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To determine if there were consistent differences in growth, mortality, and recruitment on slopes and ridge crests in tropical montane forests, which could explain the (frequent but not universal) low stature of trees in the ridgetop forests, we analyzed data from long-term plots in Jamaica (1990–1994; sixteen 200-m2 plots, six on ridge crests and five each on north and south slopes). Mortality was higher on north slopes, while growth and recruitment rates were not significantly different among positions. Soil pH and effects of recent disturbance by Hurricane Gilbert were positively correlated with growth and recruitment, while slope angle and disturbance effects were the best predictors of mortality. The patterns we found in Jamaica, that growth and recruitment were not higher on ridge crests than slopes, are different than those found by Herwitz and Young in Australia where growth and turnover were greater on a ridge crest. Therefore, it is not possible at present to make simple generalizations about dynamics of ridge crest versus slope forests in the montane tropics.
The mobile crane of the Surumoni project allowed for the first time ever a complete inventory and spatial description of the epiphytic vegetation of a tropical lowland rain forest plot (1.5 ha), at La Esmeralda on the upper Orinoco River, Venezuela. A total of 778 individual vascular epiphytes of 53 species was found, dominated by 19 orchid species and 14 species of Araceae. Fifty percent of all individual plants were obligate ant-garden epiphytes. The distribution of epiphytes was highly clumped and not random. The clumped occurrence of holoepiphytes (complete life cycle on host tree) was the consequence of the rarity of suitable phorophytes (host trees; e.g., size and age) in the plot and the preference of ants for gaps where most of the ant-garden epiphytes were found. In comparison, hemiepiphytes were distributed more evenly because of greater independence from tree suitability. The dispersal modes of epiphytes did not explain their distribution patterns. There was no consistent difference in distribution between anemochorous and zoochorous epiphytes, presumably because availability of suitable substrate is the more important factor for epiphyte establishment and growth. Whereas the vertical distribution of epiphytes could be attributed largely to deterministic factors such as physiological adaptation and requirements, horizontal distribution appeared to be governed by suitable substrate, which in turn seemed to be governed by stochastic gap formation.
A one-year phenological study of three columnar cacti, Stenocereus griseus (Haw.) Britton & Rose, Pilosocereus sp., Cereus hexagonus (L.) Mill., and a decumbent cactus Monvillea cf. smithiana (Britton & Rose) Backeberg., was carried out in the Andean arid region of La Tatacoa, Colombia. Pollinators and/or dispersers of the cacti species also were studied monthly, and fecal samples were collected for the identification of pollen and seeds. The flowering of all species was prolonged and showed bimodal, multimodal, or irregular patterns. Fruiting in all species also was prolonged and followed flowering with a lag of less than two months. Although there were no simple correlations between rainfall and flowering or fruiting, flower production during the dry season was higher for S. griseus, while Pilosocereus sp. and C. hexagonus showed higher flower production during the wet season. Fruit production was also seasonal, with higher production during the wet season for S. griseus and C. hexagonus. The patterns of flowering and fruiting in M. cf. smithiana showed no relationships with dry and wet seasons. The bats Glossophaga longirostris, Carollia perspicillata, Sturnira lilium, the birds Melanerpes rubricapillus (Picidae) and Mimus gilvus (Mimidae), and moths of the family Sphingidae, were identified as pollinators and/or fruit consumers of these cacti species.
Fates and growth of Rafflesia patma buds and insect visitation to the flowers were monitored in an evergreen tropical rain forest in the Pangandaran Nature Reserve, Java, Indonesia. Of 59 buds marked in December 1985, 44 percent died before flowering, 7 percent flowered, and 49 percent were still in bud in May 1986. Mammals and birds caused the deaths of buds before they reached maturity. Only four buds flowered between December 1985 and May 1986; all were male. Rate of increase in diameter (cm/d) was much faster in large than in small buds. Once opening began, the flower opened fully in 24 to 48 hours; the flower remained open for three to five days before rotting. Only two genera of insects visited a male flower of R. patma: Lucilia and Sarcophaga (Diptera). Insect visitation was highest in the afternoon (1300–1700 h) and on the second or third day after flower opening. The increased number of flies visiting the flower coincided with the increasing odor emitted from it. Results of the present study were combined with information from the literature to construct a life cycle diagram of R. patma.
