The Landscape Ecology of Tropical Secondary Forest in Montane Costa Rica

Multinomial logistic models of land use/land cover in montane Costa Rica and landscape pattern analysis showed that relative to agriculture, secondary forest occurred closer to old-growth forest, further from roads, in forest reserves, and at higher elevations. Collinearity between explanatory variables yielded simple multivariate models; proportion of surrounding old growth predicted secondary forest most accurately. An old-growth matrix [mean patch size (MPS) 24.5 ha], located mainly within protected areas, dominated elevations greater than 2500 m. A matrix of agriculture (MPS 23.5 ha), with smaller patches (approximately 9 ha) of secondary forest and old growth, dominated elevations from 1500 to 2500 m. Combining secondary forest with old growth decreased forest patch number and increased MPS from 7.3 to 37.1 ha. I concluded that: (a) secondary forest pattern is nonrandom, so ancillary data will aid its mapping with satellite imagery. The variables elevation, agriculture distance, road distance, and population density distinguished secondary forest from old growth with 74% accuracy; (b) socioeconomic and biological forces probably interact to create these secondary forest patterns; and (c) the strong association between secondary forest and old growth supports the concept that tropical forest recovery depends on the landscape structure of remnant forest. Received 16 February 1999; accepted 20 August 1999.     

Tree Seed Fate in a Logged and Fragmented Forest Landscape, Northeastern Costa Rica1

We compared the seed fate of two animal‐dispersed, large‐seeded timber species (Dipteryx panamensis [Fabaceae] and Carapa guianensis [Meliaceae]) in logged and fragmented forests with that for continuous forest in northeastern Costa Rica. For both species, we quantified rates of seed removal (an index of vertebrate predation) and the fate of dispersed seeds (those carried away from their original location that either germinated or were not subsequently removed within three months). We predicted that (1) fewer seeds would be dispersed by vertebrates in fragmented forest than in continuous forest due to low population abundances after hunting and/or loss of suitable habitat, and (2) seed predation rates would be higher in forest fragments than in continuous forest due to high abundance of small‐bodied seed consumers. We compared three forest fragments currently managed for timber (140–350 ha) and a large reserve of continuous forest (La Selva, 1500 ha and connected to a national park). An exclusion experiment was performed (seeds placed in the open vs. seeds within semipermeable wire cages; 5 cm mesh size) to evaluate the relative roles of large and small animals on seed removal. Seed germination capacity did not differ among all four sites for both species. Removal of Dipteryx seeds was higher in forest fragments (50% removal within 10 days and related to the activity of small rodents) compared to La Selva (50% removal after 50 days). Also, more Dipteryx seeds were dispersed at La Selva than in fragmented forests. Contrary to our predictions, removal of Carapa seeds was equally high among all four sites, and there was a trend for more seeds of Carapa to be dispersed in fragments than in La Selva. Our results suggest that fragmentation effects on tree seed fate may be specific to species in question and contingent on the animal biota involved, and that management strategies for timber production based on regeneration from seed may differ between forest patches and extensive forests.     

Long-Term Effects of Forest Regrowth and Selective Logging on the Seed Bank of Tropical Forests in NE Costa Rica1.

To investigate long-term effects of land use on the soil seed bank, we compared the abundance/density, species richness, life form distribution, and species composition of seeds stored in the soil of four 15–20 yr-old second-growth stands, two old-growth stands, and two previously selectively-logged stands in the Caribbean lowlands of Costa Rica. Surface soil (10 cm deep, 4.7 cm diameter) was collected at 10 m intervals along three 120–160 m long transects in each stand (44–48 soil cores, 22–24 combined seed bank samples per site). Seed density was highest but variable in second-growth stands (8331–14535 seeds/m²), low and homogeneous in old-growth stands (2258–2659 seeds/m²), and intermediate and highly variable in selectively-logged stands (1165–6854 seeds/m²), which also had contrasting logging intensities. Species richness was strongly dependent on seed density, but showed less variation. Life form distribution did not differ statistically among or within land-use categories. In each stand, herbs-forbs, shrubs, and vines dominated the seed bank (> 75% of the species richness and abundance), whereas trees were a minor component (< 20% of the species richness and < 5% of the abundance) and were predominandy early successional. Shrubs and vines were most abundant in second-growth stands where regrowth vegetation was repeatedly cut before abandonment, whereas grasses and sedges were most abundant in the only forest stand that was completely surrounded by pastures. In terms of species composition, old-growth stands were more similar to selectively-logged stands than to second-growth stands, but across stands, selectively-logged forests were most distinct from the other two forest types. An inventory of the standing woody vegetation in each site showed little representation of the woody taxa found in the seed bank. We discuss these results in the context of the main factors that have been postulated to influence the abundance, life form, and species composition of tropical forest seed banks, and explore the role of the latter during intermediate phases of tropical forest succession and regeneration.     

