Neotropical Wet Forests Research Articles

El Niño oscillations impact anti-predator defences to alter survival of an herbivorous beetle in a neotropical wet forest

Abstract Little is known about the effects of El Niño-Southern Oscillation (ENSO) on tropical insect communities, even though they are suffering rapid declines in complexity and stability due to climate change. We explore the impact of fluctuations in local climate imposed by ENSO on the performance of herbivore defences mediating enemy interactions. In a widespread rainforest edge community, we quantified the mortality caused by five enemy guilds on the immature stages of the herbivorous beetle, Acromis sparsa. ENSO was a significant determinant of beetle mortality. During warmer, drier El Niño years, the survival of beetles decreased. This was due to increased egg parasitism by wasps, which reduced hatching. Additionally, ant predation on beetle larvae increased. Flies and wasps were in competition for larval prey in wetter, cooler La Niña years. Experimental removal of maternal guards or chemical shields revealed which ENSO-related parameters predicted larval mortality. Guarding was most effective against social wasps in La Niña, whereas shields proved most effective in El Niño. Two ENSO-related defence–enemy breakdowns occurred: (1) decoupling whereby the efficacy of a narrow defensive adaptation was reduced to increase mortality, and (2) mismatching whereby the resistance of a narrow defensive adaptation against non-targeted enemies was further reduced to increase mortality. These results highlight that defence efficacy against natural enemies can vary predictably with biotic and abiotic environmental conditions. ENSO events will increase breakdowns in defence-mediated interactions, shifts in competition among enemies, and species loss.

Effects of climate change on plant resource allocation and herbivore interactions in a Neotropical rainforest shrub

AbstractClimate change is a mounting global issue, but its consequences will be variable across regions. Tropical species are hypothesized to have reduced climatic adaptability and plasticity. Yet, relative to temperate species, less is understood about how they will respond to climate change. Rising temperature and atmospheric CO2 could impact plant–herbivore systems directly by altering species traits or abundances, or the effects could be indirect by altering the strength and direction of the relationships that govern organismal strategies and interactions. Using open‐top chambers in a Neotropical wet forest, we applied a full‐factorial combination of active warming and CO2 fertilization to investigate the above‐ground, short‐term effects of climate change on plant–herbivore interactions in a common Neotropical shrub, Piper generalense. We aimed to answer two main questions: (1) Could climate change alter plant–herbivore systems through direct effects on plant growth rate, chemical defense, and/or insect herbivore damage rate? and (2) Could climate change affect plant–herbivore systems indirectly by altering (a) the strength of plant resource allocation trade‐offs between growth and defense or (b) the effectiveness of plant chemical defense against herbivory? None of the microclimate treatments had direct effects on plant growth, chemical defense, or herbivore damage. However, we did observe a positive relationship between growth and chemical defense in treatments mimicking climate‐change conditions, which partially supports the growth–differentiation balance hypothesis. We did not detect any effects of treatments on the effectiveness of plant chemical defense against herbivory. It appears that, in this system, increased CO2 concentration and temperature may cause indirect, cascading consequences, even where direct effects are not observable. We recommend more climate‐change experiments addressing multi‐trophic interactions that focus not only on the direct responses of organisms but also on the ways in which climate change can restructure the relationships that govern complex biotic systems.

Habitat-dependent effects of predatory spiders on prey frogs in a Neotropical wet forest

