The effect of substrate abundance in the vertical stratification of bromeliad epiphytes in a tropical dry forest (Mexico)

Diversity of epiphytes is associated with niche partitioning, through vertical strata and host preferences. However, abundance of substrate offered by hosts differs between vertical strata, misleading if epiphytes prefer a stratum or are randomly distributed. In a tropical dry forest of San Andres de la Cal Morelos, central Mexico, we tested the null hypothesis, that epiphytes follow the abundance of the substrate, rather than showing preference for a particular vertical stratum, and tested whether microclimatic variables, seed germination and seedling survival match with observed epiphyte distribution. Our data show that epiphytes could be randomly distributed inside some host; but in some host species, certain structures presented either a deficit or an excess of all, atmospheric, or tank epiphytes. In the hosts Bursera copallifera and Bursera glabrifolia, distribution of epiphytes was biased towards the upper strata, with a deficit of epiphytes in the lower strata. In Conzattia multiflora, Sapium macrocarpum and Ipomoea pauciflora, epiphyte distribution was biased towards the lower strata. Vertical gradation of light, seed germination and seedling survival did not generally match with epiphyte distribution and did not support the notion that the microclimatic gradient governs the vertical distribution of epiphytes. Our data indicate that vertical distribution of epiphytes in such tropical dry forests is mainly driven by the distribution of the structures, which apparently influence dispersion of the seeds and by the lifespan of branches, which allow the concentration of epiphytes in the stratum that optimizes seed capture and the clonal growth of epiphytes.     

Water uptake and transport in lianas and co-occurring trees of a seasonally dry tropical forest

Water uptake and transport were studied in eight liana species in a seasonally dry tropical forest on Barro Colorado Island, Panama. Stable hydrogen isotope composition (D) of xylem and soil water, soil volumetric water content (_v), and basal sap flow were measured during the 1997 and 1998 dry seasons. Sap flow of several neighboring trees was measured to assess differences between lianas and trees in magnitudes and patterns of daily sap flow. Little seasonal change in _v was observed at 90–120 cm depth in both years. Mean soil water D during the dry season was –19 at 0–30 cm, –34 at 30–60 cm, and –50 at 90–120 cm. Average values of xylem D among the liana species ranged from –28 to –44 during the middle of the dry season, suggesting that water uptake was restricted to intermediate soil layers (30–60 cm). By the end of the dry season, all species exhibited more negative xylem D values (–41 to –62), suggesting that they shifted to deeper water sources. Maximum sap flux density in co-occurring lianas and trees were comparable at similar stem diameter (DBH). Furthermore, lianas and trees conformed to the same linear relationship between daily sap flow and DBH. Our observations that lianas tap shallow sources of soil water at the beginning of the dry season and that sap flow is similar in lianas and trees of equivalent stem diameter do not support the common assumptions that lianas rely primarily on deep soil water and that they have higher rates of sap flow than co-occurring trees of similar stem size.     

Light microhabitats,growth and photosynthesis of an epiphytic bromeliad in a tropical dry forest

In the tropical dry forest of Dzibilchaltún, Yucatan, Mexico, light microhabitats, tissue acidity changes, chlorophyll fluorescence and growth were investigated for the epiphytic bromeliad Tillandsia brachycaulos Schltdl. Most individuals occurred in sites between 20 and 60% of the height of the tree, and directly on the main trunk or branches closed to the main trunk. During the dry season, individuals received about nine times more photon flux density (PFD) than during the rainy season. Individuals that occurred under 30–59% of the daily ambient PFD showed greater leaf length increase and monthly leaf production, and produced more flowers than individuals in the brighter (>60% of ambient PFD) and the darker (11–29% of ambient PFD) light microhabitats. Well-watered plants acclimated to deep shade (6% of ambient PFD) showed values of tissue acidity similar to plants under higher light treatments. Well-watered plants under 60% of daily ambient PFD showed low values of maximum quantum efficiency at midday during most of the year but showed similar values of tissue acidity than plants under 6 and 30% of ambient PFD. Leaf temperatures of plants in low sites within the canopy and on the forest floor were significantly higher than the air during about 1 h, indicating that those plants could not dissipate the excess of heat. Individuals located in the most shaded microhabitats could also receive less amount of water from rainfall and dew. Therefore, it would be less damaging for this epiphytic bromeliad to occur in the most exposed microhabitats than in the most shaded microhabitats in this tropical dry forest.     

