Responses to Fire in Selected Tropical Dry Forest Trees1

Fire is a frequent disturbance in the tropical dry forests of Central America, yet very little is known about how native species respond to such events. We conducted an experimental burn in a tropical dry forest of western Nicaragua to evaluate plant responses to fire with respect to survivorship and recruitment. Measurements of woody vegetation of all size classes were carried out prior to the prescribed burn and three successive years post fire. We selected the 15 most abundant species <10 cm DBH to assess percent survivorship and sprouting responses post fire. Changes in seedling densities for these 15 most abundant species and the 15 least abundant species were analyzed using a repeated measure ANOVA. We also assessed changes in seedling densities for three species of international conservation concern. We found three major fire‐coping strategies among common dry forests plants: resisters (low fire‐induced mortality), resprouters (vigorous sprouting), and recruiters (increased seeding post‐fire). While survivorship was generally high relative to tropical moist forest species, those species with lower survivorship used either seeding or sprouting as an alternative strategy for persisting in the forest community. Seed dispersal mechanisms, particularly wind dispersal, appear to be an important factor in recruitment success post‐fire. Burn treatment led to a significant increase in the density of seedlings for two species of conservation concern: Guaiacum sanctum and Swietenia humilis. Results of this study suggest that common dry forest species in western Nicaragua are fire tolerant. Further study of individual species and their fire responses is merited.     

The Role of Fruits and Insects in the Nutrition of Frugivorous Bats: Evaluating the Use of Stable Isotope Models1

We estimated the relative contribution of fruits and insects as sources of dietary protein in two species of Neotropical frugivorous bats (Artibeus jamaicensis and Sturnira lilium) using stable carbon and nitrogen isotope analyses. An insectivorous species (Pteronotus parnellii) was also included for comparison. We found constant patterns in stable carbon and nitrogen isotope composition in blood that separated the two species of frugivorous bats from the insectivorous bat. When we used these isotopic values (combined with those of dietary fruits and insects) to estimate the percent contribution of fruits and insects to the diet of the bats, we obtained different results, depending on assumptions and model adopted. We tested models using both 8“N and 8′³C results simultaneously and separately and further used diet‐tissue fractionation factors of 3%o for nitrogen and 1 and 3.5%o for carbon. We found that a carbon‐based model with a diet‐blood enrichment factor of 3.5%o produced the most parsimonious results. The model estimated that A. jamaicensis and S. lilium obtained most of their protein requirements from fruits, whereas P. parnellii fed mostly on insects. No sexual or seasonal variations in the diet of the two frugivorous species were detected. We found no evidence that the diet of sexually active females differed from that of nonsexually active females in the two species of frugivorous bats. We suggest that future studies better define isotopic fractionation between diet and tissues of bats using captive rearing and controlled diets.     

Patterns of Seed Dispersal and Dispersal Failure in a Hawaiian Dry Forest Having Only Introduced Birds

Dry forests are among the most endangered natural communities in the Hawaiian Islands. Most have been reduced to isolated trees and small forest fragments in which native tree species reproduce poorly. The replacement of native birds by introduced generalists may be contributing to dry forest decline through modification of seed dispersal patterns. To document seed dispersal by introduced birds, we conducted foraging observations on fleshy-fruited trees and measured seed rain under trees and in adjacent open areas for 1 year in a dry forest dominated by native trees. Although trees covered only 15.2 percent of the study area, 96.9 percent of the bird-dispersed seeds were deposited beneath them. The Japanese white-eye (Zosterops japonicus) was the principal dispersal agent. Among bird-dispersed seeds, those of the invasive tree Bocconia frutescens accounted for 75 percent of all seeds collected beneath trees (14.8 seeds/m²/yr) and the invasive shrub Lantana camara accounted for 17 percent. Although nearly 60 percent of the reserve's native woody species possess fleshy fruits, introduced birds rarely disperse their seeds. Native trees accounted for <8 percent of all bird-dispersed seeds and are consequently experiencing dispersal failure by falling directly under parent trees. Smaller-seeded non-native plants, in contrast, may be benefiting from dispersal by introduced birds. Current dispersal patterns suggest that these readily disseminated non-native plants may eventually replace the remaining native flora.     

