Maaike Y. Bader

High solar radiation hinders tree regeneration above the alpine treeline in northern Ecuador

Many tropical alpine treelines lie below their climatic potential, because of natural or anthropogenic causes. Forest extension above the treeline depends on the ability of trees to establish in the alpine environment. This ability may be limited by different factors, such as low temperatures, excess solar radiation, competition, soil properties, dispersal ability, and fires. In this paper we address the following two questions: Do trees regenerate above the present treeline, and what are the inhibiting factors for tree establishment? To answer these questions we described the spatial pattern of recent tree establishment below and above the present treeline in northern Ecuador. Also, we experimentally transplanted seedlings into the alpine vegetation (páramo) and the forest, and investigated the effect of shade, neighboring plants, and substrate on their survival. The number of naturally occurring tree sprouts (seedlings, saplings and ramets) was highest just outside the forest, and decreased with distance to the forest edge. However, only two species that were radiation-tolerant made up these high numbers, while other species were rare or absent in the páramo. In the forest, the species diversity of sprouts was high and the abundance per species was relatively low. The transplanted seedlings survived least in experimental plots without artificial shade where neighboring plants were removed. Seedling survival was highest in artificially shaded plots and in the forest. This shade-dependence of most tree species can strongly slow down forest expansion toward the potential climatic treeline. Due to the presence of radiation-tolerant species, the complete lack of forest expansion probably needs to be ascribed to fire. However, our results show that natural processes can also explain both the low position and the abruptness of tropical treelines.

Mountain Treelines: a Roadmap for Research Orientation

Abstract For over 100 years, mountain treelines have been the subject of varied research endeavors and remain a strong area of investigation. The purpose of this paper is to examine aspects of the epistemology of mountain treeline research—that is, to investigate how knowledge on treelines has been acquired and the changes in knowledge acquisition over time, through a review of fundamental questions and approaches. The questions treeline researchers have raised and continue to raise have undoubtedly directed the current state of knowledge. A continuing, fundamental emphasis has centered on seeking the general cause of mountain treelines, thus seeking an answer to the question, “What causes treeline?” with a primary emphasis on searching for ecophysiological mechanisms of low-temperature limitation for tree growth and regeneration. However, treeline research today also includes a rich literature that seeks local, landscape-scale causes of treelines and reasons why treelines vary so widely in three-dimensional patterns from one location to the next, and this approach and some of its consequences are elaborated here. In recent years, both lines of research have been motivated greatly by global climate change. Given the current state of knowledge, we propose that future research directions focused on a spatial approach should specifically address cross-scale hypotheses using statistics and simulations designed for nested hierarchies; these analyses will benefit from geographic extension of treeline research.

Vegetation Structure and Temperature Regimes of Tropical Alpine Treelines

ABSTRACT Alpine treeline ecotones can be gradual transitions, abrupt boundaries, or patchy mosaics, and these different patterns may indicate important processes and dynamic properties. We present observed spatial patterns of a wide range of tropical treelines and try to explain these patterns. Treelines were studied at seven sites in the tropical and subtropical Andes (Argentina, Bolivia, Ecuador, and Venezuela) and on a Hawaiian volcano (Haleakala, Maui). Treeline vegetation structure was described using transects perpendicular to the treeline, and air and soil temperatures were measured above and below the forest boundary. Temperature fluctuations were much larger and the average temperature was higher in alpine vegetation than in forest. Most treelines were abrupt, with surprisingly similar patterns across a wide geographical range. This abruptness could result from positive feedback processes mediated by the differences in microclimate between forest and páramo. Our data is not conclusive about the relative importance of microclimate as opposed to fire in mediating such feedbacks. However, our extensive set of comparable data from different sites in a large geographical region is an important step toward a better understanding of the nature and dynamics of tropical alpine treelines.

