Variable thermal plasticity of leaf functional traits in Andean tropical montane forests.

Tropical montane forests (TMFs) are biodiversity hotspots and provide vital ecosystem services, but they are disproportionately vulnerable to climate warming. In the Andes, cold-affiliated species from high elevations are being displaced at the hot end of their thermal distributions by warm-affiliated species migrating upwards from lower elevations, leading to compositional shifts. Leaf functional traits are strong indicators of plant performance and at the community level have been shown to vary along elevation gradients, reflecting plant adaptations to different environmental niches. However, the plastic response of such traits to relatively rapid temperature change in Andean TMF species remains unknown. We used three common garden plantations within a thermosequence in the Colombian Andes to investigate the warming and cooling responses of key leaf functional traits in eight cold- and warm-affiliated species with variable thermal niches. Cold-affiliated species shifted their foliar nutrient concentrations when exposed to warming, while all other traits did not significantly change; contrastingly, warm-affiliated species were able to adjust structural, nutrient and water-use efficiency traits from acquisitive to conservative strategies in response to cooling. Our findings suggest that cold-affiliated species will struggle to acclimate functional traits to warming, conferring warm-affiliated species a competitive advantage under climate change.

A new species of Piper (Piperaceae) with peltate leaves from Serranía de las Quinchas, Colombia.

Piperquinchasense is described and illustrated as a new species occurring in the understory of wet montane forest of the middle Magdalena Valley in Colombia, the easternmost portion of the Chocó Region. Its relationships are discussed with related taxa from the Macrostachys clade. An identification key for 35 Neotropical Piper species with peltate leaves is provided.

ResumenPiperquinchasense se describe e ilustra como una nueva especie que ocurre en el sotobosque de bosques húmedos montanos del valle medio del Magdalena en Colombia, la porción más oriental del Chocó Biogeográfico. Se discuten sus relaciones con otras especies del clado Macrostachys. Se presenta una clave de identificación para 35 especies de Piper Neotropical con hojas peltadas.

Acclimation of photosynthetic capacity and foliar respiration in Andean tree species to temperature change.

Climate warming is causing compositional changes in Andean tropical montane forests (TMFs). These shifts are hypothesised to result from differential responses to warming of cold- and warm-affiliated species, with the former experiencing mortality and the latter migrating upslope. The thermal acclimation potential of Andean TMFs remains unknown. Along a 2000 m Andean altitudinal gradient, we planted individuals of cold- and warm-affiliated species (under common soil and irrigation), exposing them to the hot and cold extremes of their thermal niches, respectively. We measured the response of net photosynthesis (Anet ), photosynthetic capacity and leaf dark respiration (Rdark ) to warming/cooling, 5 months after planting. In all species, Anet and photosynthetic capacity at 25°C were highest when growing at growth temperatures (Tg ) closest to their thermal means, declining with warming and cooling in cold-affiliated and warm-affiliated species, respectively. When expressed at Tg , photosynthetic capacity and Rdark remained unchanged in cold-affiliated species, but the latter decreased in warm-affiliated counterparts. Rdark at 25°C increased with temperature in all species, but remained unchanged when expressed at Tg . Both species groups acclimated to temperature, but only warm-affiliated species decreased Rdark to photosynthetic capacity ratio at Tg as temperature increased. This could confer them a competitive advantage under future warming.

Bracken fern does not diminish arbuscular mycorrhizal fungus inoculum potential in tropical deforested areas.

