Hypocrealean (Hypocreales,Ascomycota) Fungal Diversity in Different Stages of Tropical Forest Succession in Costa Rica1

The relationship between forest succession and microfungal diversity has been poorly studied. Fungi provide important ecosystem services that may deteriorate in deforested or highly disturbed forests. To determine the possible effects of deforestation and forest succession on microfungi, species diversity of hypocrealean fungi (Ascomycota) was compared in forest stands in Eastern Costa Rica representing three stages of succession: 1–2, 25–27 yr old, and an old growth forest. Species diversity in a second‐growth forest fragment surrounded by timber plantations and second‐growth forest was also compared to that of a stand surrounded by old growth forest. The results show that the overall diversity of hypocrealean fungi was inversely proportional to the age of the forest stand, and each family showed different successional trends. Clavicipitaceae was more diverse in the old‐growth forest and was positively related to the age of the forest stand. Nectriaceae was highly diverse in the 1‐ to 2‐yr‐old stand and less diverse in the old‐growth stand. Saprobic and plant pathogenic fungal species were more diverse in the 1‐ to 2‐yr‐old stand and their diversity was inversely proportional to the age of the forest stand. The diversity of insect pathogens was positively related to the age of the forest stand. The 20‐ to 22‐yr‐old forest fragment had the lowest number of species overall. Based on the data gathered in this study, hypocrealean fungal species diversity is related to the successional stage and fragmentation of tropical forest.     

Establishment of Epiphytic Bromeliads in Successional Tropical Premontane Forests in Costa Rica

Plant community composition is the combined result of species-specific competitive abilities and the availability of propagules. For epiphytic plants, current hypotheses consider that dispersal-related factors are most important. By controlling seed dispersal constraints, we experimentally examined whether the community composition of epiphytic bromeliads in a tropical premontane area is determined during early phases of seedling recruitment. Also, we tested whether establishment success was related to eco-physiological traits of the species. A total of 7200 seeds were artificially affixed on several host trees in two secondary forest patches and in a mature forest stand. Four bromeliad species with differing physiological characteristics (CAM, C₃-CAM, and C₃) and habitat preference (secondary vs. primary forest) were selected. We found that differences in seed germination probability among habitats and species were not likely to influence community assembly. After 2 yr, seedling survival and plant development were relatively higher in the early-successional forest. Seedling establishment success was not associated with specific physiological and morphological adaptations or habitat preference of the studied species. Our results were not consistent with the described community composition and rates of population recruitment of the studied species in the same successional habitats. The results support the hypothesis that chance and historic events related to seed dispersal have an important influence on community assembly of epiphytic plants. In addition, differences in growth rates and reproductive turnover among species are expected to influence the relative abundance and recruitment rates in a particular habitat.     

Environmental control of flowering periodicity in Costa Rican and Mexican tropical dry forests

Aim We analyse the proximate causes of the large variation in flowering periodicity among four tropical dry forests (TDF) and ask whether climatic periodicity or biotic interactions are the ultimate causes of flowering periodicity. Location The four TDFs in Guanacaste (Costa Rica), Yucatan, Jalisco and Sonora (Mexico) are characterized by a 5–7 month long dry season and are located along a gradient of increasing latitude (10–30°N). Methods To dissect the differences in flowering periodicity observed at the community level, individual tree species were assigned to ‘flowering types’, i.e. groups of species with characteristic flowering periods determined by similar combinations of environmental flowering cues and vegetative phenology. Results Large variation in the fraction of species and flowering types blooming during the dry and wet season, respectively, indicates large differences in the severity of seasonal drought among the four forests. In the dry upland forests of Jalisco, flowering of leafless trees remains suppressed during severe seasonal drought and is triggered by the first rains of the wet season. In the other forests, leaf shedding, exceptional rainfall or increasing daylength cause flowering of many deciduous species at various times during the dry season, well before the summer rains. The fraction of deciduous species leafing out during the summer rains and flowering when leafless during the dry season is largest in the Sonoran TDF. Main conclusions In many wide‐ranging species the phenotypic plasticity of flowering periodicity is large. The distinct temporal separation of spring flowering on leafless shoots and subsequent summer flushing represents a unique adaptation of tree development to climates with a relatively short rainy season and a long dry season. Seasonal variation in rainfall and soil water availability apparently constitutes not only the proximate, but also the ultimate cause of flowering periodicity, which is unlikely to have evolved in response to biotic adaptive pressures.     