AbstractIn seasonal wet Neotropical forests, many studies have suggested that species-rich terrestrial frog assemblages are regulated bottom-up by the abundance of leaf litter. However, terrestrial frogs are prey to a diverse community of predators, and no studies have tested for top-down effects of predators on this or other anuran assemblages. Here, we used an extensive field dataset to model the relative contribution of food resources, microhabitat resources and predators towards the occupancy and detection of two frog species (Craugastor bransfordiiandOophaga pumilio) at La Selva, Costa Rica. Frog occupancy was most strongly influenced by predatory spiders and secondarily influenced by the abundance of leaf litter. Predators exerted stronger effects on frogs than food resources, and frogs avoided predators more as leaf litter decreased. Detection probability was elevated when predators were present. We found support for bottom-up effects of leaf litter on the terrestrial frog assemblage, but top-down effects by predators exerted stronger effects on frog occupancy and detection. Because predator avoidance varied along a resource gradient, predator and resource effects appear to be dependent, supporting interactions between top-down and bottom-up mechanisms. Climate-driven decreases in leaf litter may drive decreased availability of frog refugia and increased interactions between frogs and predators.

Precipitation-drainage cycles lead to hot moments in soil carbon dioxide dynamics in a Neotropical wet forest.

Soil CO2 concentrations and emissions from tropical forests are modulated seasonally by precipitation. However, subseasonal responses to meteorological events (e.g., storms, drought) are less well known. Here, we present the effects of meteorological variability on short-term (hours to months) dynamics of soil CO2 concentrations and emissions in a Neotropical wet forest. We continuously monitored soil temperature, moisture, and CO2 for a three-year period (2015-2017), encompassing normal conditions, floods, a dry El Niño period, and a hurricane. We used a coupled model (Hydrus-1D) for soil water propagation, heat transfer, and diffusive gas transport to explain observed soil moisture, soil temperature, and soil CO2 concentration responses to meteorology, and we estimated soil CO2 efflux with a gradient-flux model. Then, we predicted changes in soil CO2 concentrations and emissions under different warming climate change scenarios. Observed short-term (hourly to daily) soil CO2 concentration responded more to precipitation than to other meteorological variables (including lower pressure during the hurricane). Observed soil CO2 failed to exhibit diel patterns (associated with diel temperature fluctuations in drier climates), except during the drier El Niño period. Climate change scenarios showed enhanced soil CO2 due to warmer conditions, while precipitation played a critical role in moderating the balance between concentrations and emissions. The scenario with increased precipitation (based on a regional model projection) led to increases of +11% in soil CO2 concentrations and +4% in soil CO2 emissions. The scenario with decreased precipitation (based on global circulation model projections) resulted in increases of +4% in soil CO2 concentrations and +18% in soil CO2 emissions, and presented more prominent hot moments in soil CO2 outgassing. These findings suggest that soil CO2 will increase under warmer climate in tropical wet forests, and precipitation patterns will define the intensity of CO2 outgassing hot moments.

Community assembly of glass frogs (Centrolenidae) in a Neotropical wet forest: a test of the river zonation hypothesis

Abstract:The river zonation hypothesis predicts that abiotic and biotic conditions along riparian gradients drive variation in animal communities. Glass frogs are a diverse group of Neotropical anurans that use riparian habitats exclusively for oviposition and larval development, but little is known about how glass frog communities are distributed across riparian gradients. Here, we measured glass frog community assembly across a gradient of riparian habitats from first- to fifth-order streams at La Selva Biological Station, Costa Rica. We performed repeated nocturnal frog calling surveys and built occupancy andN-mixture abundance models to test for varying patterns of species occupancy, community assembly, species richness (α-diversity) and species turnover (ß-diversity). We observed significant differences in patterns of species occupancy and community assembly across a stream-order gradient: occupancy of two species increased with stream order (Teratohyla pulverata,Hyalinobatrachium fleischmanni), one species decreased (Teratohyla spinosa), and one species did not vary (Espadarana prosoblepon). We evaluated four a priori hypotheses describing how α- and ß-diversity of centrolenids are shaped across the riparian gradient; our data were most consistent with a pattern of nested assemblages and increasing species richness along the riparian gradient. Species-specific patterns of occupancy and abundance resulted in assemblage-level differences consistent with theoretical predictions for highly aquatic organisms along riparian gradients.