Seed Dispersal by Salvin's Curassow,Mitu salvini (Cracidae), in a Tropical Forest of Colombia: Direct Measurements of Dispersal Distance1

Seed dispersal by Salvin's curassows (Mitu salvini) was studied in a lowland tropical forest at La Macarena on the border of the Macarena and Tinigua National Parks, the Department of Meta, Colombia. Continuous observations were made on the feeding and ranging behavior of a well-habituated pair of birds from 0600 to 1800 h for 19 days. Ibtal observation time was 222 h. The daily distance traveled by the birds ranged from 630 to 3750 m, with a mean of 1959 m (± 776 m). The total home range was ca 34.4 ha. Although curassows were observed consuming fruits of 13 species belonging to the families Rubiaceae, Meliaceae, Moraceae, Burseraceae, Leguminosae, and Lecythidaceae, only seeds of Geophila re fens (Rubiaceae) and Picas spbenophylla (Moraceae) were found in their feces. For G. repens, the mean and maximum retention times were 1 h 52 min (± 1 h 20 min) and 6 h 08 min, and the mean and maximum direct dispersal distances were 245 m (± 164 m) and 633 m. More than half the seeds were dispersed in canopy gaps. For F. sphenophylla, the mean and maximum retention times were 3 h 1 5 mm (± .37 min) and 7 h 08 min, and the mean and maximum direct dispersal distances were 329 m (± 46 m) and 451 m. Nearly 60 percent of the estimated seed intake of G. repens and 92-94 percent of F. sphenophylla were digested or damaged. The retention times and the dispersal distances for Streptogyna americana, which has adhesive burrs, were also measured. The mean and maximum retention times were 1 h 55 min (± 1 h 56 min) and 9 h 11 min, and the mean and maximum direct dispersal distances were 128 m (± 68 m) and 280 m. This is the first study in which direct measurements were made for retention times and dispersal distances of the epizoochory and endozoochory for birds in the field.     

Species Richness,Spatial Variation,and Abundance of the Soil Seed Bank of a Secondary Tropical Rain Forest1

Despite the importance of the soil seed bank in tropical forest regeneration, little is known about spatial variability in species composition and abundance of seeds stored in the soil. To develop sampling methods for comparative studies, we examined species richness, spatial variation, and abundance of germinants from the soil seed bank in a 16 year old secondary, tropical wet forest at La Selva Biological Station, Costa Rica. Surface soil (10 cm deep, 4.7 cm diameter) was collected at the intersection points of a gridded 1 ha plot (10 × 10-m grid, 121 samples) and in a nested 100 m² subplot (2 × 2-m grid, 36 samples). The 1 ha plot had a density of 4535 seeds/m² with 34 species observed. Based on a series of 100 randomized species accumulation curves, a Michaelis-Menten fit predicted a mean species richness of 36.3 species; the number of observed species was close to the predicted asymptote. A nonparametric, first-order jackknife species richness estimator predicted a species richness of 37.0 species. Eighty-five and 95 percent of the observed species richness is contained, on average, within 41 and 74 pooled samples, respectively. Within the 100 m² nested subplot, a density of 5476 seeds/m² was observed, comprising 26 species with an estimated species richness (Michaelis-Menten fit) of 29.1 species. The jackknife species richness estimator predicted a species richness of 36.7 species. For species richness and abundance of both plots, spatial autocorrelation statistics (Moran's I) were not significantly different from zero at lag distances from 2 to 100 m, indicating a random distribution at these spatial scales. For this site, accurate estimates of species composition depend upon the number of samples collected as well as the spatial distribution of sampling effort. Many small samples distributed over a large area provide greater accuracy and precision for estimating species richness of the soil seed bank.     

Factors Limiting Tropical Rain Forest Regeneration in Abandoned Pasture: Seed Rain,Seed Germination,Microclimate, and Soil1

Vegetation, seed rain, seed germination, microclimate, and soil physical and chemical parameters were measured in a recently abandoned pasture and adjacent primary rain forest in southern Costa Rica. The goal of this study was to assess the importance of these factors in limiting forest regeneration in abandoned pastures. Seed rain of animal dispersed species decreased dramatically in the pasture >5 m from the forest/pasture edge; fewer wind dispersed seeds fell in the pasture than in the forest, but the difference was much less than for animal dispersed seeds. Percent seed germination of most species studied was similar in the forest and in pasture with grasses; seed germination was lower during the dry season in areas of pasture cleared of grasses. Air temperature, vapor pressure deficit (VPD), and photon flux density (PFD) were much higher in the pasture than in the forest at 1 m above the ground. VPD and PFD at ground level and soil temperature were similar in the pasture and the forest, indicating that pasture grasses strongly modify microclimatic conditions near the soil surface. The lowest gravimetric water content recorded in the pasture during the dry season was 0.5 and leaf relative water contents of the two species measured in the forest and pasture were identical, suggesting that plants in the pasture were not water stressed. Levels of most soil nutrients were lower in the pasture as compared to the forest; however, aboveground and root biomass for seedlings grown in pasture and forest soils did not differ significantly. Although a number of factors impede forest recovery in abandoned pastures, these results suggest that the most imporrant limitation is lack of seed dispersal.     