Seasonal Patterns of Flowering and Fruiting in a Dry Tropical Forest in Jamaica1

Tropical dry forests occupy more area and are more endangered than rainforests, yet their regeneration ecology has received less study and is consequently poorly understood. We recorded the flowering and fruiting phenology of a tropical dry forest in Jamaica over a period of 26 mo within ten 15 × 15‐m plots. Community‐wide recruitment reached a maximum in the wet season, whereas no recruitment occurred during the dry season. We observed a unimodal peak in rainfall and fruit production, and the periodicity and intensity of seed production were significantly correlated with rainfall seasonality (the optimal time for germination). Flowering at the community and system levels lagged behind a significant increase and subsequent decrease in rainfall by 7 and 3 mo, respectively, indicating that the dominant factor controlling flowering periodicity is the passage of the major (4‐mo long) rainy season and changes in soil moisture conditions. Fruiting lagged behind flowering by 2 mo and a significant increase in fruiting occurred 2 mo prior to a significant increase in rainfall. At the population level, a correspondence analysis identified a major dichotomy in the patterns of flowering and fruiting between species and indicated two broad species groups based on their time of peak fruiting and the number of times they were in fruit. These were either individuals which were usually in peak fruit 1–2 mo prior to the start of the major rainy season or those that were in fruit more or less continuously throughout the year with no peak fruiting time. This study supports the view that seasonal variation in rainfall and hence soil water availability constitutes both the proximate and the ultimate cause of flowering periodicity in tropical dry forests.     

Distribution Patterns of Tropical Dry Forest Trees Along a Mesoscale Water Availability Gradient

Tropical dry forests (TDFs) host a large diversity of tree species but little is known of potential mechanisms that contribute to its maintenance. Given the paramount importance of water availability in such forests, tree species would be expected to show nonrandom patterns along water availability gradients, as well as differential individual species responses. In this work we explored whether that was true for 50 dominant tree species. Within a total area of 5.2 ha, divided into 26 transects each with ten 20 × 10 m plots, we registered presence–absence of these tree species with diameter at breast height ≥5 cm. We assessed the response of trees to four environmental variables differentially related to water availability in three steps: (1) identifying the shape of the response to individual environmental variables, (2) testing for artifacts in previous patterns due to spatial autocorrelation of presence–absence, and (3) identifying the environmental variable or combination of variables that best explained the pattern. We then classified the species with respect to their probability of occurrence along the gradient, and explored which components of the water cycle were likely to be driving the observed patterns. We found that 14 species were generalists, 16 drought tolerant, 9 intermediate, 3 water demanding and 8 showed mixed responses. Lateral flow and access to ground water most likely underlie such patterns. Our results confirm the key role played by water availability in tree species distribution. Water‐related niche differentiation seems to be crucial for maintaining the high diversity of this TDF. Abstract in Spanish is available at http://www.blackwell‐synergy.com/loi/btp .     

Effects of Season and Vegetation Type on Community Organization of Dung Beetles in a Tropical Dry Forest1

Dung beetles are important components of most terrestrial ecosystems. In tropical rain forests, dung beetle communities can be very rich in number of species and individuals, and they are known to be useful bioindicators of habitat disturbance. In contrast, very little is known about the organization of dung beetle communities in tropical dry forests. The aim of this study was to describe in detail the dung beetle community of a Mexican tropical dry forest and to assess the relative importance of rainfall seasonality and forest structure in affecting the temporal and spatial dynamics of this community. Dung beetles were captured with pitfall traps at the beginning of the rainy season, the middle of the rainy season, and the middle of the dry season, in two distinct forest types: deciduous forest (DF) and semideciduous forest (SDF) at the Estación de Biología Chamela. Both rainfall seasonality and forest structure affected the community organization of dung beetles. During both rainy periods, 14 species were captured, but only three during the dry season. Dung beetles captured during the dry season were only found in the SDF. When comparing the beginning and the middle of the rainy season, differences in abundance and guild structure were also observed between both periods and between forest types, but these differences were much less pronounced.     