Growth of Mantled Howler Groups in a Regenerating Costa Rican Dry Forest

We examined population dynamics in mantled howlers (Alouatta palliata palliata) in a regenerating tropical dry forest in Santa Rosa National Park (SRNP), Costa Rica. The population has grown at a rate of about 7% per annum during the past decade. The growth in numbers from 342 in 1984 to 554 in 1992 reflects an increase in the number of groups (from 25 to 34) and a slight increase in their average size (from 13.6 to 16.3). Population density has increased from 4.9 to 7.9 individuals per km ². Santa Rosa's population density and group compositions are similar to those at several other mantled howler sites, but densities of mantled howlers are much higher at two other well-studied sites: La Pacifica and Barro Colorado Island (BCI). We relate the low density of howlers at Santa Rosa to local historical and ecological factors. Howler populations at high and low densities differ in average group size and sex ratio. At high population densities, groups are larger and include more adult females. The number of male howlers per group appears to be more strictly limited and less variable than the number of females is. However, there is greater variation in male group membership at Santa Rosa than at La Pacifica or BCI, and at Santa Rosa there are more generating forests available into which males and females can disperse and form new groups. We present case studies describing two ways in which new howler groups are formed, and we suggest that, compared to females and compared to males at high density sites, males are relatively advantaged in the uncrowded habitats at Santa Rosa and other low density sites.     

Arthropod Assemblages in Epiphyte Mats of Costa Rican Cloud Forests

Tropical cloud forests are functionally important ecosystems, but are severely threatened due to deforestation and fragmentation. Epiphyte mats, accumulations of live vegetation and dead organic matter on tree trunks and branches, are a conspicuous component of cloud forests and harbor diverse assemblages of meso- and microarthropods. We compared the morphospecies richness, composition, and abundance of arthropods in epiphyte mats between primary and secondary forests of Monteverde, Costa Rica, and at two nearby replicate sites. Epiphyte mats were thinner and less structurally diverse in secondary forest. We collected ca 36,000 micro- and mesoarthropods from epiphyte mats in the 2-yr study. Whereas arthropod morphospecies richness did not differ among forest types, arthropod abundance was significantly higher in secondary forest due to larger numbers of ants, especially Solenopsis spp. Arthropod assemblages showed a high degree of taxonomic overlap both within and between primary and secondary forests (Jaccard abundance-based similarity = 0.93–0.96). Although characteristics of the arthropod fauna proved to be similar among sites and between forest types, there was a significant temporal effect: arthropod morphospecies richness in epiphyte mats generally was lower in the dry season (February–May), when many taxa probably became dormant or sought shelter against desiccation in deeper portions of mats.     

Retention of Inorganic Nitrogen by Epiphytic Bryophytes in a Tropical Montane Forest1

We developed and evaluated a model of the canopy of a tropical montane forest at Monteverde, Costa Rica, to estimate inorganic nitrogen (N) retention by epiphytes from atmospheric deposition. We first estimated net retention of inorganic N by samples of epiphytic bryophytes, epiphyte assemblages, vascular epiphyte foliage, and host tree foliage that we exposed to cloud water and precipitation solutions. Results were then scaled up to the ecosystem level using a multilayered model of the canopy derived from measurements of forest structure and epiphyte mass. The model was driven with hourly meteorological and event‐based atmospheric deposition data, and model predictions were evaluated against measurements of throughfall collected at the site. Model predictions were similar to field measurements for both event‐based and annual hydrologic and inorganic N fluxes in throughfall. Simulation of individual events indicated that epiphytic bryophytes and epiphyte assemblages retained 33–67 percent of the inorganic N deposited in cloud water and precipitation. On an annual basis, the model predicted that epiphytic components retained 3.4 kg N ha/yr, equivalent to 50 percent of the inorganic N in atmospheric deposition (6.8 kg N ha/yr). Our results indicate that epiphytic bryophytes play a major role in N retention and cycling in this canopy by transforming highly mobile inorganic N (ca. 50% of atmospheric deposition is NO^?₃) to less mobile (exchangeable NH⁺₄) and recalcitrant forms in biomass and remaining litter and humus.     

