Effects of light,moisture, temperature,and litter on the regeneration of five tree species in the tropical montane wet forest of Puerto Rico

Field experiments were conducted to investigate the effects of light, moisture, temperature, and litter on the regeneration of two early-, one mid-, and two late-successional tropical tree species. High light and litter seem to be universally good cues for regeneration, increasing seed/seedling survival for all species except for Cecropia (an early-successional species) whose small seeds may not be able to penetrate the litter layer. In addition, the high temperature environment in both artificially shaded and nonshaded areas of a natural gap exhibits less seed loss, an increase in the percent and rate of germination, and an increase in seedling survival for Dacryodes (a late-successional species), than the lower temperature environment under an intact canopy. Low soil water is also a good cue for Dacryodes germination as it is for Prestoea and Cecropia. Finally, the lower temperature environment found under the forest canopy (compared to the natural gap) leads to less seed loss and more germination for Guarea (a mid-successional species). Our results suggest that a good patch for regeneration of many species in this forest, early- as well as late-successional species, would have high light and a litter layer that moderates temperature and moisture extremes. The substantial variation in suitability among regeneration filters and species could: (1) contribute to low establishment success, i.e., most dispersed propagules do not become trees, (2) make it difficult to group species into germination strategies, and (3) make it hard to generalize about a net effect of any specific environmental variable on establishment. We suggest that tropical disturbances should be viewed in terms of their impact on a variety of environmental cues, which may signal germination and impact subsequent growth and survival.     

A Flood Plain Palm Forest in the Luquillo Mountains of Puerto Rico Five Years After Hurricane Hugo1

Long-term studies are needed to understand the dynamics of tropical forests, particularly those subject to periodic disturbances such as hurricanes. We studied a flood plain Prestoea montana palm forest in the Luquillo Mountains of Puerto Rico over a 15-yr period (1980–1995), which included the passage of Hurricane Hugo in September 1989. The passage of the hurricane caused the dominant species to become more dominant and created low instantaneous tree mortality (1% of stems) and reductions in tree biomass (-16 Mg/ha/yr) and density, although not in basal area. Five years after the hurricane, the palm flood plain forest had exceeded its prehurricane aboveground tree biomass, tree density, and basal area. Aboveground tree biomass accumulated at a rate of 9.2 Mg/ha/yr, 76 percent of which was due to palms. Before the hurricane this rate was on the order of 3 Mg/ha/yr. Forest floor litter decreased to prehurricane levels (6.7 Mg/ha), within 5 yr, mostly due to the disappearance of woody litter. Thirteen tree species not represented in the canopy entered the forest by regeneration, and 2 species suffered almost 20 percent/yr mortality over a 5-yr period after the storm (floodplain average of 2%/yr). Delayed tree mortality was twice as high as instantaneous tree mortality after the storm and affected dicotyledonous trees more than it did palms. Regencration of dicotyledonous trees, palms, and tree ferns was influenced by a combination of factors including hydroperiod, light, and space. Redundancy Data Analysis showed that the area near the river channel was the most favorable for plant regeneration. Palm regeneration was higher in locations with longer hydroperiods, while regeneration of dicotyledonous trees was higher in areas with low risk of flooding. This study shows how a periodic disturbance provides long-term opportunities for species invasions and long-term ecosystem response at the patch scale of < 1 ha.     

Soil Phosphorus Fractionation during Forest Development on Landslide Scars in the Luquillo Mountains,Puerto Rico1

Mineral soils from a chronosequence of landslide scars ranging in age from 1 to more than 55 years in a subtropical montane rain forest of eastern Puerto Rico were used to determine the rate at which labile P capital recovers during primary succession. Nine organic and inorganic soil P fractions were measured using the Hedley sequential extraction procedure. Deep soil cores (9 m) from a nearby site were also analyzed to determine the distribution of P fractions below the solum. Litterfall P was measured for two years in the landslide scars to estimate allochthonous litter P inputs, and published precipitation data were used to estimate annual atmospheric inputs of P to the recovering forests. In the upper solum (0–10 cm), organic matter increased with landslide age, as did resin‐Pi, labile P (defined here as resin‐Pi + HCO₃‐Pi + HCO₃‐Po) and total organic P. Occluded P decreased with increasing landslide age. No significant changes in P concentrations or pools were observed in 10 to 35 or in 35 to 60 cm depth intervals across the chronosequence. Labile soil P increased to approximately two‐thirds of the pre‐disturbance levels in the oldest landslide scar (>55 yr). Thus, plants, their associated microflora/fauna, and P inputs from off‐site substantially altered the distribution of soil P fractions during forest recovery. Across the chronosequence, the increase in labile P accumulated in soil and biomass appeared to be greater than the estimated allochthonous inputs from litter and precipitation, indicating that as the forest developed, some occluded P may have been released for use by soil biota. Resin‐Pi and labile P were correlated with soil organic matter content, suggesting, as in other highly weathered soils, organic matter accumulation and turnover are important in maintaining labile P pools. Primary mineral P (apatite) was scarce, even in deep soil cores.     

Decay rate and substrate quality of fine roots and foliage of two tropical tree species in the Luquillo Experimental Forest,Puerto Rico

Decomposition rates, initial chemical composition, and the relationship between initial chemistry and mass loss of fine roots and foliage were determined for two woody tropical species, Prestoea montana and Dacryodes excelsa, over a gradient of sites in two watersheds in the Luquillo Experimental Forest, Puerto Rico. At all locations, fine roots decayed significantly more slowly than foliage during the initial 6 months.Substrate quality of the initial tissue showed marked differences between roots and foliage when using cell wall chemistry, secondary chemistry and total elemental analysis as indices. Quantity of acid detergent fiber (ADF) (non-digestible cell wall fiber) and lignin content were higher for roots than leaves: D. excelsa roots had 55.3% ADF and 28.7% lignin while leaves had 36.2% ADF and 11.8% lignin; P. montana roots had 68.0% ADF and 26.8% lignin while leaves had 48.5% ADF and 16.1% lignin. Aluminum concentrations were higher in fine roots (843 mg kg^–1 in D. excelsa, 1500 mg kg^–1 in P. montana) than leaves (244 mg kg^–1 in D. excelsa, 422 mg kg^–1 in P. montana), while calcium concentrations were higher in foliage (5.5 mg g^–1 in D. excelsa, 7.8 mg g^–1 in P. montana) than roots (3.4 mg g^–1 in D. excelsa, 3.1 mg g^–1 in P. montana). Nitrogen did not show any trend with tissue or species type. A linear model between mass remaining after 6 months and initial tissue chemistry could be developed only for calcium (r²=0.64).