Tropical montane forests are threatened by uncontrolled fire events because of agricultural expansion. Consequently, deforested areas frequently are dominated by the bracken fern, Pteridium spp., for long periods, and forest regeneration is limited. Despite considerable research on bracken-dominated ecosystems, little is known about the relationship between bracken mycorrhizal fungi and tree seedlings. Arbuscular mycorrhizal fungi (AMF) form symbiotic relationships with terrestrial plants, providing nutrients and protection against pathogens and promoting seedling growth and establishment. Therefore, AMF inoculum have high potential for forest restoration programs. Here, we compare the species diversity of AMF spores, root colonization, and seedling growth of Clusia trochiformis 1 year after the addition of different liquefied root inocula: forest conspecific, forest heterospecific, and from Pteridium rhizomes. Thirteen morphospecies of arbuscular mycorrhizal fungi were identified on the roots of C. trochiformis, and Glomus spp. were the most abundant in all treatments. No differences were observed in spore species richness and diversity among treatments, but spore density was the highest subsequent to the Pteridium inoculum. There was no significant difference in mycorrhizal root colonization and seedling growth of C. trochiformis among inoculated treatments. We found a positive relation between root colonization and total biomass. This study shows that the AMF communities in bracken areas and forests present similar characteristics and that the bracken fern does not limit AMF inoculum potential, favouring seedling growth of Clusia.

Nutrient cycling drives plant community trait assembly and ecosystem functioning in a tropical mountain biodiversity hotspot.

Community trait assembly in highly diverse tropical rainforests is still poorly understood. Based on more than a decade of field measurements in a biodiversity hotspot of southern Ecuador, we implemented plant trait variation and improved soil organic matter dynamics in a widely used dynamic vegetation model (the Lund-Potsdam-Jena General Ecosystem Simulator, LPJ-GUESS) to explore the main drivers of community assembly along an elevational gradient. In the model used here (LPJ-GUESS-NTD, where NTD stands for nutrient-trait dynamics), each plant individual can possess different trait combinations, and the community trait composition emerges via ecological sorting. Further model developments include plant growth limitation by phosphorous (P) and mycorrhizal nutrient uptake. The new model version reproduced the main observed community trait shift and related vegetation processes along the elevational gradient, but only if nutrient limitations to plant growth were activated. In turn, when traits were fixed, low productivity communities emerged due to reduced nutrient-use efficiency. Mycorrhizal nutrient uptake, when deactivated, reduced net primary production (NPP) by 61-72% along the gradient. Our results strongly suggest that the elevational temperature gradient drives community assembly and ecosystem functioning indirectly through its effect on soil nutrient dynamics and vegetation traits. This illustrates the importance of considering these processes to yield realistic model predictions.

Up in the air: Threats to Afromontane biodiversity from climate change and habitat loss revealed by genetic monitoring of the Ethiopian Highlands bat.

While climate change is recognized as a major future threat to biodiversity, most species are currently threatened by extensive human-induced habitat loss, fragmentation and degradation. Tropical high-altitude alpine and montane forest ecosystems and their biodiversity are particularly sensitive to temperature increases under climate change, but they are also subject to accelerated pressures from land conversion and degradation due to a growing human population. We studied the combined effects of anthropogenic land-use change, past and future climate changes and mountain range isolation on the endemic Ethiopian Highlands long-eared bat, Plecotus balensis, an understudied bat that is restricted to the remnant natural high-altitude Afroalpine and Afromontane habitats. We integrated ecological niche modelling, landscape genetics and model-based inference to assess the genetic, geographic and demographic impacts of past and recent environmental changes. We show that mountain range isolation and historic climates shaped population structure and patterns of genetic variation, but recent anthropogenic land-use change and habitat degradation are associated with a severe population decline and loss of genetic diversity. Models predict that the suitable niche of this bat has been progressively shrinking since the last glaciation period. This study highlights threats to Afroalpine and Afromontane biodiversity, squeezed to higher altitudes under climate change while losing genetic diversity and suffering population declines due to anthropogenic land-use change. We conclude that the conservation of tropical montane biodiversity requires a holistic approach, using genetic, ecological and geographic information to understand the effects of environmental changes across temporal scales and simultaneously addressing the impacts of multiple threats.

Dynamics and multi-annual fate of atmospherically deposited nitrogen in montane tropical forests.

