Simulated seed predation reveals a variety of germination responses of neotropical rain forest species

Seed predation, an omnipresent phenomenon in tropical rain forests, is an important determinant of plant recruitment and forest regeneration. Although seed predation destroys large amounts of the seed crop of numerous tropical species, in many cases individual seed damage is only partial. The extent to which partial seed predation affects the recruitment of new individuals in the population depends on the type and magnitude of alteration of the germination behavior of the damaged seeds. We analyzed the germination dynamics of 11 tropical woody species subject to increasing levels of simulated seed predation (0-10% seed mass removal). Germination response to seed damage varied considerably among species but could be grouped into four distinct types: (1) complete inability to germinate under damage ≥1%, (2) no effect on germination dynamics, (3) reduced germination with increasing damage, and (4) reduced final germination but faster germination with increasing damage. We conclude that partial seed predation is often nonlethal and argue that different responses to predation may represent different proximal mechanisms for coping with partial damage, with potential to shape, in the long run, morphological and physiological adaptations in tropical, large-seeded species.     

Pollen dispersal and representation in a neotropical rain forest

Results of 3 years of pollen trapping on Barro Colorado Island, Panama, are presented. Minimum pollen dispersal distances are estimated for the most abundant pollen taxa. Dispersal distances for some taxa appear to be as low as 5 m, while for other taxa at least 50% of the trapped pollen had travelled more than 40 m. No consistent pattern of spatial variation was reflected in the pollen rain, however, samples close to large canopy gaps had exceptional abundances of 'disturbance' taxa. A preliminary analysis of the representation of canopy components indicates that as much as 19% of pollen caught in the traps was derived from large tree species.     

Nitrogen and phosphorus resorption in a neotropical rain forest of a nutrient-rich soil

In tropical forests with nutrient-rich soil tree's nutrient resorption from senesced leaves has not always been observed to be low. Perhaps this lack of consistence is partly owing to the nutrient resorption methods used. The aim of the study was to analyse N and P resorption proficiency from tropical rain forest trees in a nutrient-rich soil. It was hypothesised that trees would exhibit low nutrient resorption in a nutrient-rich soil. The soil concentrations of total N and extractable P, among other physical and chemical characteristics, were analysed in 30 samples in the soil surface (10 cm) of three undisturbed forest plots at 'Estaci6n de Biologia Los Tuxtlas' on the east coast of Mexico (18 degrees 34' - 18 degrees 36' N, 95 degrees 04' - 95 degrees 09' W). N and P resorption proficiency were determined from senescing leaves in 11 dominant tree species. Nitrogen was analysed by microkjeldahl digestion with sulphuric acid and distilled with boric acid, and phosphorus was analysed by digestion with nitric acid and perchloric acid. Soil was rich in total N (0.50%, n = 30) and extractable P (4.11 microg g(-1) n = 30). As expected, trees showed incomplete N (1.13%, n = 11) and P (0.11%, n = 1) resorption. With a more accurate method of nutrient resorption assessment, it is possible to prove that a forest community with a nutrient-rich soil can have low levels of N and P resorption.     

Resource partitioning and interspecific interactions among sympatric rain forest arboreal mammals of the Western Ghats, India

Resource partitioning in a community of diurnal arboreal mammalsconsisting of the lion-tailed macaque Macaca silenus, bonnetmacaque (BM) Macaca radiata, Nilgiri langur Semnopithecus johnii,and the Indian giant squirrel Ratufa indica of the Western Ghats,southern India, was studied. Differences in their diet, verticalstratification, food resource niche breadth, niche overlap,and behavioral interactions were examined. Resource partitioningwas through differential habitat use, resource use, and verticalstratification. Of the four species, the BM was not a residentspecies and made frequent forays into evergreen forest fromthe adjacent deciduous forest during the flowering season ofCullenia exarillata and fruiting season of Ficus microcarpa.The macaques had narrower niches, and the langur and the squirrelhad wider niches. Niche overlap was highest between the twomacaques. Overlap among the study species was particularly pronouncedduring the flowering of C. exarillata. There was significantcorrelation between niche overlap and intolerance among thestudy species. Certain species pairs showed little or no mutualintolerance despite high overlap. Cooperative interactions suchas alarm calls occurred more frequently among the resident species.Interaction matrices revealed an underlying pattern of interspecificdominance hierarchy, with the BM dominating over the other threespecies. Our study suggests that the BM do not coexist withthe other three because of high overlap with its congener andlow occurrence of cooperative interactions.     