Phosphorus retention in a lowland Neotropical stream following an eight-year enrichment experiment

AbstractHuman alteration of the global P cycle has led to widespread P loading in freshwater ecosystems. Much research has been devoted to the capacity of wetlands and lakes to serve as long-term sinks for P inputs from the watershed, but we know much less about the potential of headwater streams to serve in this role. We assessed storage and retention of P in biotic and abiotic compartments after an 8-y experimental P addition to a 1st-order stream in a Neotropical wet forest. Sediment P extractions indicated that nearly all P storage was in the form of Fe- and Al-bound P (∼700 μg P/g dry sediment), similar to nearby naturally high-P streams. At the end of the enrichment, ∼25% of the total P added over the 8-y study was still present in sediments within 200 m of the injection site, consistent with water-column measurements showing sustained levels of high net P uptake throughout the experiment. Sediment P declined to baseline levels (∼100 μg P/g dry sediment) over 4 y after the enrichment ended. Leaf-lit...

Molecular data reveal spatial and temporal patterns of diversification and a cryptic new species of lowland Stenocercus Duméril & Bibron, 1837 (Squamata: Tropiduridae).

Phylogenetic studies have uncovered biogeographic patterns and the associated diversification processes of Neotropical wet forest taxa, yet the extensive open and drier biomes have received much less attention. In the Stenocercus lizard radiation, restricted sampling and phylogenetic information have limited inferences about the timing, spatial context, and environmental drivers of diversification in the open and dry lowland settings of eastern and southern South America. Based on new DNA sequence data of previously unsampled species, we provide an updated historical biogeographic hypothesis of Stenocercus. We infer phylogenetic relationships, estimate divergence times, and track ancestral distributions, asking whether cladogenetic events within the genus correlate to reported shifts in South American landscapes during the past 30millionyears, focusing in the open and drier areas. To examine correlations between genetic and ecological divergence, we extracted environmental data from occurrence records and estimated climatic envelopes occupied by lowland taxa. Our results suggest that Stenocercus began to diversify around the South American Midwest by the late Oligocene. We recovered two main lowland and two main Andean clades within the genus; within both Andean clades, most cladogenetic events date back to the Miocene, synchronously with the most intense phase of Andean uplift. In the western clade of lowland Stenocercus, species ranges and divergence times are consistent with major landscape shifts at the upper Guaporé and Paraguay River basins as a result of Andean orogeny, suggesting vicariant speciation. By contrast, in the 'horned' lowland clade, we find evidence that dispersal and ecological differentiation have shaped species divergences and current ranges in the Brazilian Cerrado, Caatinga, Pampas and Atlantic Forest, possibly under a vanishing refuge scenario. Lastly, our phylogenetic results indicate two divergent clades within the formerly recognized taxon S. sinesaccus, and further evaluation of morphological data corroborates the existence of a distinct, new species of Stenocercus, here described. The new taxon occurs in the Chapada dos Parecis massif in the Brazilian states of Mato Grosso and Rondônia.

Diversity in neotropical wet forests during the Cenozoic is linked more to atmospheric CO 2 than temperature

Models generally predict a response in species richness to climate, but strong climate-diversity associations are seldom observed in long-term (more than 10(6) years) fossil records. Moreover, fossil studies rarely distinguish between the effects of atmospheric CO2 and temperature, which limits their ability to identify the causal controls on biodiversity. Plants are excellent organisms for testing climate-diversity hypotheses owing to their strong sensitivity to CO2, temperature and moisture. We find that pollen morphospecies richness in an angiosperm-dominated record from the Palaeogene and early Neogene (65-20 Ma) of Colombia and Venezuela correlates positively to CO2 much more strongly than to temperature (both tropical sea surface temperatures and estimates of global mean surface temperature). The weaker sensitivity to temperature may be due to reduced variance in long-term climate relative to in higher latitudes, or to the occurrence of lethal or sub-lethal temperatures during the warmest times of the Eocene. Physiological models predict that productivity should be the most sensitive to CO2 within the angiosperms, a prediction supported by our analyses if productivity is linked to species richness; however, evaluations of non-angiosperm assemblages are needed to more completely test this idea.