Research Priorities for Neotropical Dry Forests1

Our understanding of the human and biophysical dimensions of tropical dry forest change and its cumulative effects is still in the early stages of academic discovery. The papers in this special section on Neotropical dry forests cover a wide range of sites and problems ranging from the use of multispectral and hyperspectral remote sensing platforms to the impact of hurricanes on tropical dry forest regeneration. Here, we present to the scientific community the results of a workshop on which research priorities for tropical dry forests were discussed. This discussion focuses on the need to develop linkages between remote sensing, ecological, and social science research. The incorporation of social sciences into ecological research could contribute dramatically to our understandings of tropical dry forests by providing important contextual information to ecologists, and by helping to develop an important science–policy–public nexus on which environmental management can succeed.     

Biomass,Carbon, and Nitrogen Pools in Mexican Tropical Dry Forest Landscapes

Tropical dry forest is the most widely distributed land-cover type in the tropics. As the rate of land-use/land-cover change from forest to pasture or agriculture accelerates worldwide, it is becoming increasingly important to quantify the ecosystem biomass and carbon (C) and nitrogen (N) pools of both intact forests and converted sites. In the central coastal region of México, we sampled total aboveground biomass (TAGB), and the N and C pools of two floodplain forests, three upland dry forests, and four pastures converted from dry forest. We also sampled belowground biomass and soil C and N pools in two sites of each land-cover type. The TAGB of floodplain forests was as high as 416 Mg ha^–1, whereas the TAGB of the dry forest ranged from 94 to 126 Mg ha^–1. The TAGB of pastures derived from dry forest ranged from 20 to 34 Mg ha^–1. Dead wood (standing and downed combined) comprised 27%–29% of the TABG of dry forest but only about 10% in floodplain forest. Root biomass averaged 32.0 Mg ha^–1 in floodplain forest, 17.1 Mg ha^–1 in dry forest, and 5.8 Mg ha^–1 in pasture. Although total root biomass was similar between sites within land-cover types, root distribution varied by depth and by size class. The highest proportion of root biomass occurred in the top 20 cm of soil in all sites. Total aboveground and root C pools, respectively, were 12 and 2.2 Mg ha^–1 in pasture and reached 180 and 12.9 Mg ha^–1 in floodplain forest. Total aboveground and root pools, respectively, were 149 and 47 kg ha^–1 in pasture and reached 2623 and 264 kg ha^–1 in floodplain forest. Soil organic C pools were greater in pastures than in dry forest, but soil N pools were similar when calculated for the same soil depths. Total ecosystem C pools were 306. The Mg ha^–1 in floodplain forest, 141 Mg ha^–1 in dry forest, and 124 Mg ha^–1 in pasture. Soil C comprised 37%–90% of the total ecosystem C, whereas soil N comprised 85%–98% of the total. The N pools lack of a consistent decrease in soil pools caused by land-use change suggests that C and N losses result from the burning of aboveground biomass. We estimate that in México, dry forest landscapes store approximately 2.3 Pg C, which is about equal to the C stored by the evergreen forests of that country (approximately 2.4 Pg C). Potential C emissions to the atmosphere from the burning of biomass in the dry tropical landscapes of México may amount to 708 Tg C, as compared with 569 Tg C from evergreen forests.     