Changes in diversity along a successional gradient in a Costa Rican upper montane Quercus forest

Changes in terrestrial vascular plant diversity along a successional gradient were studied in a Costa Rican upper montane Quercus forest. In 1991 and 1992 species presence and cover were recorded in 12 successional 0.1 ha forest plots. A total of 176 species in 122 genera and 75 families were found. Asteraceae was the most speciose family. With the help of TWINSPAN three successional phases were classified: (i) Early Secondary Forest (ESF, 145 spp.), (ii) Late Secondary Forest (LSF, 130 spp.) and (iii) Primary Forest (PF, 96 spp.). Detrended Correspondence Analysis (DCA) species ordination using DECORANA illustrates that different ecological species groups can be distinguished along the time sequence. Alpha diversity (Shannon-Wiener index, among others) in ESF and LSF was significantly greater than in PF. This is probably explained by downslope migration of numerous sub(alpine) species to cleared and recently abandoned montane sites. Beta diversity applying Sørensen's similarity coefficients declined during succession. Using linear regression, the minimum time required for floristic recovery following disturbance and abandonment was calculated at 65.9 years. A comparison with other studies shows that secondary forests in upper montane Costa Rica can be as diverse as in neotropical lowlands.     

Diversity of mesophilic clostridia in Costa Rican soils

Costa Rica is located in the Tropic, one of the most biologically diverse regions of the world; its soil is an important epicenter of biodiversity and Clostridium spp. are among the most frequent bacteria. The diversity of clostridia in Costa Rican soils and its possible association with geographic zone, pH or type of soil was studied in 117 soil samples: 18 from the Atlantic Zone, 30 from the Central Plateau, 30 from the Dry Pacific, 13 from the North Zone, and 26 from the South Pacific. The pH and the mesophilic clostridia species were determined for each sample. For bacterial isolation, a selective methodology for spores and pre-reduced anaerobically sterilized media were used. A total of 1945 strains of clostridia were isolated, 98% were identified and corresponded to 54 species; the most frequent species were C. subterminale (56%), C. oceanicum (51%), C. bifermentans and C. glycolicum (50%, each), C. sporogenes (49%), and C. sordellii (42%). An average of 7.1 species per sample was obtained; the Atlantic Zone showed the greatest diversity: 8.6 species per sample and a total of 45 species. Except for C. chauvoei, all described toxigenic clostridia species were isolated; C. sordellii (42%) and C. perfringens (38%) were the most frequent. No statistical relation could be established between geographic zone or type of soil and any species, showing that clostridia had a high adaptation capability to grow in different soil conditions; only some clostridia were isolated from very acidic samples while others from soils with a wide range of pH. In general, a uniform distribution of most species and a high variety of clostridia in Costa Rican soils were observed, in agreement with the high biodiversity described for other living beings in this country.     

Ecophysiology in Relation to Exposure of Pendant Epiphytic Bryophytes in the Canopy of a Tropical Montane Oak Forest1

We investigated ecophysiological and morphological traits affecting light and water use of four commercially important pendant epiphytic bryophyte species in a montane oak‐bamboo forest in Costa Rica. Predictions based mostly on ecophysiological studies of temperate bryophytes and vascular plants were experimentally tested on tropical montane bryophyte species ranked on the basis of their distributions in the canopy from the most protected (1) to the most exposed sites (4): (1) Phyllogonium viscosum; (2) Pilotrichella flexilis; (3) Dendropogonella rufescens; and (4) Frullania convoluta. As predicted, highly exposed species tended to have higher light saturation and compensation points, higher dark respiration rates, more chlorophyll, higher chlorophyll a:b ratios, and higher nitrogen concentrations. Contrary to predicted trends, exposed‐site species had lower water contents at full saturation; there were no detectable differences among species in the rate of water loss. Rates of carbon gain in all species reached asymptotes with increasing water content, but the moisture compensation point for carbon uptake of the moss D. rufescens was substantially higher than in the other species. Observed patterns result from interactions among processes related to water storage and transport operating at different scales and are determined by various morphological traits including the density, size, and disposition of phylloids, as well as by whole‐clump architecture.     

