae) in the Colombian Andes.

Topography is a factor that can significantly affect the diversity and the distribution of trees species in tropical forests. Aniba perutilis, a timber species listed as vulnerable to extinction, is widely distributed in Andean forest fragments, especially in those with highly variable topography. Based on field surveys and logistic regression analyses, we studied the population structure and the effect of highly variable topography on the spatial distribution of this tree in three protected forest fragments in the central Andes of Colombia. Individuals of A. perutilis were mainly found on mountain ridges and hills with gentle slopes; no individuals were found in valleys. Using a species distribution model with presence/absence data, we showed that the available habitat for A. perutilis is significantly smaller than the extension of the fragments and much smaller than the extension of the currently protected areas. Our results have important implications for the conservation ofA. perutilis and likely for other threatened Andean tree species, which can also have locally restricted distributions due to highly variable local topography.

The effect of highly variable topography on the spatial distribution of Aniba perutilis (Lauraceae) in the Colombian Andes

Topography is a factor that can significantly affect the diversity and the distribution of trees species in tropical forests. Aniba perutilis, a timber species listed as vulnerable to extinction, is widely distributed in Andean forest fragments, especially in those with highly variable topography. Based on field surveys and logistic regression analyses, we studied the population structure and the effect of highly variable topography on the spatial distribution of this tree in three protected forest fragments in the central Andes of Colombia. Individuals of A. perutilis were mainly found on mountain ridges and hills with gentle slopes; no individuals were found in valleys. Using a species distribution model with presence/absence data, we showed that the available habitat for A. perutilis is significantly smaller than the extension of the fragments and much smaller than the extension of the currently protected areas. Our results have important implications for the conservation of A. perutilis and likely for other threatened Andean tree species, which can also have locally restricted distributions due to highly variable local topography.

La topografía es un factor que puede afectar considerablemente la diversidad y la distribución de las especies de árboles tropicales. Aniba perutilis, una especie de árbol maderable vulnerable a la extinción, está ampliamente distribuida en fragmentos de bosques andinos, especialmente en aquellos con topografía altamente variable. A partir de trabajo de campo y análisis de regresión logística, estudiamos la estructura de la población y los efectos de la topografía sobre la distribución espacial de este árbol en tres fragmentos de bosque en la cordillera central de Colombia que actualmente se encuentran protegidos. Los individuos de A. perutilis se encontraron principalmente en los filos de montaña y colinas con gradientes topográficos suaves; no se encontraron individuos en los valles. A partir de un modelo de distribución de especies usando datos de presencia/ausencia, mostramos que el hábitat disponible para A. perutilis es considerablemente más pequeño que la extensión de los fragmentos y mucho más reducido que la extensión actual de las áreas protegidas. Nuestros resultados tienen implicaciones importantes para la conservación de A. perutilis y probablemente otras especies de árboles andinos amenazados, los cuales pueden estar restringidos de forma similar debido a la variabilidad topográfica local.