Stability structures tropical woody plant diversity more than seasonality: Insights into the ecology of high legume-succulent-plant biodiversity

Phylogenies of legume taxa are ecologically structured along a tropical seasonality gradient, which suggests phylogenetic niche conservatism. This seasonality gradient spans Neotropical wet forests, savannas, and highly seasonal drought-prone woody vegetation known as the succulent biome. Ecological phylogenetic structure was investigated using a community phylogenetic approach. We further analyzed bioclimatic and other independent variables that potentially explained phylogenetic beta diversity among 466 floristic sites that spanned the savanna and succulent biomes in eastern South America. Explanatory variables were selected using variance inflation factors, information criteria, and the ability to explain both species and phylogenetic beta diversity. A model involving annual precipitation suggests that a threshold of <1200mm explains community phylogenetic structure along the savanna–succulent biome transition. Variables involving temperatures or measures of seasonality were notably lacking from top-ranked models. The abundance and diversity of legumes across the tropical seasonality gradient suggest that a high nitrogen metabolism confers an advantage in one of two ways, both of which are related to rapid growth rates. Legumes adapted to the succulent biome may be responding to regular post-dry-season leaf-flush opportunities. Legumes adapted to the savanna biome may be responding to intermittent post-disturbance growing opportunities. A seasonal predominance of leaf flushing by woody plants implicates the role of ecological stability in the succulent biome because of the need to recover the cost of regenerating short-lived leaves. Ecological stability may be the fundamental cause of ecological phylogenetic structure across the tropical seasonality gradient and required for maintaining high levels of both leaf-flushing legume and succulent plant biodiversity.

Ecology and Distribution of Neotropical Podocarpaceae

Podocarps are a frequent, but rarely a dominant, component of neotropical wet forests extending from South America into central Mexico and the Greater Antilles. Although podocarps are often considered to be predominantly montane taxa, several species occur in lowland forest and are locally abundant on some Pacific and Atlantic coastal islands in Central America. Here we review literature on the origins and distribution of neotropical podocarps and highlight their apparent association with resourcepoor environments. As a consequence of forest conversion and logging, many podocarps that were already habitat specialists are now further restricted to small and increasingly fragmented populations. Unfortunately, there is little information on the regeneration ecology of podocarps with which to assess the recruitment potential of these populations. An exception is the long-term studies of the dynamics of Podocarpus urbanii, a common species in montane forest in Jamaica. Podocarpus urbanii is moderately shade tolerant and successfully regenerates beneath undisturbed forest. The low juvenile mortality rate of P. urbanii, coupled with relatively high diameter growth, suggests that this species and possibly other podocarps may have greater utility for reforestation than is currently recognized.

'Nucleating’ succession in recovering neotropical wet forests: The legacy of remnant trees

AbstractQuestion: What is the influence of remnant trees on secondary forest structure and composition in tropical pastures many years after abandonment?Location: Neotropical lowland wet forest, La Selva Biological Station, Costa Rica.Methods: Tree and sapling density, basal area, and species richness were quantified at three distances from remnant trees, 0–10 m (inner), 20–30 m (intermediate), and ca. 50 m (distal) zones. A total of 15 remnant trees were sampled in pastures ~23 years after abandonment.Results: Tree density decreased along a gradient from inner (1117 ± 377 individuals/ha) to distal (592 ± 282 individuals/ha) zones, and the number of large‐seeded individuals (seeds &gt; 1 cm diameter) was significantly greater in the inner zone. Basal area of tree individuals was greater in the inner (25.6 ± 12 m2/ha) and intermediate (28.3 ± 15.6 m2/ha) zones than the distal zone (14.7 ± 7.2 m2/ha), but there were no differences between inner and intermediate zones. Similar patterns are reported for species richness. Additionally, saplings (1 ‐ 5 cm DBH) had higher density directly beneath and adjacent to remnants, suggesting that remnant trees can affect recruitment even many years after pasture abandonment and the formation of a surrounding secondary forest.Conclusions: Results indicate that remnant trees facilitate forest recovery over a broad temporal range, and appear to ‘nucleate’ forest regeneration by expanding their sphere of influence outward over time.