A Simple Spatial Model Exploring Positive Feedbacks at Tropical Alpine Treelines

ABSTRACT Climate change could cause alpine treelines to shift in altitude or to change their spatial pattern, but little is known about the drivers of treeline dynamics and patterning. The position and patterns of tropical alpine treelines are generally attributed to land use, especially burning. Species interactions, in particular facilitation through shading, may also be important for treeline patterning and dynamics. We studied how fire in alpine vegetation and shade dependence of trees may affect the position and spatial pattern of tropical alpine treelines and their response to climatic warming, using a spatial minimal model of tree growth at treeline. Neighboring trees provided shade and protection from fire. The positive feedback that resulted from these neighbor interactions strongly affected the emergent treelines and always reduced the distance and speed of treeline advance after a temperature increase. Our model demonstrated that next to fire, shade dependence of trees can also lead to abrupt treelines and relatively low treeline positions. This implies that these patterns do not necessarily indicate human disturbance. Strong abruptness of a treeline may indicate that it will respond slowly to climatic changes.

Altitudinal changes in temperature responses of net photosynthesis and dark respiration in tropical bryophytes

BACKGROUND AND AIMS There is a conspicuous increase of poikilohydric organisms (mosses, liverworts and macrolichens) with altitude in the tropics. This study addresses the hypothesis that the lack of bryophytes in the lowlands is due to high-temperature effects on the carbon balance. In particular, it is tested experimentally whether temperature responses of CO(2)-exchange rates would lead to higher respiratory carbon losses at night, relative to potential daily gains, in lowland compared with lower montane forests. METHODS Gas-exchange measurements were used to determine water-, light-, CO(2)- and temperature-response curves of net photosynthesis and dark respiration of 18 tropical bryophyte species from three altitudes (sea level, 500 m and 1200 m) in Panama. KEY RESULTS Optimum temperatures of net photosynthesis were closely related to mean temperatures in the habitats in which the species grew at the different altitudes. The ratio of dark respiration to net photosynthesis at mean ambient night and day temperatures did not, as expected, decrease with altitude. Water-, light- and CO(2)-responses varied between species but not systematically with altitude. CONCLUSIONS Drivers other than temperature-dependent metabolic rates must be more important in explaining the altitudinal gradient in bryophyte abundance. This does not discard near-zero carbon balances as a major problem for lowland species, but the main effect of temperature probably lies in increasing evaporation rates, thus restricting the time available for photosynthetic carbon gain, rather than in increasing nightly respiration rates. Since optimum temperatures for photosynthesis were so fine tuned to habitat temperatures we analysed published temperature responses of bryophyte species worldwide and found the same pattern on the large scale as we found along the tropical mountain slope we studied.

Differences in desiccation tolerance do not explain altitudinal distribution patterns of tropical bryophytes

Abstract Bryophyte biomass and diversity vary strongly with altitude in the tropics. Low abundance and low species numbers in lowland rain forests are most likely due to reduced diurnal activity times combined with high nocturnal respiration rates at high temperatures. This may exclude many montane species from the warm lowlands. However, an alternative hypothesis explains the observed pattern, namely a limited desiccation tolerance of montane species, precipitation being more concentrated but less frequent in most lowland forests compared to montane cloud forests. To test this hypothesis, we studied the desiccation tolerance of four montane and four lowland bryophyte species. The effects of prolonged drought were quantified with chlorophyll fluorescence (Fv/Fm) and the extent of electrolyte leakage. Both montane and lowland species survived dry periods of ≧80 days, which far exceeds the duration of dry periods in the wet lowland tropics. We can thus exclude intolerance to long dry spells as an explaination for the absence of the tested montane species in the lowlands. We should continue to focus on other mechanisms to explain the altitudinal gradient of bryophyte abundance and diversity in the tropics, in order to understand this pattern, as well as to predict future trends under climatic warming.

Early establishment of trees at the alpine treeline: idiosyncratic species responses to temperature-moisture interactions

Alpine treelines globally may move upslope due to climatic warming. Such movement would need, as the first steps, seed germination and seedling establishment above current treelines. These processes were studied experimentally in five common European treeline tree species. Surprisingly, each species responded very differently to moisture and temperature gradients, with positive and negative responses possible. These results match the heterogeneity observed in treeline dynamics and spatial patterns globally. They strongly emphasize the need for species-specific parameterisations in predictive models of treeline responses to climatic change.