Tree Diversity,Environmental Heterogeneity,and Productivity in a Mexican Tropical Dry Forest1

Species diversity–environmental heterogeneity (D–EH) and species diversity–productivity (D–P) relationships have seldom been analyzed simultaneously even though such analyses could help to understand the processes underlying contrasts in species diversity among sites. Here we analyzed both relationships at a local scale for a highly diverse tropical dry forest of Mexico. We posed the following questions: (1) are environmental heterogeneity and productivity related?; (2) what are the shapes of D–EH and D–P relationships?; (3) what are individual, and interactive, contributions of these two variables to the observed variance in species diversity?; and (4) are patterns affected by sample size, or by partitioning into average local diversity and spatial species turnover? All trees (diameter at breast height ≥5 cm) within twenty‐six 0.2‐ha transects were censused; four environmental variables associated with water availability were combined into an environmental heterogeneity index; aboveground standing biomass was used as a productivity estimator. Simple and multiple linear and nonlinear regression models were run. Environmental heterogeneity and productivity were not correlated. We found consistently positive log‐linear D–EH and D–P relationships. Productivity explained a larger fraction of among‐transect variance in species diversity than did environmental heterogeneity. No effects of sample size were found. Different components of diversity varied in sensitivity to environmental heterogeneity and productivity. Our results suggest that species' differentiation along water availability gradients and species exclusion at the lowest productivity (driest) sites occur simultaneously, independently, and in a scale‐dependent fashion on the tree community of this forest.     

Mating Systems of Three Tropical Dry Forest Tree Species1

Polymorphic allozyme loci were used to estimate outcrossing rates for three tree species from a disturbed dry forest in southern Costa Rica. Estimates of the multilocus outcrossing rates of Cedrela odorata and Jacaranda copaia were 0.969 and 0.982, respectively, and suggest that these species may be self-incompatible. The subcanopy tree Stemmadenia donnell-smithii also demonstrated little self-fertilization based on an estimated outcrossing rate of 0.896. Significant heterogeneity in pollen allele frequencies among maternal trees was detected for at least two enzyme loci for each species. A test of correlated mating between progeny of S. donnell-smithii revealed that all seeds within a fruit were singly sired. In addition, the low estimates of biparental inbreeding and significant differences in pollen and ovule allele frequencies for this species suggest that gene flow into the sampled forest fragment may occur. The implications of deforestation on the mating systems of these tropical tree taxa are discussed.     

The Role of Arboreal Seed Dispersal Groups on the Seed Rain of a Lowland Tropical Forest1

Most tropical plants produce fleshy fruits that are dispersed primarily by vertebrate frugivores. Behavioral disparities among vertebrate seed dispersers could influence patterns of seed distribution and thus forest structure. This study investigated the relative importance of arboreal seed dispersers and seed predators on the initial stage of forest organization–seed deposition. We asked the following questions: (1) To what degree do arboreal seed dispersers influence the species richness and abundance of the seed rain? and (2) Based on the plant species and strata of the forest for which they provide dispersal services, do arboreal seed dispersers represent similar or distinct functional groups? To answer these questions, seed rain was sampled for 12 months in the Dja Reserve, Cameroon. Seed traps representing five percent of the crown area were erected below the canopies of 90 trees belonging to nine focal tree species: 3 dispersed by monkeys, 3 dispersed by large frugivorous birds, and 3 wind‐dispersed species. Seeds disseminated by arboreal seed dispersers accounted for ca 12 percent of the seeds and 68 percent of the seed species identified in seed traps. Monkeys dispersed more than twice the number of seed species than large frugivorous birds, but birds dispersed more individual seeds. We identified two distinct functional dispersal groups, one composed of large frugivorous birds and one composed of monkeys, drop dispersers, and seed predators. These groups dispersed plants found in different canopy strata and exhibited low overlap in the seed species they disseminated. We conclude it is unlikely that seed dispersal services provided by monkeys could be compensated for by frugivorous birds in the event of their extirpation from Afrotropical forests.     