Changes in light or water availability can result in synchronous leaf production, concentrating food availability for herbivores of young leaves to only a few months. To determine the importance of food availability on herbivory, leaf phenology and leaf damage were studied in the Luquillo Experimental Forest (LEF) of Puerto Rico. We studied 20 individuals of eight species for two years. Every month, new leaves were marked; the following month, leaf area and area of damage were measured. Over two years, comparison of leaf production and percent herbivory were performed for each species, and for all species taken together. More than 30 percent of the annual leaf production occurred in May and June. Leaf production was associated with an increase in PFD (photon flux density) and was not related to the patterns of rainfall. Although leaf production was synchronous, there were no differences in herbivory between the peak and non-peak periods of leaf production. Possible explanations for the constant levels of herbivory throughout the year are the presence of a generalist herbivore community, the ability of herbivores to track changes in food availability, or high densities of herbivore predators that control herbivore populations.
We compared the community structure, nutritive quality, and aboveground biomass of grazing lawns (patches of shortgrass communities) to neighboring grasslands in the Terai of western Nepal. Grazing lawns differed from the adjacent grasslands in species composition and community structure. Species diversity and species richness were higher on grazing lawns (H = 1.60, S = 20.93) than the grasslands (H = 0.97, S = 8.97). Fencing that excluded grazers for 150 days made areas of grazing lawns indistinguishable from neighboring grasslands in terms of plant height and biomass. Growing shoots of forage from grazing lawns had higher digestibility, crude protein, and sodium than forage from the grasslands. Grazing lawns appear to be maintained by continuous grazing and are enriched by deposition of urine, dung, and by certain plant species not found in the the adjacent grasslands.
Tropical monodominant forests in which one tree species dominates the canopy occur in all three major tropical regions, but few studies have focused on the mechanisms responsible for dominance. This study tests the hypothesis that relative to other species in the community, dominant species are well defended and escape herbivore and pathogen damage. We surveyed the rate of damage on young expanding leaves of seedlings and saplings belonging to eight species within both monodominant Gilbertiodendron dewevrei forests and adjacent mixed-species forests in eastern Congo. Results showed that escape from herbivore and pathogen damage is not a mechanism by which Gilbertiodendron achieves dominance, as it suffered the highest damage level of any species surveyed. Similarly, other sub-dominant common species also suffered high rates of damage. These results are discussed in relation to the phenolic, fiber, and nitrogen content of leaves, and in the context of current theories pertaining to plant–herbivore interactions.
Characteristic secondary metabolites are currently acknowledged to play a pivotal role in the circumscription of a plant's insect fauna. A newly discovered association between wood-boring longicorn beetles (Cerambycidae) and their host trees belonging to the Brazil nut family (Lecythidaceae) inspired a yearlong rearing project in the lowland Neotropical rain forest of central French Guiana. Branches severed from five species of Lecythidaceae yielded 1813 cerambycids belonging to 37 species. Two of the five tree species, Couratari stellata and Gustavia hexapetala, yielded impoverished complements of cerambycids that included large percentages of generalists. Both tree species are characterized by fetid odors. Wood samples collected from the malodorous C. stellata, as well as from two well-colonized species, Lecythis poiteaui and Eschweilera coriacea, were analyzed for their volatile components. Sulfur compounds accounted for almost 15 percent of the volatiles detected from C. stellata, while they were at most minor components of the two well-colonized tree species. S-methylmethionine, isolated from a C. stellata wood sample, appeared to be the major sulfur compound contributing to the distinctive smell. We hypothesize that this foul odor is a deterrent to specialist cerambycids seeking oviposition sites.
The effects of dispersal pattern (seeds in small clumps vs. seeds scattered in pairs) and distance to the nearest Carapa procera (Meliaceae; a tree that produces seeds preferred by terrestrial vertebrates) on survival of seeds and seedlings were examined for the animal-dispersed tree species Virola michelii (Myristicaceae) in a mature forest at Paracou, French Guiana, in 1992 and 1993. We assessed the putative role of ground-dwelling mammalian herbivores, rodents, and ungulates that filter the seed shadow, acting either as dispersers or predators and thus modifying the original pattern of seed dispersal made by frugivores. We measured the effects of simulated seed burial by rodents using marked seeds and quantified the effect of protecting seeds and seedlings from ground-dwelling vertebrates on seedling germination and survival with fence exclosures in 1992. Dispersal pattern had short-term but no long-term effects on the proportion of V. michelii seeds that survived one year later as seedlings. In the short term, within six weeks, clumped seeds survived better than scattered seeds in both years. Marked seeds that were removed from their site of dispersal were eaten; rodents only rarely buried seeds of V. michelii, and seed burial reduced seed and seedling survivorship. The combined effect of the factors year and Carapa proximity significantly affected seed survival within six weeks. Although six-week seed survival was greater in 1993 than in 1992, seedling establishment was lower in 1993 than in 1992 following a lower rainfall regime during the key period of seed germination (February). One-year seed and seedling survivorship was similar between treatments and years. Seed survival and seedling establishment in V. michelii was dependent on vertebrates in the short term and on climate in the long term. Overall, seed and seedling survivorship depended on a combination of these factors.