Differences in epiphyte biomass and community composition along landscape and within-crown spatial scales

Vascular epiphytes contribute to the structural, compositional, and functional complexity of tropical montane cloud forests because of their high biomass, diversity, and ability to intercept and retain water and nutrients from atmospheric sources. However, human-caused climate change and forest-to-pasture conversion are rapidly altering tropical montane cloud forests. Epiphyte communities may be particularly vulnerable to these changes because of their dependence on direct atmospheric inputs and host trees for survival. In Monteverde, Costa Rica, we measured vascular epiphyte biomass, community composition, and richness at two spatial scales: (1) along an elevation gradient spanning premontane forests to montane cloud forests and (2) within trees along branches from inner to outer crown positions. We also compared epiphyte biomass and distribution at these scales between two different land-cover types, comparing trees in closed canopy forest to isolated trees in pastures. An ordination of epiphyte communities at the level of trees grouped forested sites above versus below the cloud base, and separated forest versus pasture trees. Species richness increased with increasing elevation and decreased from inner to outer branch positions. Although richness did not differ between land-cover types, there were significant differences in community composition. The variability in epiphyte community organization between the two spatial scales and between land-cover types underscores the potential complexity of epiphyte responses to climate and land-cover changes.     

Reproduction of the Bat Glossophaga commissarisi (Phyllostomidae: Glossophaginae) in the Costa Rican Rain Forest During Frugivorous and Nectarivorous Periods1

I studied the reproduction of the 9 g nectarivorous bat Glossophaga commissarisi (Phyllostomidae: Glossophaginae) in relation to its food choice in a Costa Rican lowland rain forest over more than 2 yr. Reproduction was bimodal with birth peaks in April/May and October/November. The first birth peak coincided with a period of frugivory, whereas the second occurred during a period of almost exclusive nectarivory. All adult females recaptured in consecutive reproductive periods were reproductively active on both occasions, indicating that individual G. commissarisi females regularly reproduce twice per year at the study site. Throughout the annual cycle the mean testis length of the males correlated best with the proportion of females lactating, suggesting a post partum estrous pattern. Animals became reproductively active at about 1 yr of age. Fecal samples showed that females consumed significantly more insects than males.     

Effects of Treefall Gap Disturbances on Ant Assemblages in a Tropical Montane Cloud Forest

The study of gap dynamics and the effects of gaps on diversity has been at the center of tropical ecology for decades. While most studies have focused on the responses of plant species and communities to gap formation, in this study, we consider the effects of treefall gap disturbances on leaf litter ant assemblages in a Neotropical montane cloud forest. We sampled leaf litter ant assemblages and estimated a suite of abiotic parameters in 12 large (>80‐m²) treefall gaps across a chronosequence and in 12 paired adjacent intact forest sites in the Monteverde Cloud Forest Preserve in Costa Rica. No species were more common in gaps than in intact forests, and in fact, species that were common in gaps were also among the most common in forests. The Chao2 estimate of species richness, however, was higher in gap sites than in intact forest sites. In addition, ant assemblages in gap sites did not become more similar to those in adjacent intact sites as gaps aged. In contrast to other studies, our work demonstrates that ant assemblages in the Monteverde Cloud Forest Preserve are weakly affected by the formation of treefall gaps. Together, these results indicate that treefall gap dynamics probably play little role in promoting ant diversity at more regional scales, or coexistence among species at more local scales.     

Disperser- vs. Establishment-Limited Distribution of a Riparian Fig Tree (Ficus insipida) in a Costa Rican Tropical Rain Forest1