Seed predation by mammals and forest dominance by Quercus oleoides,a tropical lowland oak

Summary Quercus oleoides Cham. and Schlecht is an unusual tree in several respects: it is an oak found in neotropical lowland forests, its distribution is not continuous but ratherdivided into many patches of various sizes, and it is a dominant in all the forests in which it occurs, attaining densities far higher than most species of tropical trees. This density pattern is related to the vulnerability of Q. oleoides acorns to predation by mammals. Observations of agoutis, deer, peccaries, squirrels, pocket mice and other seed consumers in Santa Rosa National Park, Costa Rica, showed that these mammals act only as predators, not dispersers, of Q. oleoides acorns. Experiments which involved placing acorns in deciduous forest where Q. oleoides does not occur, demonstrated that, due to high predation rates, the number of acorns produced by an isolated tree is far too low for adults to replace themselves.In oak forest, on the other hand, where the combined acorn crops of many oaks satiate the seed predators, acorn survivorship until germination is high enough to maintain the population. Furthermore, acorn survivorship in oak forest areas is inversely proportional to the apparent mammal density in those areas. Thus the pattern of forest dominance and patchy distribution is related to positively density-dependent acorn survivorship: where Q. oleoides is the forest dominant, it will survive, but if its density falls to the level typical of tropical trees, it will go locally extinct.     

Environmental and neighbourhood effects on tree fern distributions in a neotropical lowland rain forest

Questions: To what extent are the distributions of tropical rain forest tree ferns (Cyatheaceae) related to environmental variation, and is habitat specialization likely to play a role in their local coexistence? Location: Lowland rain forest at La Selva Biological Station, Costa Rica. Methods: Generalized linear (GLM) and generalized additive (GAM) logistic regression were used to model the incidence of four tree fern species in relation to environmental and neighbourhood variables in 1154 inventory plots regularly distributed across 6 km² of old‐growth forest. Small and large size classes of the two most abundant species were modelled separately to see whether habitat associations change with ontogeny. Results: GLM and GAM model results were similar. All species had significant distributional biases with respect to micro‐habitat. Environmental variables describing soil variation were included in the models most often, followed by topographic and forest structural variables. The distributions of small individuals were more strongly related to environmental variation than those of larger individuals. Significant neighbourhood effects (spatial autocorrelation in intraspecific distributions and non‐random overlaps in the distributions of certain species pairs) were also identified. Overlaps between congeners did not differ from random, but there was a highly significant overlap in the distributions of the two most common species. Conclusions: Our results support the view that habitat specialization is an important determinant of where on the rain forest landscape tree ferns grow, especially for juvenile plants. However, other factors, such as dispersal limitation, may also contribute to their local coexistence.     

Landscape variation of liana communities in a Neotropical rain forest

We studied local and landscape variation of liana communities across habitats differing in soil and topography in the Lacandon tropical rain forest, southeast Mexico. All liana stems 1 cm diameter breast height (DBH) were sampled within each one of eight 0.5 ha plots. Two plots were sampled in each of the following habitats: alluvial-terrace, flood plain, low-hill, and karst-range. In the whole sampled area, we recorded 2092 liana stems ha^–1 representing a total basal area of 1.95 m² ha^–1 and 90 species within 34 families. Lianas showed a strong clumped spatial pattern and a high taxonomic diversity at the scale of 50 m². On average (± s.e.), we found 10.4±0.6 stems, 4.4±0.2 species and 3.4±0.2 families per 50-m² quadrat. Bignonaceae (19 species), Malpighiaceae (9), and Fabaceae (8) comprised about 40% of total number of recorded species, and almost 50% of the total liana biomass, as expressed by an importance value index that combines species relative abundance, spatial frequency and basal area. Nineteen families (56%) were represented by just one species and Cydista (Bignoniaceae) and Machaerium (Fabaceae) were the most diverse genera with four species each. In the landscape, lianas showed a geometric diversity-dominance relationship with only three species (Combretum argenteum, Hiraea fagifolia and Machaerium floribundum) accounting for more than 50% of total biomass. More than 30% of the species were rare (<15 stems ha^–1) and showed low spatial frequency (recorded in just one of the eight plots). Liana communities differed in structure and composition among sites and habitats. Among sites, lianas exhibited four-fold variation both in stem density and basal area and two-fold variation in species richness. Liana density was significantly and positively correlated with treefall disturbance. Ordination analysis indicated a strong habitat differentiation of lianas at the family and species levels. Most species with non-random distribution among habitats (69% from 25 species) were significantly most abundant in low-hill or flood plain sites, and some (12%) were preferentially found at the karst-range sites. The karst-range habitat was well differentiated from the others in species composition and structure, and shared only 50% of common species with other habitats.Soil water availability, treefall dynamics, as well as tree host identity and abundance may play an important role in the organisation of the liana communities at the Lacandon forest.     