The effects of nitrogen (N) deposition on forests largely depend on its fate after entering the ecosystem. While several studies have addressed the forest fate of N deposition using 15 N tracers, the long-term fate and redistribution of deposited N in tropical forests remains unknown. Here, we applied 15 N tracers to examine the fates of deposited ammonium ( NH 4 + ) and nitrate ( NO 3 - ) separately over 3 years in a primary and a secondary tropical montane forest in southern China. Three months after 15 N tracer addition, over 60% of 15 N was retained in the forests studied. Total ecosystem retention did not change over the study period, but between 3 months and 3 years following deposition 15 N recovery in plants increased from 10% to 19% and 13% to 22% in the primary and secondary forests, respectively, while 15 N recovery in the organic soil declined from 16% to 2% and 9% to 2%. Mineral soil retained 50% and 35% of 15 N in the primary and secondary forests, with retention being stable over time. The total ecosystem retention of the two N forms did not differ significantly, but plants retained more 15 NO 3 - than 15 NH 4 + and the organic soil more 15 NH 4 + than NO 3 - . Mineral soil did not differ in 15 NH 4 + and 15 NO 3 - retention. Compared to temperate forests, proportionally more 15 N was distributed to mineral soil and plants in these tropical forests. Overall, our results suggest that atmospherically deposited NH 4 + and NO 3 - is rapidly lost in the short term (months) but thereafter securely retained within the ecosystem, with retained N becoming redistributed to plants and mineral soil from the organic soil. This long-term N retention may benefit tropical montane forest growth and enhance ecosystem carbon sequestration.

Contrasting Dependencies of Photosynthetic Capacity on Leaf Nitrogen in Early- and Late-Successional Tropical Montane Tree Species.

Differences in photosynthetic capacity among tree species and tree functional types are currently assumed to be largely driven by variation in leaf nutrient content, particularly nitrogen (N). However, recent studies indicate that leaf N content is often a poor predictor of variation in photosynthetic capacity in tropical trees. In this study, we explored the relative importance of area-based total leaf N content (Ntot) and within-leaf N allocation to photosynthetic capacity versus light-harvesting in controlling the variation in photosynthetic capacity (i.e. V cmax, J max) among mature trees of 12 species belonging to either early (ES) or late successional (LS) groups growing in a tropical montane rainforest in Rwanda, Central Africa. Photosynthetic capacity at a common leaf temperature of 25˚C (i.e. maximum rates of Rubisco carboxylation, V cmax25 and of electron transport, J max25) was higher in ES than in LS species (+ 58% and 68% for V cmax25 and J max25, respectively). While Ntot did not significantly differ between successional groups, the photosynthetic dependency on Ntot was markedly different. In ES species, V cmax25 was strongly and positively related to Ntot but this was not the case in LS species. However, there was no significant trade-off between relative leaf N investments in compounds maximizing photosynthetic capacity versus compounds maximizing light harvesting. Both leaf dark respiration at 25˚C (+ 33%) and, more surprisingly, apparent photosynthetic quantum yield (+ 35%) was higher in ES than in LS species. Moreover, Rd25 was positively related to Ntot for both ES and LS species. Our results imply that efforts to quantify carbon fluxes of tropical montane rainforests would be improved if they considered contrasting within-leaf N allocation and photosynthetic Ntot dependencies between species with different successional strategies.

Why size matters: the interactive influences of tree diameter distribution and sap flow parameters on upscaled transpiration.

In stands with a broad range of diameters, a small number of very large trees can disproportionately influence stand basal area and transpiration (Et). Sap flow-based Et estimates may be particularly sensitive to large trees due to nonlinear relationships between tree-level water use (Q) and tree diameter at breast height (DBH). Because Q is typically predicted on the basis of DBH and sap flow rates measured in a subset of trees and then summed to obtain Et, we assessed the relative importance of DBH and sap flow variables (sap velocity, Vs, and sapwood depth, Rs) in determining the magnitude of Et and its dependence on large trees in a tropical montane forest ecosystem. Specifically, we developed a data-driven simulation framework to vary the relationship between DBH and Vs and stand DBH distribution and then calculate Q, Et and the proportion of Et contributed by the largest tree in each stand. Our results demonstrate that variation in how Rs is determined in the largest trees can alter estimates up to 26% of Et while variation in how Vs is determined can vary results by up to 132%. Taken together, these results highlight a great need to expand our understanding of water transport in large trees as this hinders our ability to predict water fluxes accurately from stand to catchment scales.