The genus Dombeya is an important component of tropical forests in west Africa, Madagascar, and the Mascarene islands, but little is known about the reproductive biology of its members. In this study, we quantified gender variation and flower-size variation in three populations of the dioecious tree Dombeya ciliata, an endemic of La Réunion Island (Indian Ocean). Variation among the three populations was observed for three characters. First, the high elevation population had strict males and females, whereas in the two low elevation populations, males were inconstant, with > 50 percent of the males set a small number of fruit. Second, females showed greater seed production at high elevation. Third, flower-size dimorphism was greater at high elevation because male flowers were larger than female flowers.
Few data exist on seed dispersal by frugivorous birds in fragmented landscapes, originating from tropical dry forests, in contrast to more abundant data from tropical rain forests. In this study, we assessed the effect of frugivorous birds in a fragmented landscape of Veracruz, Mexico, now occupied by remnant fragments of tropical semi-deciduous forest and dry deciduous forest, grassland, and shrubby patches on sand dunes. We determined four characteristics related to seed dispersal by birds: the interacting species of plants and birds, the characteristics of these species, spatio-temporal variation in the dispersal system, and the outcome of the process. During one year, we recorded 54 frugivorous bird species and 33 ornithochorous plant species, which engaged in 176 different bird–plant species interactions. Similarity (Sørensen index) of frugivorous bird communities using different vegetation types was high (>70%), suggesting that many bird species used all of the vegetation types. In contrast, the similarity of ornithochorous plant communities among vegetation types commonly was low (<37%), suggesting that most plant species were restricted to particular sites in this landscape. At the landscape level, as well as for tropical deciduous forest, we detected a significant positive relationship (Spearman's correlation of rank coefficient >0.65, P < 0.05) among richness per month of frugivorous birds and plant species bearing fleshy fruits. Seeds of many plant species previously detected in studies of seed rain at the site were eaten by birds during this study. Most seeds of zoochorous species, which are deposited in the dry and decidous tropical forests patches, are produced within these vegetation types (i.e., they are autochthonous species), whereas bird-dispersed seeds arriving in grassland and shrubby patches are produced outside (i.e., allochthonous) and are mostly woody species. Birds are important seed dispersers among vegetation types in this landscape but they have different effects in each one. The four characteristics studied, as well as the landscape approach of this research, allowed us to detect spatial and temporal patterns that otherwise would have remained undetected.
Figs are a critical resource for many tropical frugivores, yet they often are referred to as low quality fruits. To determine their nutritional value, both as a group and for individual species, we analyzed 14 fig species from Barro Colorado Island (BCI), Panama, for fiber, tannins, lipids, protein, carbohydrates, amino acids, and minerals. Seeds and pulp were analyzed separately. Fig fruit pulp consisted of about one-third digestible components, mostly carbohydrates with some lipids and proteins. Tannin, lignin, and water-soluble carbohydrates showed considerable variation among species, as did fruit size. Figs contained high amounts of amino acids, such as leucine, lysine, valine, and arginine, and minerals, such as potassium, calcium, magnesium, sodium, and phosphorus. One species, Ficus insipida, contained the highest concentrations of almost all amino acids, many minerals, and protein. Small figs had as much nutritional value per gram as large figs. Free-standing figs had higher percentages of protein, complex carbohydrates, and ash than strangler figs, which had higher percentages of water-soluble carbohydrates, tannins, and hemicellulose. The guild of fruit-eating bats on BCI included ten common species with diets dominated by figs. Fecal analyses and captures at ripe fig trees showed a consistent pattern of resource partitioning. Small bats preferentially ate small-fruited and strangler figs while large bats consumed mostly large-fruited and free-standing figs. Small bats most often ate F. bullenei, which has high levels of lipid and carbohydrates, and F. yoponensis, which has high levels of protein. Medium and large bats most often ate F. insipida, a nutritionally superior species; their second most eaten species was F. obtusifolia, in which the large size may make it efficient to eat. Each bat ate a variety of fig species, supporting the idea that although no single species of fig may be sufficient to sustain frugivores, a mix of fig species can provide a complete set of nutrients.