Tropical rain forest fragmentation,howler monkeys (Alouatta palliata), and dung beetles at Los Tuxtlas,Mexico

In Neotropical rain forests, fresh mammal dung, especially that of howler monkeys, constitutes an important resource used by dung beetles as food and for oviposition and further feeding by their larvae. Tropical rain forest destruction, fragmentation, and subsequent isolation causing reductions in numbers of and the disappearance of howler moneys may result in decreasing numbers of dung beetles, but this has not been documented. In this study, we present information on the presence of howlers and dung beetles in 38 isolated forest fragments and 15 agricultural habitats. Howler monkeys were censused by visual means, while dung beetles were sampled with traps baited with a mixture of howler, cow, horse, and dog dung. Results indicated that loss of area and isolation of forest fragments result in significant decrements in howlers and dung beetles. However, dung beetle abundance was found to be closely related to the presence of howler monkeys at the sites and habitats investigated. Scenarios of land management designed to reduce isolation among forest fragments may help sustain populations of howler monkeys and dung beetles, which may have positive consequences for rain forest regeneration. Am. J. Primatol. 48:253–262, 1999. © 1999 Wiley‐Liss, Inc.     

Postdispersal survivorship of north Queensland rainforest seeds and fruits: Effects of forest,habitat and species

Abstract Diaspore (seed and/or fruit) survivorship of 12 species of North Queensland rainforest trees was monitored over a 4 week period in two rainforests on the Atherton Tableland, north Queensland, Australia, with replicated sites in forest interiors, canopy gaps, forest edges and adjacent pastures. General linear modelling was used to estimate the relative importance of forest site, habitat, species and diaspore type on survival. The probability of survival varied significantly among species, between habitat, forest and diaspore types. Survival averaged 41.2% at day 28, was greater in one forest (Curtain Fig, 56.1%) than the other (Lamins Hill, 25.3%) and was apparently inversely related to the abundance of rodents. In both forests, survival declined in the order: pasture ≥ forest edge ≥ forest interior = canopy gap. A lack of significant difference between the forest interior and canopy gap is consistent with findings in other rainforest sites within Australia, but contrasts with most results from other continents. The study also indicated that diaspore weight was an important intrinsic variable affecting survival, the predators (mostly rodents) taking small to moderately large diaspores rather than the very large ones.     

Use of Fruiting Trees by Birds in Continuous Forest and Riparian Forest Remnants in Los Tuxtlas,Veracruz, Mexico1

The composition of plant species in fragmented landscapes may be influenced by the pattern of visitation by birds to fruiting trees and by the movement of seeds among and within fragments. We compared bird visitation patterns to two tree species (Dendropanax arboreus, Araliaceae; Bursera simaruba, Burseraceae) in continuous forest and remnants of riparian vegetation in a region dominated by pasture in Los Tuxtlas, Veracruz, Mexico. We quantified frequency of visitation, fruit consumption, consistency of visitation (percentage of total tree observation periods during which a given bird species was recorded), and species composition of birds at individuals of both tree species in continuous forest and riparian remnants. Bird visitation rate, species richness, and fruit consumption rates were similar within both tree species in the two habitats. Species assemblages at D. arboreus were different between continuous forest and remnants. Species assemblages at B. simaruba did not differ by habitat. Our results demonstrate that habitat disturbance may influence avian visitation patterns, which may in turn affect subsequent recruitment patterns in some tree species. Our results, however, were not consistent between the tree species, suggesting that it is difficult to generalize concerning the effects of forest disturbance on avian species assemblages in fruiting trees.     

Patterns of Seed Size,Germination and Seed Viability of Tropical Tree Species in Southern India1

Seed weight, days to germination and seed viability were observed for 99 species growing in the Western Ghats of Karnataka, India. Seed size was strongly correlated with days to germination; smaller seeds germinated faster than larger seeds. Species which flowered during the rainy season had lighter seeds than species which flowered during the dry season. It was also found that seed size and viability of seeds were related to the season of fruiting. Species which fruit during the rainy season had heavier seeds and shorter viability than species which fruit during the dry season. These flowering and fruiting patterns and varying seed sizes are argued to be adaptations to the time of dispersal, time of moisture availability in the habitat and seedling survival.