Ciclos biogeoquímicos en bosques naturales y plantaciones de coníferas en ecosistemas de alta montaña de Colombia

Biogeochemical cycles in natural forest and conifer plantations in the high mountains of Colombia. Plant litter production and decomposition are two important processes in forest ecosystems, since they provide the main organic matter input to soil and regulate nutrient cycling. With the aim to study these processes, litterfall, standing litter and nutrient return were studied for three years in an oak forest (Quercus humboldtii), pine (Pinus patula) and cypress (Cupressus lusitanica) plantations, located in highlands of the Central Cordillera of Colombia. Evaluation methods included: fine litter collection at fortnightly intervals using litter traps; the litter layer samples at the end of each sampling year and chemical analyses of both litterfall and standing litter. Fine litter fall observed was similar in oak forest (7.5Mg ha/y) and in pine (7.8Mg ha/y), but very low in cypress (3.5Mg ha/y). Litter standing was 1.76, 1.73 and 1.3Mg ha/y in oak, pine and cypress, respectively. The mean residence time of the standing litter was of 3.3 years for cypress, 2.1 years for pine and 1.8 years for oak forests. In contrast, the total amount of retained elements (N, P, S, Ca, Mg, K, Cu, Fe, Mn and Zn) in the standing litter was higher in pine (115kg/ha), followed by oak (78kg/ha) and cypress (24kg/ha). Oak forests showed the lowest mean residence time of nutrients and the highest nutrients return to the soil as a consequence of a faster decomposition. Thus, a higher nutrient supply to soils from oaks than from tree plantations, seems to be an ecological advantage for recovering and maintaining the main ecosystem functioning features, which needs to be taken into account in restoration programs in this highly degraded Andean mountains. Rev. Biol. Trop. 59 (4): 1883-1894. Epub 2011 December 01.

La caída y descomposición de hojarasca representan los principales ingresos de materia orgánica y nutrientes a los ecosistemas forestales. Se estudiaron la caída y acumulación de hojarasca fina y el retorno de nutrientes, en un robledal (Quercus humboldtii) y en plantaciones de pino (Pinus patula) y ciprés (Cupressus lusitanica) en tierras altas de Colombia. La caída de hojarasca fina fue similar entre el robledal (7.5Mg/ha.a) y el pinar (7.8Mg/ha.a), y muy inferior en el cipresal (3.5Mg/ha.a). El mantillo representó 1.76, 1.73 y 1.3Mg/ha.a en el robledal, pinar y cipresal, respectivamente. El tiempo medio de residencia (TMR) del mantillo siguió la secuencia: cipresal (3.3 años)>pinar (2.1 años)>robledal (1.8 años). La cantidad de nutrientes retenidos en el mantillo siguió la secuencia: pinar (115kg/ ha)>robledal (78kg/ha)>cipresal (24kg/ha). Los menores TMR de nutrientes se presentaron para la hojarasca foliar del robledal, en su mayoría inferiores a 1.0 años. En términos de la función ecosistémica en el robledal los procesos estudiados son muy superiores vía provisión de nutrientes al suelo y regulación de los ciclos biogeoquímicos, aspectos que deben ser considerados previa implementación de programas de repoblamiento forestal.

Carabidae diversity along an altitudinal gradient in a Peruvian cloud forest (Coleoptera).

Carabid beetles were sampled at five sites, ranging from 1500 m to 3400 m, along a 15 km transect in the cloud forest of Manu National Park, Perú. Seasonal collections during a one year period yielded 77 morphospecies, of which 60% are projected to be undescribed species. There was a significant negative correlation between species richness and altitude, with the number of carabid species declining at the rate of one species for each 100 m increase in altitude. The majority of species (70.1 %) were restricted to only one altitudinal site and no species was found at more than three of the five altitudinal sites. Only one genus, Pelmatellus (Tribe Harpalini), was found at all five sites. Active (hand) collections yielded approximately twice as many species per individuals collected than passive (pitfall trap) collections. This study is the first systematic sampling ofcarabid beetles of a high altitude gradient in the cloud forests of southeastern Perú and supports the need to conserve the zone of extremely high biodiversity present on the eastern slopes of the Peruvian Andes.