Physiological Ecology of Tropical Bryophytes

Bryophytes in the tropics occur from cool alpine grasslands to warm lowland sites and from cloud forests to dry forests, varying markedly in abundance and diversity in these habitats. This chapter deals with the current knowledge of the ecophysiology of tropical bryophytes attempting to explain some of these abundance patterns, in particular the marked increase in bryophyte biomass with altitude in rain and cloud forests. As data are scarce, we include data on, physiologically rather similar, lichens in our account where appropriate. We focus mostly on carbon relations, and water, nutrients, light, CO2 and temperature are discussed as co-determinants of the carbon balance. In particular, we address the hypothesis that the surprisingly low bryophyte abundance in lowland rainforests is due to the limitation of net carbon gain by fast drying and low light levels during the day combined with moist and warm conditions at night, which promote high respiration rates. The timing of hydration is crucial in determining this diel balance between photosynthesis and respiration. Temperature is important in determining moisture loss rates and nocturnal carbon loss through respiration – if respiration does not acclimatize to higher temperatures. Since carbon balance precariously depends on daily hydration patterns, future climate change may pose a serious problem to tropical lowland bryophytes.

The temperature acclimation potential of tropical bryophytes.

Bryophyte biomass and diversity in tropical moist forests decrease dramatically from higher altitudes towards the lowlands. High respiratory carbon losses at high temperatures may partly explain this pattern, if montane species are unable to acclimatise their metabolic rates to lowland temperatures. We transplanted ten bryophyte species from two altitudes (1200 and 500 m a.s.l.) to lower (warmer) altitudes (500 m and sea level) in Panama. We studied short-term temperature acclimation of CO2 exchange for 2.5 months, and survival and growth for 21 months following transplantation. Short-term acclimation did not occur, and on a longer time scale mortality was highest and growth lowest in the transplanted samples. A few transplanted samples of most species, however, survived the whole experiment and finished with growth rates similar to controls. This recovery of growth rate suggests temperature acclimation, in spite of no measurable metabolic changes in smaller random samples. This acclimation even compensated for shorter periods of CO2 uptake due to more rapid drying. Nevertheless, these species are not abundant in lowland forests, perhaps due to dispersal or establishment limitation. The apparent heterogeneity of the acclimation potential within species may allow populations to adapt locally and avoid being forced uphill under climatic warming.

How to minimize the sampling effort for obtaining reliable estimates of diel and annual CO 2 budgets in lichens

Estimating carbon budgets for poikilohydric organisms, such as lichens and bryophytes, requires methods other than those for homoiohydric plants due to a strong dependency of carbon gain on fluctuating hydration. This paper provides guidance with respect to optimal sampling strategies for estimating annual carbon budgets of lichens and bryophytes, based on a one-year dataset of half-hourly CO 2 -exchange readings on the epilithic placodioid lichen Lecanora muralis (syn. Protoparmeliopsis muralis ) and tests the effects of reduced sampling frequencies and different temporal sampling schemes on carbon budget estimates. Both fine-scale sampling (measurements within a day) and large-scale sampling (selection of days within a year) are addressed. Lowering the sampling frequency within a day caused large deviations for 24-h (diel) budget estimates. Averaged over a larger number of days, these errors did not necessarily cause a large deviation in the annual budget estimate. However, the occurrence of extreme deviations in diel budgets could strongly offset the annual budget estimate. To avoid this problem, frequent sampling ( c . every 1·5 hours) is necessary for estimating annual budgets. For estimating diel budgets and patterns a more frequent sampling (every c . 0·5 hours, balancing data resolution and disturbance) is often needed. Sampling fewer than 365 days in a given year inevitably caused estimates to deviate from the ‘true’ carbon budget, i.e. the annual budget based on half-hourly measurements during 365 days. Accuracy increased with total sample frequency, and blocking days caused larger deviations than sampling randomly or regularly spaced single days. Restricting sampling to only one season led to strongly biased estimates. The sampling effort required for a reliable estimate of the annual carbon balance of lichens based on simple extrapolations of diel carbon budgets is impracticably large. For example, a relatively large sample of 52 random days yielded an estimate within 25% of the true annual budget with only 60% certainty. Supporting approaches are therefore suggested, in particular extrapolating diel budgets using ‘weather response types’, possibly aided by diel activity patterns from chlorophyll fluorescence, or modelling CO 2 exchange as a function of climatic conditions.