Flocking behaviour of foraging birds in a neotropical rain forest and the antipredator defence hypothesis

Randomly encountered foraging birds were recorded in a primary rain forest of French Guiana (13,550 records of 216 species), together with their size, diet and habitat use, to assess the relative frequencies of different types of flocking behaviour and some of their ecological correlates. Overall, 42% of birds foraged singly, primarily carnivores (raptors), nectarivores (hummingbirds) and lek-mating frugivores (manakins, some cotingas). For-aging in pairs (26.6%) was widespread, notably among insectivores in the 17–32-g size class. The remaining 31.4% of records were birds in groups of different composition and function, including, in order of decreasing frequency, (1) multispecies upper canopy flocks (83 member species identified)—the largest and most mobile associations of small insectivores, nectarivores and frugivores, mostly tanagers; (2) understorey mixed species flocks of small insectivores, at midlevels of closed forest interior, with 12 core, obligate members and 74 occasional species, mostly active foliage or bark gleaners and probers sharing a unique set of ecological characteristics; (3) monospecific groups (29 species), either gregarious foragers but solitary breeders (large frugivores in canopy) or also breeding colonially or several permanently group living cooperative breeders; (4) opportunistic gatherings of frugivores at fruiting trees (at least 40 species); (5) army ant followers near ground of closed understorey (29 species of mid- to large-sized insectivores); (6) followers of Red-throated Caracaras Daptrius americanus (23 species, usually canopy frugivores entering understorey with caracaras); (7) two raptors following monkeys. Attributes of vulnerability to predators defined by habitat structure (vegetation density or openness) and foraging behaviour (conspicuousness, speed, degree of vigilance) were important determinants of flocking propensity, at least in flocks that were not attracted by a particular food source. The results suggest that the permanent mixed-species flocks in the mature forest under-storey may be an antipredator defence to compensate for the conspicuousness and reduced vigilance resulting from active foraging behaviour in semi-open vegetation, where early detection of predators is difficult.     

Leaf and soil nitrogen and phosphorus availability in a neotropical rain forest of nutrient-rich soil

The nitrogen and phosphorus supply in a lowland rain forest with a nutrient-rich soil was investigated by means of the leaf N/P quotient. It was hypothesised a high N and P supply to the forest ecosystem with a N and P rich soil. Total N and extractable P were determined in the surface (10 cm) soil of three plots of the forest. Total N was analysed by the Kjeldahl method, and P was extracted with HCI and NH4F. The leaf N/P quotient was evaluated from the senesced leaves of 11 dominant tree species from the mature forest. Samples of 5 g of freshly fallen leaves were collected from three trees of each species. Nitrogen was analysed by microkjeldahl digestion with sulphuric acid and distilled with boric acid, and phosphorus was analysed by digestion with nitric acid and perchloric acid, and determined by photometry. Concentrations of total N (0.50%, n = 30) and extractable P (4.11 microg g(-1), n = 30) in the soil were high. As expected, P supply was sufficient, but contrary to expected, N supply was low (N/P = 11.8, n = 11).     

Spatial and temporal variation in soil CO2 efflux in an old-growth neotropical rain forest, La Selva, Costa Rica

Our objectives were to quantify and compare soil CO₂ efflux of two dominant soil types in an old-growth neotropical rain forest in the Atlantic zone of Costa Rica, and to evaluate the control of environmental factors on CO₂ release. We measured soil CO₂ efflux from eight permanent soil chambers on six Oxisol sites. Three sites were developed on old river terraces (old alluvium) and the other three were developed on old lava flows (residual). At the same time we measured soil CO₂ concentrations, soil water content and soil temperature at various depths in 6 soil shafts (3 m deep). Between old alluvium sites, the two-year average CO₂ flux rates ranged from 117.3 to 128.9 mg C m^–2 h^–1. Significantly higher soil CO₂ flux occurred on the residual sites (141.1 to 184.2 mg C m^–2 h^–1). Spatial differences in CO₂ efflux were related to fine root biomass, soil carbon and phosphorus concentration but also to soil water content. Spatial variability in CO₂ storage was high and the amount of CO₂ stored in the upper and lower soil profile was different between old alluvial and residual sites. The major factor identified for explaining temporal variations in soil CO₂ efflux was soil water content. During periods of high soil water content CO₂ emission decreased, probably due to lower diffusion and CO₂ production rates. During the 2-year study period inter-annual variation in soil CO₂ efflux was not detected.     

The value of radio tracking in the study of neotropical rain forest monkeys

Although radio tracking has been used increasingly in primate field studies over the past 15 years, some primatologists have been reluctant to use it. We present data that demonstrate benefits of radio tracking in the study of rain forest primates. Data were collected during an ecological study of Ateles geoffroyi and Cebus capucinus in hilly, dense rain forest habitat with poor visibility, in northeastern Costa Rica. We SHOW that radio tracking decreased search time for both species, which led to increased contact time and facilitated continuous data collection. Mean search time for both primate species was significantly reduced using radio tracking (Kruskal-Wallis Test, P < 0.05). Search times for both species increased at the end of the study, when the transmitters ceased functioning. These increased search times occurred despite high levels of familiarity with the ranging patterns of the animals. The rate of marking feeding trees increased significantly with radio tracking and decreased significantly when the radios failed (Kruskal-Wallis Test, P < 0.01). Other benefits of radio tracking include: finding animals far off trail, in dense vegetation, and in inclement weather; maintaining continuous contact with the animals, which allows for more complete knowledge of ranging and foraging patterns; and monitoring group and subgroup composition. © 1994 Wiley-Liss, Inc.     