Studies were conducted at the La Selva Biological Station in Costa Rica and in a greenhouse in California to determine the factors accounting for the nonrandom distribution of the riparian fig tree Ficus insipida Willd. along streams in the La Selva Biological Reserve and adjacent deforested lands. We also evaluated the potential seed dispersers of this tree relative to the role of the fruit‐eating fish Brycon guatemalensis that previously was proposed to be an important disperser of F. insipida seeds in this system. At La Selva, we recorded the fig‐foraging activities of vertebrates at fruiting F. insipida trees, surveyed for the presence or absence of F. insipida along streams of different sizes, and determined the fate of fig seedlings transplanted in different riparian habitats. In the greenhouse, we measured seed germination and seedling survival and growth under different light and soil pH conditions mimicking natural conditions. The findings provided evidence that (1) the tree occurs along the larger streams running through forest habitat and only along smaller streams with relatively high light availability; (2) bats (Artibeus spp.) and fish are the major dispersers of F. insipida seeds; (3) the seedlings are subject to mortality not only from low light conditions but also from treefalls, frequent flooding, and bank erosion; and (4) high light levels and near neutral soil pH result in relatively better seed germination, faster growth, and higher survival rates of seedlings. Overall, our results suggest that this fig tree is dispersed mainly by bats and fish and is more establishment‐limited than disperser‐limited in its local distribution in the La Selva rain forest habitat.     

Diversity and spatial clustering of shade trees affect cacao yield and pathogen pressure in Costa Rican agroforests

The importance of the spatial organisation of individuals in explaining species coexistence within a community is widely recognised. However, few analyses of spatial structure have been performed on tropical agroforests.The main objective of this study was to highlight the links between spatial organisation of shade trees on the one hand, and shade tree species richness and cacao yield on the other, using data from 29 cacao agroforests in Costa Rica.A method of spatial statistics, Ripley's K-function, was used to analyse the spatial organisation of shade and cacao trees in the study plots. For each stand, the X and Y coordinates of ≥2.5-m-tall trees were recorded. In each plot we also assessed shade tree species richness and cacao yield (with total number of pods = number of pods damaged by frosty pod rot + number of healthy pods).Three types of stands were identified: the first was characterised by significant clustering of shade trees, the highest shade tree species richness (S = 6), and the highest number of damaged pods (139 pods ha^?1 year^?1). The second type was characterised by random spatial organisation of shade trees. The third type showed a trend towards regular organisation. Species richness of shade trees did not differ significantly between the last two types (S = 4 for both), nor did the number of damaged pods (56 pods ha^?1 year^?1 and 67 pods ha^?1 year^?1 respectively).Although the trends were not statistically significant for all the variables in our data set, the clustered spatial structure appears to favour a synergy between environmental (tree species richness), and provisioning (cacao production) services.     

Evidence for a Novel Mutualism in the Tropical Understory Shrub Piper urostachyum1

We tested the impact of a predatory species of the family Miridae (Hemiptera) on its host plant Piper urostachyum (Piperaceae) by comparing folivory in mirid‐excluded plants and controls. We also tested the effect of resident anyphaenid spiders on mirid behavior and levels of folivory. Plants without mirids suffered significantly more folivory than plants with mirids. The impact of spider removal was less clear. Together, these results support the idea that resident mirids serve in anti‐herbivore defense for P. urostachyum plants.     

Epiphytic bromeliad communities in secondary and mature forest in a tropical premontane area

We analyzed the differences in species richness, community composition, population structure and within-tree location of epiphytic bromeliads in contiguous secondary and mature forests in a premontane area in Costa Rica. Diversity in the mature forest was highest, and the communities differed in their composition as well as in the recruitment rates of the dominant species. Guzmania monostachia and Catopsis nutans dominated the secondary forests, whereas Tillandsia fasciculata and T. tricolor were more abundant in the mature forest. The secondary forest species showed high rates of seedling recruitment while the opposite was found for the mature forest species. Species presence and abundance among and within habitats did not correlate with their physiological (i.e. CAM vs. C₃ photosynthesis) or morphological attributes. The spatial distribution patterns were similar among habitats; bromeliads tended to aggregate on a few relatively large phorophytes. The species shared a similar vertical stratification within habitats, except for the two dominant species in the early and mid-successional stages, although its ecological implication is not clear. With some exceptions, conspecifics of different ages were located on similar substrate types (i.e. stems, primary, secondary, or tertiary branches) within the tree-crowns, which suggests limited within-tree dispersion. Differences in species composition and rates of seedling recruitment among secondary and mature forest may arise from ecophysiological differences among species; however, the combined effect of seed availability and dispersal differences may have a larger influence. Thus, epiphyte community assembly can only be understood when the differences in habitat conditions, the availability of propagules, their dispersal characteristics and requirements for seedling establishment are known.