MULTIGENERATIONAL GENETIC ANALYSIS OF TROPICAL SECONDARY REGENERATION IN A CANOPY PALM

Iriartea deltoidea (Arecaceae) is an abundant canopy palm with a wide geographic distribution in Neotropical wet forests. We analyzed the genetic profile across three generations of Iriartea within a 43-ha area encompassing two areas of second-growth and adjoining old-growth forest at La Selva Biological Field Station in northeastern Costa Rica. A total of 311 reproductively mature trees, 99 large saplings, 207 small saplings, and 601 seedlings were genotyped using 141 AFLP loci. Parentage analysis revealed high dispersal distances, both for seed (over 2.3 km) and pollen (over 3.8 km), indicating a large genetic neighborhood within La Selva Biological Station. In a 20-ha area of second growth, the founding palm population was dominated by a small number of parental trees located in the adjacent old-growth forest; two old-growth trees contributed 48% of the second-growth genes. The genetic diversity of reproductively mature trees in this second-growth forest was significantly reduced compared to adjacent old-growth forest. Within 400 m of the border with old-growth forest, we observed a similar reduction of genetic diversity in saplings, and an even greater loss of genetic diversity in the second generation of seedlings. Nearly half of these seedlings were offspring of local parents. In contrast, in the distant portion of second-growth forest (400-800 m from the old-growth border), parentage analysis showed that 40% of seedlings originated from outside the study area and only 10% were offspring of local parents. These high levels of gene flow maintained genetic diversity in saplings and seedlings similar to levels observed in old-growth forest. Our findings highlight the importance of gene flow from diverse seed and pollen sources for sustaining levels of genetic diversity of tree populations in second-growth forests.

Placing an Omnivore in a Complex Food Web: Dietary Contributions to Adult Biomass of an Ant1

ABSTRACT Workers of Paraponera clavata, a common Neotropical ant, collect both nectar and insect prey. Previous reports show that nectar accounts for up to 90 percent of the ants' food loads, while calculations suggest that nectar contributes only 10 percent of colonies' energy supply. We assessed the trophic source of carbon and nitrogen in adult workers using stable isotope analysis. Carbon in adult workers was largely derived from plant sources. Worker nitrogen isotopic ratios varied significantly among colonies and were enriched compared to prey. Prey nitrogen isotope ratios suggest considerable intercolonial variation in diet, with some colonies collecting prey from lower trophic levels than other colonies. The importance of nectar as a source of metabolic carbon in adult worker biomass, coupled with the high frequency of nectar collection, supports the conclusion that omnivory is a key to supporting this species' biomass in Neotropical wet forests.

Wood specific gravity and anatomy in Heliocarpus appendiculatus (Tiliaceae)

Specific gravity exhibits extremely large radial increases with distance from the pith in Heliocarpus appendiculatus Turcz. (Tiliaceae), a pioneer of neotropical wet forests. To determine some of the wood anatomical changes associated with this increase, wood samples taken at breast height from three trees were divided into 1.0‐cm‐long segments from pith to bark. Measurements were made of fiber wall thickness, fiber lumen diameter, and percentages of fibers, axial parenchyma, ray parenchyma, and vessels on sections prepared from each segment. The extreme radial increases in specific gravity were associated with increases in fiber wall thickness, decreases in fiber diameter, decreases in fiber lumen diameter, and changes in the relative proportions of fibers and parenchyma. The increase in percent fiber concomitant with a decrease in axial parenchyma was the most important contributor to the increase in specific gravity in this species. The best predictor of specific gravity was percent fibers (r = 0.91, 0.92, 0.94) or percent axial parenchyma (r = ‐0.92, ‐0.91, ‐0.95), two variables that were highly intercorrelated (r = ‐0.95, ‐0.98, ‐0.99).