Forest Structure and Biomass of a Tropical Seasonal Rain Forest in Xishuangbanna,Southwest China1

Xishuangbanna is a region located at the northern edge of tropical Asia. Biomass estimates of its tropical rain forest have not been published in English literature. We estimated forest biomass and its allocation patterns in five 0.185–1.0 ha plots in tropical seasonal rain forests of Xishuangbanna. Forest biomass ranged from 362.1 to 692.6 Mg/ha. Biomass of trees with diameter at 1.3 m breast height (DBH) ≥ 5 cm accounted for 98.2 percent of the rain forest biomass, followed by shrubs (0.9%), woody lianas (0.8%), and herbs (0.2%). Biomass allocation to different tree components was 68.4–70.0 percent to stems, 19.8–21.8 percent to roots, 7.4–10.6 percent to branches, and 0.7–1.3 percent to leaves. Biomass allocation to the tree sublayers was 55.3–62.2 percent to the A layer (upper layer), 30.6–37.1 percent to the B layer (middle), and 2.7–7.6 percent to the C layer (lower). Biomass of Pometia tomentosa, a dominant species, accounted for 19.7–21.1 percent of the total tree biomass. The average density of large trees (DBH ≥100 cm) was 9.4 stems/ha on two small plots and 3.5 stems/ha on two large plots, illustrating the potential to overestimate biomass on a landscape scale if only small plots are sampled. Biomass estimations are similar to typical tropical rain forests in Southeast Asia and the Neotropics.     

Regional‐Scale Variation in Litter Production and Seasonality in Tropical Dry Forests of Southern Mexico1

Highly seasonal rainfall creates a pulse of litterfall in the southern Yucatan peninsula region, with cascading effects on the timing of essential nutrient fluxes, microbial dynamics, and vegetation growth. I investigated whether forest age or a regional environmental gradient related to rainfall has a greater effect on patterns of litterfall in this increasingly human‐dominated landscape. Litterfall was sampled in 10–13 stands in each of three locations spanning a rainfall gradient of ca 900–1400 mm/yr. Litter was collected monthly from November 1998 through January 2000 in mature forests and in secondary forests aged 2–25 yr. Despite a substantial precipitation gradient, age was the only significant predictor of annual litter mass. Two‐ to five‐yr‐old forests produced significantly less litter than 12–25‐yr‐old secondary forests (4.6 vs. 6.2 Mg/ha/yr), but the difference between older secondary forests and mature forests (9 percent) was not significant. Litter production increased with rainfall, but not significantly so. The pattern of litterfall was similar across locations and age classes, with a peak during late March or early April. However, litterfall seasonality was most pronounced in the old secondary and mature forests. Litterfall was more evenly distributed throughout the year in forests under 10 yr old. Seasonality of litterfall was also less pronounced at the wettest site, with less disparity between peak litterfall and off‐peak months. Seasonality was not related to soil texture. Forest age and rainfall are important drivers of litterfall dynamics; however, both litter mass and degree of seasonality depended more strongly on forest age. Thus, the impact of land‐use change on litter nutrient cycling is as great, if not greater, than the constraint imposed by the major natural environmental factor affecting tropical dry forests.     

Abundance and Leaf Damage Caused by Gall‐Inducing Insects in a Mexican Tropical Dry Forest1

We evaluated the temporal and spatial patterns of abundance and the amount of damage caused by gall‐inducing insects (GII) in deciduous and riparian habitats in a seasonal tropical dry forest in Mexico. Plants occurring in these habitats differ in their phenology and moisture availability. Deciduous habitats are seasonal and xeric, while riparian habitats are aseasonal and mesic. We found 37 GII species and each one was associated with a specific plant species. In total, 19 species (51.3%) were present in deciduous habitats, 13 species (35.2%) in riparian habitats, and only 5 species (13.5%) occurred in both. Abundance and leaf damage by GII were greater in deciduous than in riparian habitats during the wet season. For each GII species that occurred in both habitats, host plant species supported greater abundance and leaf damage by GII in deciduous habitats during the wet season. These results indicate a greater association of GII species with host plants in deciduous than in riparian habitats during the wet season. In riparian habitats, 11 plant species (61.1%) had greater density of GII in the dry than in the wet season. Similarly, leaf damage by GII was significantly greater in the dry than in the wet season in riparian habitats for 12 plant species (66.7%). Dry forest plants of riparian habitats presented two peaks of leaf‐flushing: GII colonized leaves produced in the first peak at the beginning of the wet season, and accumulated or recolonized leaves in the second peak at the beginning of the dry season. The levels of leaf damage by GII detected in this study in the rainy season were considerably higher than those obtained for folivorous insects in other neotropical forests, suggesting that this GII guild might have an important impact on their host plant species in this tropical community.     