Leaf trait integration mediates species richness variation in a species-rich neotropical forest domain

Environmental gradients are known to drive changes in mean trait values, but changes in the trait integration strength across local communities are less well understood, particularly with regard to possible links with species richness variation. Here, we tested if climate, soil, and topography gradients drive species richness indirectly via constraints on trait integration in the Atlantic Forest of South America. We evaluated seven traits (from leaf, wood, seed, and plant size) of 1456 species occurring across 84 local communities. Generalized least square models and a path model were applied to test direct and indirect relationships. Correlations were higher between leaf traits (average r?=?0.28) and lower when other traits were included (average r?=?0.16). In line with this result, species richness was related to a multivariate index of interspecific trait integration (ITI) computed for leaf traits, but not to the ITI for all the seven traits. Abiotic gradients influenced species richness both directly and indirectly through the leaf trait integration. A total of 33% and 26% of the variation in species richness and ITI, respectively, were explained by the models, with climatic conditions showing higher contribution than topographic and edaphic factors. These results support a significant but reduced environmental selection role behind the trait-based community assembly and may suggest that other processes are involved in the constrain of trait integration at larger spatial scales. In addition, different directional trends in trait–trait relationships across local communities suggest that global trait relationships may not necessarily hold at local contexts.

     

Members of Sebacinales subgroup B form mycorrhizae with epiphytic orchids in a neotropical mountain rain forest

Previous investigations revealed that epiphytic orchids in a mountain rain forest in southern Ecuador formed mycorrhizae with diverse members of Tulasnellales. Using specific primers, we now show that the same orchids are also associated with Sebacinales. Ultrastructural observations confirmed the Sebacinales mycobionts in situ. Mycorrhizae of flowering individuals of Stelis hallii, S. superbiens, S. concinna and Pleurothallis lilijae were sampled in different forest types of the mountain rain forest of southern Ecuador along an altitudinal gradient between 1,850 and 2,100 m a.s.l. Phylogenetic analysis of fungal nuclear rDNA sequences coding for the ribosomal large subunit (nucLSU) showed the presence of eight sequence types based on proportional differences of <1% bp. All sequence types clustered in the Sebacinales subgroup B which also contained sequences of mycobionts from ericads and terrestrial orchids. Sequences of the nuclear rDNA 5.8S subunit, including parts of the internal transcribed spacers ITS1 and ITS2 (5.8-ITS) from the mycobionts of the epiphytic orchids, were distinct from published sequences of sebacinoid mycobionts of green terrestrial orchids and ericads. Sebacinales sequences from different epiphytic orchid species differed at least by 1% bp as was previously found for Tulasnella sequences. Sebacinales occurred less frequently and with a lower number of sequence types than Tulasnellales, but distribution along the altitudinal gradient was similar.     

Seed rain across fire-created edges in a neotropical rainforest

Human-induced wildfires are increasing in frequency in tropical forests, and their deleterious consequences for biodiversity include decreases in seed rain, which may be affected directly by fire or indirectly by the creation of edges between forest and non-forest environments. Understanding seed rain is key to assess the potential for natural regeneration in plant communities. We assessed the impact of fire and fire-created edges on seed rain species richness, abundance, size, weight, and dispersal syndromes in Atlantic Forest remnants in Bahia, Brazil. We assessed seed rain at monthly intervals for an entire year along seven 300 m-long transects placed perpendicular to the edge. We installed seed traps at the edge and at 20, 40, 60, 80, 100, and 150 m into the burnt area and into the forest from forest edge. We recorded a total of 9050 seeds belonging to 250 morphospecies. We did not observe edge influence; however, we detected a lower abundance and proportion of animal-dispersed seeds in the burnt than in the unburnt areas. The seed abundance in the burnt areas was lower and seeds were smaller and lighter than those in the unburnt area. Seed rain in the burnt area was not greater near to the forest than far from it. The abundance and richness of seed rain was positively correlated with tree density. Our findings highlight the lack of seed rain in burnt areas and differences in community composition between the burnt and unburnt areas. Collectively, these results indicate negative consequences on natural regeneration, which can lead to permanent secondarization of the vegetation and challenges for early regeneration of burnt areas, which will initially have impoverished forests due to low seed richness.