Genetic effects of tank-forming bromeliads on the associated invertebrate community in a tropical forest ecosystem

Within the area of community genetics there is an expanding volume of literature demonstrating how within-species genetic variation in temperate trees can have important effects on structuring animal and plant communities. The influence of intraspecific genetic variation on associated communities in relatively more complex ecosystems is only starting to be appreciated. Within tropical forests, epiphytic bromeliad plants often grow high in the canopy and create unique nutrient-rich microhabitats on which many invertebrate and vertebrate species depend. We investigated the influence of within-species genetic variation in the bromeliad Aechmea bracteata on the invertebrate microhabitat community. We found that more genetically similar bromeliad plants were host to more similar communities of juvenile-stage invertebrates, but not adult invertebrates. We discuss possible mechanisms for this, including differential survival and active female oviposition choice. Our work shows that the impact of within-species genetic variation on associated ecological communities may be more general than previously considered. These results agree with recent research suggesting that within-species genetic variation may perform a supporting ecosystem service for maintaining community and ecological processes.     

Oxygen isotope ratio stratification in a tropical moist forest

Summary Oxygen isotope ratios were determined in leaf cellulose from two plant species at Barro Colorado (Republic of Panama) in 4 different plots, two of which were undergoing an irrigation treatment during the dry season. There is a gradient in ¹⁸O values of leaf cellulose from the understory to canopy leaves, reflecting the differences in relative humidity between these two levels of the forest. This gradient is most pronounced in irrigated plots. For irrigated plots there was a highly significant correlation between ¹⁸O and ¹³C values, which was not observed in control plots. This relationship can be explained by humidity controlling stomatal conductance. Low humidity affects ¹⁸O values of leaf water during photosynthesis, which isotopically labels cellulose during its synthesis. Low humidity also decreases stomatal conductance, which affects discrimination against carbon-13 by photosynthetic reactions, thus affecting the ¹³C values of photosynthates. WUE values calculated by using plant carbon and oxygen isotope ratios were similar to those observed with gas exchange measurements in other tropical and temperate area. Thus the concurrent analysis of carbon and oxygen isotope ratios of leaf material can potentially be useful for long term estimation of assimilation and evapotranspiration regimes of plants.     

Generous-like flowers: nectar production in two epiphytic bromeliads and a meta-analysis of removal effects

Animal-pollinated angiosperm plants that respond positively to nectar removal by replenishment invest energy that can entail a reproductive cost. We investigated whether or not nectar removal stimulates replenishment in two hummingbird-pollinated bromeliad species. Nectar replenishment rates were also assessed by removing nectar from manually pollinated flowers because pollination events might be used as signals to save energy by preventing allocation to post-pollination nectar production. Then we synthesized the current understanding of nectar removal effects by reviewing existing published studies with a meta-analysis. The magnitude and significance of estimated nectar removal effects and factors associated with variation in size and direction of nectar removal effects were elucidated with the meta-analysis. We found that both Tillandsia species strongly respond to repeated nectar removal by producing >3 times additional nectar. Nectar secretion patterns were not altered by pollination (stigmatic pollen deposition) and we found no evidence of nectar reabsorption. Although the effect size varied widely across systems and/or environmental conditions, the meta-analysis showed that nectar removal had overall a positive effect on nectar replenishment (mainly among species inhabiting wet tropical habitats such as Tillandsia), and a negative effect on the secretion of additional sugar, suggesting that those plants are resource limited and conservative in the secretion of additional sugar.Electronic Supplementary Material Supplementary material is available for this article at     

Effect of canopy position on germination and seedling survival of epiphytic bromeliads in a Mexican humid montane forest