Distribution and effects on tree growth of lianas and woody hemiepiphytes in a Costa Rican tropical wet forest

ABSTRACTWe evaluated occurrence and abundance of lianas and woody hemiepiphytes on canopy and emergent tree species in primary tropical wet forest at the La Selva Biological Station, Costa Rica. Two pioneers, Cecropia obtusifolia and C. insignis, lacked both lianas and hemiepiphytes. The seven non-pioneer species differed significantly in their loads of lianas and hemiepiphytes. For all non-pioneer species, two measures of liana and hemiepiphyte loads (the percentage of the crown occupied and the combined basal area of descending hemiepiphyte and liana roots and stems) increased significantly with tree diameter. In all non-pioneer species, most trees ∵ 70 cm diameter (50–97%) were colonized. Lianas occupied more trees and had a smaller mean host diameter than did hemiepiphytes; however, basal area of descending roots or stems was equivalent for the two life forms in occupied trees. We used partial correlations controlling for tree diameter to evaluate the relationship between annual tree diameter growth and loads of hemiepiphytes and lianas for six non-pioneer species. Five of the six species showed a significant negative correlation between loads and diameter growth. Existing published data show that the high incidence of lianas and hemiepiphytes at La Selva is paralleled in most other Neotropical wet forests.

Leaf Herbivores Decrease Fitness of a Tropical Plant

Damage by insect herbivores to neighboring individuals of the shrub Piper arieianum in a neotropical wet forest varies greatly. This differential damage has a genetic basis and results in a 2-year decrease in growth, seed production, and seed viability, with larger plants recovering before smaller plants. The results provide evidence that leaf herbivores represent a potentially strong selective force for the evolution of plant defenses.

The Behavioral Response Race Between Predator and Prey

On tente de preciser par la methode de regression lineaire ce qui determine les rapports de force relatifs en ce qui concerne les relations predateur-proie (on donne comme exemple Notonectidae-Culicidae)

The etTects of food presentation and microhabitat upon resource monopoly in a ground-foraging ant (Hymenoptera: Formicidae) community

In Neotropical wet forests several species of omnivorous, resource-defending ants, live and forage in close proximity to one another. Although the forest floor is heterogeneous in microhabitat and food quantity, little is known about the impact of microhabitat and food variation upon resource monopoly among ants. We investigated how food type and microhabitat influence food monopoly in resource-defending ants in old-growth tropical wet forest in the Caribbean lowlands of Costa Rica. We measured several microhabitat characteristics at 66 points in a 0.5 hectare plot, and baited each point with two categories of tuna bait. These baits were presented in "split" and "clumped" arrangements. We measured the frequency of bait monopoly by a single species, as well as the number of recruited ant foragers at a bait. Out of five common species, two (Wasmannia auropunctata and Pheidole simonsi) more frequently monopolized one bait type over the other, and one (P. simonsi) recruited more ants to the split baits. We then considered the recruitment response by all ant species in the community. We found that the frequency of monopoly, sharing, and the absence of ants at a given point in the rainforest differed with bait type. The frequency of monopoly was associated with microhabitat type in two out of eight microhabitat variables (leaf litter depth and palms); variation in two other types (canopy tree distance and leafcutter ant trails) was associated with changes in forager number. In at least two ant species, food presentation affected monopoly at baits; among all resource-defending ants, the microhabitats where ants foraged for food and the type of food located determined in part the frequency of monopoly and the number of foragers at the food item. These results suggest that the location and presentation of food items determines in part which ant species will utilize the resource.