LEAF GROWTH AND PHENOLOGY OF THE DIMORPHIC HERBACEOUS LAYER FERN DANAEA WENDLANDII (MARATTIACEAE) IN A COSTA RICAN RAIN FOREST

Leaf growth and spore release in Danaea wendlandii were monitored under temperate greenhouse conditions and in a rain forest at the La Selva Biological Station in Costa Rica. Sterile leaf (trophophyll) growth occurs in four phases, and fertile leaf (sporophyll) growth occurs in five phases. Trophophylls show an initial rapid increase in length for a 4-week period, attain full maturity at 8 weeks, and remain on the plant for about 3 years. There appears to be a seasonal pattern of trophophyll growth, with crozier emergence rates in May and June three times higher than the mean annual rate. Sporophyll growth shows a double sigmoid pattern. For about 3 weeks after crozier emergence, there is a rapid increase in length. This phase is followed by a 10-week period with no appreciable change in length. Rapid elongation over a 2-week period precedes spore release; this is followed within 2 weeks by disappearance of the sporophyll. Sporophylls in the rain forest are subject to 14% mortality prior to maturation, with most mortality occurring in later stages of growth. A projected phenology based on the sporophyll growth curve shows peaks in spore release late in the wet season with a concomitant peak in sporophyll emergence early in the wet season during the 15-month period from June 1986 through September 1987.     

LEAF OPTICAL PROPERTIES ALONG A VERTICAL GRADIENT IN A TROPICAL RAIN FOREST CANOPY IN COSTA RICA

Leaf optical properties (400–1,100 nm) were compared for four species of rain forest trees with crowns in understory, mid-canopy, and canopy positions to test whether optical properties change with light environment. The species tested represent a spectrum of regeneration patterns ranging from shade tolerant to light demanding. Overall, leaf optical properties of the four species were similar. Differences in absorptance were small, but statistically significant among the species and positions along the canopy gradient. Species absorptance differences corresponded somewhat to shade tolerance; two of the shade species showed higher absorptance in lower light environments, while the sun species showed the reverse pattern. Specific leaf mass (leaf weight per unit area) and chlorophyll content per unit leaf weight also changed along the canopy gradient. Specific leaf mass was positively correlated and chlorophyll per unit leaf weight was negatively correlated with increasing light environment. Consequently, the efficiency of absorption, as represented by the absorptance per unit leaf weight, increased as light level decreased, largely due to changes in specific leaf mass. In contrast, efficiency of absorption per unit leaf chlorophyll was relatively constant with light environment for the two species measured for chlorophyll.     

RELATIONS BETWEEN SUNFLECK SEQUENCES AND PHOTOINHIBITION OF PHOTOSYNTHESIS IN A TROPICAL RAIN FOREST UNDERSTORY HERB

We report the photosynthetic characteristics of a C₃ shade plant native to the tropical rain forest understory. It was shown that Elatostema repens Lour. (Hall) f. (Urticaceae) presents a large light adjustment capacity. The effects of several lightfleck sequences on photoinhibition of photosynthesis and carbon gain are analyzed. Photoinhibition is measured both as a decrease in leaf net CO₂ uptake in limiting light (shown to be linearly correlated to quantum yield of O₂ evolution measured at saturating CO₂) and as a decrease of the ratio of variable fluorescence (Fv) to maximum fluorescence (Fmax) measured in liquid nitrogen. It is shown that lightflecks (from 10 to 30 min in duration) of 700 μmol m^–2 s^–1 (high light) induce photoinhibition, and that the effects of those successive high light periods are additive; there is apparently no recovery from photoinhibition during the low light periods (from 10 to 45 min in duration). In contrast, the Fv/Fmax ratio, though decreasing similarly to quantum yield of net CO₂ uptake on leaves submitted to a continuous illumination of 700 μmol m^–2 s^–1, is only decreased a little on leaves submitted to lightfleck sequences of the same photon flux density. Lightflecks of 250 μmol m^–2 s^–1 are not photoinhibitory. Compared to the control maintained under light growth condition (40 μmol m^–2 s^–1) carbon gain is increased on leaves submitted to lightflecks; this gain remains high throughout the light cycles on leaves submitted to nonphotoinhibitory lightflecks and to the photoinhibitory lightflecks followed by the shortest low light period. In the other cases, carbon gain, higher than that of the control at the beginning of the treatments, decreases and becomes lower than the control carbon gain. Finally, the relevance of photoinhibition in the tropical rain forest understory environment is discussed.     