BACKGROUND AND AIMS: Seeds of epiphytes must land on branches with suitable substrates and microclimates to germinate and for the resulting seedlings to survive. It is important to understand the fate of seeds and seedlings in order to model populations, but this is often neglected when only established plants are included in analyses. METHODS: The seeds of five bromeliad species were exposed to different canopy positions in a Mexican montane forest, and germination and early seedling survival were recorded. Additionally, the survival of naturally dispersed seedlings was monitored in a census over 2.5 years. Survival analysis, a procedure rarely used in plant ecology, was used to study the influence of branch characteristics and light on germination and seedling survival in natural and experimental populations. KEY RESULTS: Experimental germination percentages ranged from 7.2 % in Tillandsia deppeana to 33.7 % in T. juncea, but the seeds of T. multicaulis largely failed to germinate. Twenty months after exposure between 3.5 and 9.4 % of the seedlings were still alive. There was no evidence that canopy position affected the probability of germination, but time to germination was shorter in less exposed canopy positions indicating that higher humidity accelerates germination. More experimental seedlings survived when canopy openness was high, whereas survival in census-seedlings was influenced by moss cover. While mortality decreased steadily with age in juveniles of the atmospheric Tillandsia, in the more mesomorphic Catopsis sessiliflora mortality increased dramatically in the dry season. CONCLUSIONS: Seedling mortality, rather than the failure to germinate, accounts for the differential distribution of epiphytes within the canopy studied. With few safe sites to germinate and high seedling mortality, changes of local climate may affect epiphyte populations primarily through their seedling stage.     

Studies on epiphytic flora of a tropical rain forest in Southwestern Nigeria

A survey of epiphytes in Gambari Forest Reserve in Southwestern Nigeria was undertaken. A total of 26 epiphytes were recorded. These include five pteridophytes and 21 angiosperm species belonging to the following families; Acanthaceae, Araceae, Asteraceae, Commelinaceae, Moraceae, Orchidaceae, Palmae, Urticaceae Piperaceae, and Portulacaceae. The smooth-barked phorophytes had only one or two epiphytes on them because they are unable to accumulated dust, debris and moisture for germination and growth of the epiphytes. The palm trees with rough barks had points at which they can collect soil, nutrients and moisture for epiphytic growth, hence all the epiphytic species recorded on the site were present on palm trees. Presence of many of the epiphytes is thought to be due to the availability of propagules from reproducing adults in the surrounding vegetation. Also, epiphytes appear to have a dressing effect on the palm trees.     

Studies on epiphytic flora of a tropical rain forest in Southwestern Nigeria

Bark microflora of 12 rainforest trees were examined. Crustose, foliose, and fruticose forms of lichens were observed. Bacillus sp., Erwinia sp., Micrococcus sp., Proteus sp., and Pseudomonas sp. were the five genera of bacteria recorded while three genera of Cyanobacteria; Entophysalis, Gleocapsa and Stigonema were observed. In the algal division, three genera of Chlorophyta found are Chlorococcum, Pleurococcus and Physolinum. Only one diatom, Cocconeis sp. was present on just one tree. Most of the epiphytic microflora found are mainly terrestrial and their spores might have been deposited on the barks by air, or dust-currents. While moisture is regarded as the most critical factor in growth of bark microflora, other factors like texture of the bark, insolation and especially chemical characteristics of the barks may play a key role in occurrence and distribution of micro-organisms on them.     

Contrasting patterns of vertical stratification in two moth families in a Costa Rican lowland rain forest

The vertical stratification of two species-rich moth families (Lepidoptera: Arctiidae, Geometridae) was analysed in a lowland rain forest in Costa Rica. Moths were trapped with automatic 8 W ultraviolet light traps at three sites in the understorey and canopy of a primary forest at the La Selva Biological Station (10.4° N, 84.0° W). A total of 846 arctiid moths (148 species) and 946 geometrid moths (140 species) were analysed. Species richness and diversity of arctiid moths was significantly lower in the understorey than in the canopy (Fisher's alpha: 24 vs. 49) whereas geometrid moths showed an inverse pattern (Fisher's alpha: 44 vs. 27). This resulted in an overall increase in the proportion of conspicuously coloured species towards the canopy. Moth ensembles were clearly separated in multidimensional scaling ordinations, and differed significantly in their faunal composition and dominance between the strata. The available host plant data suggest that the flight height of moths was determined by larval resource availability. Examples include understorey flyers such as the geometrid genus Eois feeding on Piper, and canopy flyers such as the arctiid moth genera Aclytia, Macrocneme and Poliopastea which feed on lianas.     

Highly efficient uptake of phosphorus in epiphytic bromeliads

Background and Aims

Vascular epiphytes which can be abundant in tree crowns of tropical forests have to cope with low and highly intermittent water and nutrient supply from rainwater, throughfall and stem flow. Phosphorus rather than nitrogen has been suggested as the most limiting nutrient element, but, unlike nitrogen, this element has received little attention in physiological studies. This motivated the present report, in which phosphate uptake kinetics by leaves and roots, the subsequent distribution within plants and the metabolic fate of phosphate were studied as a step towards an improved understanding of physiological adaptations to the conditions of tree canopies.