Effects of Forest Fragmentation on the Colima Long-nosed Bat (Musonycteris harrisoni) Foraging in Tropical Dry Forest of Jalisco, Mexico1

We determined the effect of forest fragmentation on the nectarivorous Colima long‐nosed bat (Musonycteris harrisoni) by observing foraging behavior of this species in disturbed and undisturbed forests on the flowers of Ceiba grandiflora (Bombacaceae). The study was conducted in the area of the Chamela‐Cuixmala Biosphere Reserve in Jalisco, Mexico. Musonycteris harrisoni was observed visiting flowers during six nights (88 visits), exclusively in undisturbed forest. This species feeds on the nectar and serves as a pollinator of C. grandiflora.     

Flying Foxes Prefer to Forage in Farmland in a Tropical Dry Forest Landscape Mosaic in Fiji

To test flying fox adaptations to a habitat mosaic with extreme deforestation, the abundance, habitat choice and feeding behavior of the Pacific flying fox, Pteropus tonganus, were investigated across 16 islands of the Yasawa archipelago, Fiji. The habitat mosaic is formed by 4.3 percent tropical dry forest and 3.3 percent farmland, leaving exotic grasslands and stands of Leucaena leucocephala to overrun the vast majority of land. Pteropus tonganus abundance was high (5757 bats) despite deforestation and hunting. Roosting sites were restricted to native forest fragments. Grasslands and stands of L. leucocephala were completely void of bats at all times. The mean foraging density in farmland was four times higher than in forests and foraging competition was routinely observed in farmland but was extremely rare in forests. The author suggests that during the study, extensive foraging in farmland was supporting the high P. tonganus population. Additionally, the preferential foraging in farmland was responsible for the low foraging densities within forests and dramatically less intraspecies competition for forest resources. Further research is needed on seed dispersal within forests and to test for seasonal variations in bat abundance and feeding.     

Diversity and Coexistence of Ectoparasites in Small Rodents in a Tropical Dry Forest

To understand the mechanisms driving species diversity is central to community ecology. Here, we explored if habitat partitioning is associated with a species‐rich ectoparasite community in small rodents from a tropical dry forest in western Mexico. We trapped 199 mice in three 0.5 ha‐plots from eight small rodent species for every two months, from July 2011 to April 2012, and collected their ectoparasites. We identified 17 species of mites, two sucking lice species, two phoretic species, and one commensal species. The most abundant ectoparasite species was Steptolaelaps liomydis, representing 42 percent of all ectoparasites collected; seven ectoparasite species had < 10 individuals. Eighteen ectoparasite species (of 22 species) were collected from the most abundant rodent Liomys pictus. C‐score and the number of checkerboard species pairs were significantly higher against a random expectation. Ectoparasite species in L. pictus mice showed host microhabitat partitioning; Fahrenholzia ehrlichi and Fahrenholzia texana were found only in the anterior dorsal area, Ornithonysus sp. occurred along the dorsal part, Ixodes species were restricted to the ears, and Steptolaelaps liomydis was found throughout the body. We also identified ectoparasite communities with distinct species composition in two rodent species that use contrasting macrohabitats (L. pictus, strictly terrestrial; Peromyscus perfulvus, mostly arboreal). The remaining and low abundant rodent species showed a species‐poor ectoparasite community composition. We conclude that habitat partitioning at both macro and microhabitat scales appeared to characterize the species‐rich ectoparasite community. Conversely, most rodent host species with low abundances showed a species‐poor ectoparasite community.