INTERACTIONS BETWEEN CROWN STRUCTURE AND LIGHT ENVIRONMENT IN FIVE RAIN FOREST PIPER SPECIES

Measurements of light variation among leaves within crowns of five Piper species were compared with estimates of spatial variation in light within understory, forest edge, and clearing habitats to estimate the extent to which crown structure contributes to variation in leaf light environment. Daily photon flux density (PFD) varied greatly within and among crowns. Coefficients of variation for daily PFD among sensors within a single crown ranged from 26 to 79%. Within a single crown located in a clearing, the range in daily PFD among leaves was nearly as great as the range over the entire sample of plants. In the understory, localized sunfleck activity contributed to a high degree of spatial variation in instantaneous and total PFD among leaves within individual crowns. Much of the microsite variation in sunfleck activity, however, reflected environmental conditions within the understory habitat. Within an array of sensors placed next to Piper crowns in the understory, correlations were poor for light sensors spaced only 0.2 m apart, and only 8% of the variance in light readings was explained by measurements made 0.5 m away. In the clearing habitats, microsite heterogeneity among leaves was more strongly influenced by leaf positions within crowns and leaf angles than by spatial heterogeneity within the habitat.     

VARIATION IN SEED SIZE WITHIN A CROP OF A COSTA RICAN MUCUNA ANDREANA (LEGUMINOSAE)

The mature viable seeds within the seed crop of one individual of Costa Rican Mucuna andreana varied in fresh weight from 2.6 to 9.4 g (average of 6.81 g, s.d. = 1.09 g, n = 282). The seed coats weighed 14 to 17 percent of the fresh seed weight and variation in their weight was not the cause of the variation in seed weight. There was no significant difference in average seed weight of seeds from pods with 1 to 5 filled seeds. The great range of seed weights within one plant's seed crop is interpreted as possibly of adaptive significance in generating a more homogeneous seed shadow than would be the case with a narrow range of seed weights.     

WOOD SPECIFIC GRAVITY GRADIENTS IN TROPICAL DRY AND MONTANE RAIN FOREST TREES

Pith to bark specific gravity (SG) trends were investigated in 18 tropical dry forest and six montane rain forest tree species of Costa Rica. Eleven dry forest species showed statistically significant increases in SG with distance from pith. The increases ranged from 20–80%; the greatest changes were exhibited by species which are known to occur in tropical wet as well as tropical dry forests. The other seven species showed no change in SG with distance from pith. Of the montane forest species, one showed a significant decrease of 20%, and three showed significant increases ranging from 20–40%. Two species exhibited no change in SG. Comparison of these changes with trends found in tropical wet forest and temperate forest suggests that the increase in SG with size is most common in tropical wet forest, least common in temperate forest, and intermediate in tropical dry and montane forests.     

WIND STRESS AND ELFIN STATURE IN A MONTANE RAIN FOREST TREE: AN ADAPTIVE EXPLANATION

Physiognomic trends in a population of Didymopanux pittieri Marchal (Araliaceae), a dominant shade-intolerant tree of the elfin forests of Costa Rica, are related to a gradient of wind stress. Wind stress in this forest increases with proximity to ridgecrests. For a given tree height, trunk girth increases with proximity to the ridgecrest. At the same time twig slenderness decreases. These responses are produced in part by slower elongation of twigs which are exposed to stronger winds. These trends suggest that elfin stature is an adaptive response to greater wind stress along exposed ridges.     