Methods

Radioactively labelled [³²P]phosphate was used to study uptake kinetics and plant distribution of phosphorus absorbed from bromeliad tanks. The metabolism of low molecular phosphorus metabolites was analysed by thin-layer chromatography followed by autoradiography.

Key Results

Uptake of phosphate from tanks is an ATP-dependent process. The kinetics of phosphorus uptake suggest that epiphytes possess effective phosphate transporters. The K_m value of 1·05 µm determined for leaves of the bromeliad Aechmea fasciata is comparable with values obtained for the high affinity phosphate transporters in roots of terrestrial plants. In this species, young leaves are the main sink for phosphate absorbed from tank water. Within these leaves, phosphate is then allocated from the basal uptake zone into distal sections of the leaves. More than 80 % of the phosphate incorporated into leaves is not used in metabolism but stored as phytin.

Conclusions

Tank epiphytes are adapted to low and intermittent nutrient supply by different mechanisms. They possess an effective mechanism to take up phosphate, minimizing dilution and loss of phosphorus captured in the tank. Available phosphorus is taken up from the tank solution almost quantitatively, and the surplus not needed for current metabolism is accumulated in reserves, i.e. plants show luxury consumption. Young, developing leaves are preferentially supplied with this nutrient element. Taken together, these features allow epiphytes the efficient use of scarce and variable nutrient supplies.Key words: Epiphytic bromeliads, phosphorus uptake, forest canopies, luxury consumption, phytotelms, plant nutrition, Aechmea fasciata     

Structure and dynamics of a tropical dry forest in Ghana

Forest under low rainfall (averaging 745 mm yr^-1) on the Shai Hills in S.E. Ghana has redeveloped following cessation of farming in the 1890s. Forest stature is low, with a canopy at about 11 m, principally of three species, Diospyros abyssinica, D. mespiliformis and Millettia thonningii. Drypetes parvifolia and Vepris heterophylla are common understorey trees. Twelve species of woody liane were recorded. Species of thicket vegetation in the area were also present at low density. Most species are evergreen.Tree mortality averaged 2.3% yr^-1 and exceeded recruitment (1.5% yr^-1). Differences between species in mortality and recruitment were pronounced: canopy species showed a small decline in density; understorey species increased markedly and the thicket species declined. Seed production was very variable, but seedling establishment was very poor for all species. Seedling mortality was high (11% yr^-1) especially for small seedlings. These population trends probably represent the latter stages of succession of forest regrowth after farming about 100 years ago.Compared with tropical rain forest, Shai Hills forest has similar relative tree diameter growth (1–3.5% yr^-1), mortality and recruitment rates, and small-litter fall (5.52 t ha^-1 yr^-1).Shai Hills forest differs from rain forest by its short stature, relatively few (evergreen) tree species, poor regeneration from seed, high soil nutrient status and low rainfall. Similar forests have been reported in east Africa and in parts of New Guinea.Abbreviations dbh diameter at breast height (1.3 m) - gbh girth at breast height died May 1984     

Genetic variation in a tropical tree species influences the associated epiphytic plant and invertebrate communities in a complex forest ecosystem

Genetic differences among tree species, their hybrids and within tree species are known to influence associated ecological communities and ecosystem processes in areas of limited species diversity. The extent to which this same phenomenon occurs based on genetic variation within a single tree species, in a diverse complex ecosystem such as a tropical forest, is unknown. The level of biodiversity and complexity of the ecosystem may reduce the impact of a single tree species on associated communities. We assessed the influence of within-species genetic variation in the tree Brosimum alicastrum (Moraceae) on associated epiphytic and invertebrate communities in a neotropical rainforest. We found a significant positive association between genetic distance of trees and community difference of the epiphytic plants growing on the tree, the invertebrates living among the leaf litter around the base of the tree, and the invertebrates found on the tree trunk. This means that the more genetically similar trees are host to more similar epiphyte and invertebrate communities. Our work has implications for whole ecosystem conservation management, since maintaining sufficient genetic diversity at the primary producer level will enhance species diversity of other plants and animals.     