Beetles were collected on 64 trees of four species (Cynometra alexandri C. H. Wright, Rinorea beniensis (Welwitsch ex Olivier) Kuntze, Teclea nobilis Delile, and Trichilia rubescens Olivier) in Budongo Forest, Uganda, using an insecticidal fogging technique. Selected tree species were abundant, taxonomically not closely related, and different in the shape of leaves, growth form, and size, with heights between 7 and 35 m. Trees were fogged in an old primary forest stand, in an area of secondary forest where selective logging was performed, and in a swamp forest. Eight conspecific trees per forest type were fogged. A total of 29,736 beetles were collected from all trees that could be assigned to 1433 (morpho)-species; 41.6 percent were singletons and 89.6 percent of species were found with less than ten individuals. Abundant beetle taxa included Latridiidae (N = 4093), Chrysomelidae (3952), Staphylinidae (2931), Apioninae (2621), and Curculionidae (2457). Most species-rich groups were Staphylinidae (N = 196 spp.), Curculionidae (189), and Chrysomelidae (148). Abundance increased in the order: primary < secondary < swamp forest. Due to the relatively high dominance of some species in the secondary forest, species richness increased in the order: secondary < primary < swamp forest. Beta diversity measures and factor analysis showed distinct differences among forest types but higher similarity of beetle communities on different tree species within one forest type. The taxonomic distribution of beetles in the secondary forest was more heterogeneous than in the primary forest. Analyses of the data revealed low host specificity even for phytophagous beetles, underlining the importance of habitat structure and chance effects on the spatial distribution of beetles in the canopy of Budongo Forest.
Tropical rain forests are characterized by their rich plant diversity and highly diverse insect faunas containing mainly rare species. Phytophagous and parasitoid insects utilizing such fragmented resources often must travel considerable distances to find suitable hosts. For small, weak-flying insects, entry into the fast-flowing air above the canopy can provide one way by which long-distance dispersal is achieved. Using sticky traps placed at different heights in a lowland rain forest of Borneo, we compared the diurnal and nocturnal flight heights of chalcids, a group of mainly very small parasitoids and phytophages, to determine if the air above the canopy was used for dispersal. Most families were represented throughout the range of trap heights, including those above the general canopy. A higher proportion of individuals were trapped above the canopy at night than during the day. Fig wasps were exceptional in that they were trapped almost entirely above the canopy. They included species associated with host trees that do not fruit in the canopy, suggesting that these short-lived, slow-flying insects actively fly up above the canopy and then use the wind to passively carry them the long distances needed to reach their highly localized and ephemeral hosts. Once the fig wasps detect the species-specific volatiles released by their host figs, they then may fly down into the canopy, where the lower wind speeds would allow them to fly actively upwind to their hosts.
The Resplendent Quetzal (Pharomachrus mocinno) is an altitudinal migrant that nests in high elevation cloud forests and migrates toward lower areas during the summer rainy season. It has been suggested that its migratory movements are related to the abundance of ripe Lauraceae fruits. We studied the quetzal diet during two consecutive years, as well as changes in fruit abundance of the plant species on which the bird feeds at El Triunfo Biosphere Reserve, southeastern Mexico. The quetzal was observed feeding on 32 plant species; of these, 24 are new records in its diet. We chose 20 of these 32 species and studied their fruit phenology for two years in order to describe the relationship between fruit and quetzal abundance. Our results showed that quetzal abundance in the breeding area was correlated with the total number of fruiting species, whereas the correlation between quetzal abundance and the number of fruiting Lauraceae species was only marginal. Additionally, a correlation test showed that quetzal abundance was marginally correlated with total fruit availability (total no. of fruits per month); however, the correlation between quetzal abundance and the number of fruits in the Lauraceae was not significant. Our results suggest that the dynamics of food resources may be playing a major role in the quetzal's migratory behavior. Knowing the bird's diet may aid in characterizing the type of habitat adequate for its conservation. Our observations in this respect suggest that conservation efforts to preserve this bird species should concentrate on the protection of its habitat, including both breeding and nonbreeding (migration) locations.
Data on fruit abundance and ecological overlap among Ateles belzebuth, Lagothrix lagothricha, Cebus apella, and Alouatta seniculus were gathered during 13 months at Tinigua National Park (Colombia), in an effort to test the following hypotheses concerning competition for fruits. Coexistence is possible because: (1) during periods when fruit availability is limited, the species utilize different resources; and (2) the species have different fruit preferences independent of fruit production in the forest. Differences were found in resource use (diet and habitat) for all four species. Despite these differences, all four devoted large proportions of time feeding on fruit. Fruit abundance influenced their activity patterns. Ninety percent of all interspecific aggressive interactions (N = 69) were seen in fruiting trees. The first hypothesis was best supported, given that all species significantly increased their intake of the vegetative parts of plants during periods of fruit scarcity. Fruit partitioning during periods of scarcity was observed clearly only for one pair of species (C. apella and L. lagothricha). In general, the second hypothesis was not supported as a mechanism for reducing competition because most fruit species were consumed by more than one primate species. Fruit preferences, however, may be particularly important in explaining differences in niche overlap between the most ecologically similar species: A. belzebuth relied heavily on the fruits of one palm species (Oenocarpus bataua) during periods of fruit scarcity and abundance, while L. lagothricha completely ignored this fruit.
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