ECOLOGICAL CONSTRAINTS ON WOOD DENSITY IN A TROPICAL MONTANE RAIN FOREST

In a Costa Rican tropical lower montane rain forest the wood densities of canopy tree species are related to the windiness of their preferred habitats, and to their abilities to tolerate shade. Shade-intolerant species tend to have less dense wood than shade-tolerant species from the same habitat. Species characteristic of windy sites tend to have denser wood than species characteristic of sheltered habitats. Stand mean wood density, the average of species’ wood densities weighted by their proportional contributions to stand basal area, increases with exposure to the wind. These trends in wood density should at least partially counteract the damaging effects of wind on exposed sites. Since investment in wood must come at the expense of growth elsewhere, such trends in wood density may help explain the small stature of elfin forest and montane thicket formations in tropical mountains.     

SUBSTRATE COMPETITION BETWEEN A SALT MARSH DIATOM AND A BACTERIAL POPULATION1

Cylindrotheca closterium Ehrenberg, a benthic marine diatom, competes successfully with Aeromonas sp. a bacterium from the same environment, for low molecular weight organic substrates when they are presented at natural concentrations (1–10 μM). In short term (1 h) experiments, the uptake of mannose by C. closterium was enhanced in light. Seventy percent of the total uptake of mannose by both species was effected by C. closterium over a 1 h period in light. The diatom population was also competitive in darkness. The algal portion of glucose uptake over 1 h was 71% when both populations were given the substrate initially. Percentages of total amino acid uptake for C. closterium ranged from 33% glycine in light to 73% leucine in darkness when both species were given substrate initially. Cylindrotheca had smaller percentages of glucose and aspartic acid total uptake in competition experiments run to stationary phase (10 days).     

海南岛霸王岭热带雨林的种间分离

基于地理信息系统软件绘制了海南岛霸王岭热带雨林 1个 0 .5hm2 永久样地所有DBH≥ 1cm树木的分布图 ,进而借助最近邻体分析扩展模块判定每个个体的最近邻体植株 ,并得到每个基株 最近邻体种对的距离。采用N×N最近邻体列联表的截表法 ,研究了这个多物种群落的种间分离。结果表明 ,随机毗邻种对占优势 ,正分离种对次之 ,负分离种对最少。多数灌木和一些小乔木 (平均胸径 <1 5cm)彼此呈正分离 ,少数负分离 ;灌木和乔木间的分离关系复杂 ;大乔木种类 (平均胸径≥ 65cm)间不存在负分离。种间分离存在种间差异。种间分离与分布格局明显相关 ,聚集分布的物种与其它物种正分离比例远比随机分布、均匀分布大 ;相反 ,聚集分布物种与其它物种负分离比例要比随机分布、均匀分布小。聚集 聚集分布的种对最可能出现正分离 ;聚集 均匀分布的种对以及随机 随机分布的种对则最可能出现负分离。但是 ,无论物种的分布格局或种对相对分布格局如何 ,随机毗邻的种对都占优势。与种间联结不同 ,种间分离反映的是小尺度的物种空间关系。在此基础上 ,初步阐述了种间分离的可能原因及其生态学涵义     

AUTOGAMY AND COMPETITION FOR POLLINATORS IN HEPATICA AMERICANA (RANUNCULACEAE)

Hepatica americana is one of the earliest flowering vernal herbs in the deciduous forests of piedmont North Carolina. Unlike most other members of the spring wildflower community, its flowers are nectarless and autogamous. The number of insect visits a flower receives varies markedly through the blooming season and is lowest when H. americana is in full bloom. Potential for outcrossing as measured by the seed set of emasculated flowers follows the same pattern. The mid-season decline in insect-pollination is most likely the result of competition with Erythronium umbilicatum, which flowers concurrently with H. americana and is much more attractive to the solitary bees that are the principal early season pollen vectors. Because of this competitive disadvantage, seed set of H. americana in piedmont North Carolina would be pollination-limited if the flowers were not autogamous. Autogamy may also be favored by flowering early in the spring before pollinators are available. Opportunities for outcrossing in H. americana are primarily the result of protogyny and blooming earlier than competitors.