Root biomass of a dry deciduous tropical forest in Mexico

The deciduous tropical dry forest at Chamela (Jalisco, Mexico) occurs in a seasonal climate with eight rainless (November through June) and four wet months (700 mm annual precipitation). The forest reaches a mean height of 10 m. Tree density in the research area was 4700 trees per ha with a basal area at breast height of 23 m² per ha. The above-and below-ground biomass of trees, shrubs, and lianas was 73.6 Mg ha^–1 and 31 Mg ha^–1, respectively. A root:shoot biomass ratio of 0.42 was calculated. Nearly two thirds of all roots occur in the 0–20 cm soil layer and 29% of all roots have a diameter of less than 5 mm.     

Seedling recruitment patterns in a tropical dry forest in Ghana

Abstract. Studies of seedling demography and dynamics were carried out in a dry forest at Pinkwae, Ghana (rainfall 1100 mm). Seedlings of tree and liana species in permanent transects were tagged, identified, measured and recensused at bimonthly intervals. A total of 1931 seedlings in 53 species were followed over 14 censuses during a 2-yr period. Germination and mortality were seasonal in their distribution; mortality was highest in dry periods. Seedling density varied seasonally, with the highest densities in comparatively wet periods. Diversity of seedlings was low, corresponding to that of the adult assemblage. Survivorship of tagged seedlings was recorded during the study period. The cohort tagged at the first census, comprising 1033 individuals of mixed ages and species, lost 61.9% during a 24-month period. Many seedlings were found to be rather large at the initial post-germination stage, with strong root development. Vegetative reproduction by root suckers or epicormic shoots was quite common at the site, although the majority of regeneration was by seeds. The current seedling assemblage appears to be representative, in terms of composition and diversity, of the adult species assemblage at Pinkwae.     

Drought effects on seedling survival in a tropical moist forest

The amount and seasonality of rainfall varies strongly in the tropics, and plant species abundance, distribution and diversity are correlated with rainfall. Drought periods leading to plant stress occur not only in dry forests, but also in moist and even wet forests. We quantified experimentally the effect of drought on survival of first year seedlings of 28 co-occurring tropical woody plant species in the understory of a tropical moist forest. The seedlings were transplanted to plots and subjected to a drought and an irrigation treatment for 22 weeks during the dry season. Drought effects on mortality and wilting behavior varied greatly among species, so that relative survival in the dry treatment ranged from 0% to about 100% of that in the irrigated treatment. Drought stress was the main factor in mortality, causing about 90% (median) of the total mortality observed in the dry treatment. In almost half of the species, the difference in survival between treatments was not significant even after 22 weeks, implying that many of the species are well adapted to drought in this forest. Relative drought survival was significantly higher in species associated with dry habitats than in those associated with wet habitats, and in species with higher abundance on the dry side of the Isthmus of Panama, than in those more abundant on the wet side. These data show that differential species survival in response to drought, combined with variation in soil moisture availability, may be important for species distribution at the local and regional scale in many tropical forests.     

Leaf age and the timing of leaf abscission in two tropical dry forest trees

We used experimental defoliations to examine the effect of leaf age on the timing of leaf shedding in two tropical dry forest trees. Trees of the deciduous Bombacopsis quinata (bombacaceae, a.k.a. Pachira quinata) and the brevi-deciduous Astronium graveolens (anacardiaceae) were manually defoliated for three times during the rainy season. All trees started to produce a new crown of leaves 2 weeks after defoliation, and continued expanding leaves throughout the rainy season. At the transition to the dry season, the experimental groups consisted of trees with known differences in maximum leaf age. Defoliations resulted in declines in stem growth but did not affect the mineral content or water relations of the leaves subsequently produced. There was no effect of leaf age on the timing of leaf abscission in B. quinata. In A. graveolens, the initiation of leaf shedding followed in rank order, the maximum leaf age of the four treatments, but there was substantial coherence among treatments in the major period of leaf abscission such that trees completed leaf shedding at the same time. In the two species, leaf water potential (Ψ_L) and stomatal conducantce (g _S) declined with the onset of the dry season, reaching minimum values of –0.9 MPa in P. quinata and <–2.0 MPa in A. graveolens. Within each species, leaves of different age exhibited similar Ψ_L and g _S at the onset of drought, and then decreased at a similar rate as the dry season progressed. Overall, our study suggests that the environmental factors were more important than leaf age in controlling the timing of leaf shedding.