A Late Holocene Pollen and Charcoal Record from La Selva Biological Station, Costa Rica

Pollen and charcoal analysis of a 5.9-m-long sediment profile from a swamp on an alluvial terrace on the edge of La Selva biological station, Costa Rica (10°26'23" N, 84°00'24" W, 36 m asl), documents three millennia of human and natural disturbance within a lowland tropical rain forest. The record indicates that the highly diverse rain forest that presently surrounds the Cantarrana swamp regrew following forest clearing and maize agriculture that ended only a few centuries ago. The first maize pollen appears in sediments deposited ∼1070 calendar-year BP. Older sediments below the 'maize zone' contain macroscopic charcoal, abundant microscopic charcoal, and possible pollen indicators of forest clearance, perhaps signaling local cultivation of root crops that left no pollen in the sediments. Interpretation of local archaeology indicates that La Selva reserve may have been most heavily populated during the El Bosque and La Selva regional archaeological phases from 2250–950 yr BP. However, the distribution of maize pollen in the sediments is clear evidence that the reserve was also occupied during the later La Cabaña phase (950–400 yr BP), from which few artifacts have yet surfaced. Natural forest disturbance from treefalls and stream dynamics, and hydrological shifts associated with late-Holocene climate variability, form a backdrop to the human land-use history preserved in the Cantarrana sediment profile.     

Pleistocene Plant Fossils in and near La Selva Biological Station, Costa Rica1

Radiocarbon dating and ⁴⁰Ar/³⁹Ar analysis of overlying tephra indicate that plant fossil assemblages exposed by stream erosion and well construction in and near La Selva Biological Station in eastern lowland Costa Rica are Pleistocene in age. We identified plant taxa based on wood, leaves, fruits, seeds, pollen, and spores examined from three sites at ca 30 m elevation. Extrapolating from modern ranges and surface temperature lapse rates suggests paleotemperatures 2.5–3.1°C cooler than at present     

Insights into the Witheringia solanacea (Solanaceae) Complex in Costa Rica. II. Insect Visitors and Pollination Biology of W. asterotricha and W. meiantha1

Isolating mechanisms are important in maintaining the taxonomic integrity of closely related sympatric taxa. A previous study found strong post‐zygotic isolating barriers between two species, Witheringia asterotricha and W. meiantha, of the W. solanacea (Solanaceae) species complex in Costa Rica. This study examines the presence of pre‐zygotic barriers between the two species at La Selva Biological Station in Costa Rica. Both species offer pollen and nectar as floral rewards and are visited primarily by solitary or semi‐social bees, some of which sonicate (“buzz”) the anthers to discharge pollen. No evidence was found for phenological differences in flowering time between W. asterotricha and W meiantha, but pre‐zygotic factors, such as ethological isolation and possibly fine‐scale ecological or geographic barriers, may be responsible for restricting gene flow between the two species.     

Rainfall and parasitic wasp (Hymenoptera: Ichneumonoidea) activity in successional forest stages at Barro Colorado Nature Monument, Panama, and La Selva Biological Station, Costa Rica

1 In 1997, we ran two Malaise insect traps in each of four stands of wet forest in Costa Rica (two old‐growth and two 20‐year‐old stands) and four stands of moist forest in Panama (old‐growth, 20, 40 and 120‐year‐old stands). 2 Wet forest traps caught 2.32 times as many ichneumonoids as moist forest traps. The average catch per old‐growth trap was 1.89 times greater than the average catch per second‐growth trap. 3 Parasitoids of lepidopteran larvae were caught in higher proportions in the wet forest, while pupal parasitoids were relatively more active in the moist forest. 4 We hypothesize that moisture availability is of key importance in determining parasitoid activity, community composition and trophic interactions.     

High foliicolous lichen alpha-diversity on individual leaves in Costa Rica and Amazonian Ecuador

Two individual, dicotyledoneous leaves (125 and 98 cm² in size) and one composed palm leaf (c. 6800 cm² in size), gathered at La Selva Biological Station, Costa Rica, and Jatun Satcha Biological Station, Amazonian Ecuador, were screened for small-scale foliicolous lichen diversity. On the dicotyledoneous leaf from Costa Rica, 49 lichens and one lichenicolous fungus were found, while a comparable leaf from Ecuador revealed 46 lichens and two lichenicolous fungi. The palm leaf yielded 81 lichens and one lichenicolous fungus. This is the highest alpha-diversity so far reported for foliicolous lichens on individual leaves and invites for comparison with tree diversity in tropical rain forests. Due to the high proportion of species represented by a single thallus, the taxonomic diversity of lichens on individual leaves (or trees in selected plots) cannot be self-supporting, but reflects a high degree of dispersion or entropy within the community of which the individual leaf (or selected plot) is part. Diversity is therefore fractal, showing similar patterns at different scales, each part of a given community reflecting the entire community. Thus, mechanisms that result in high small-scale diversity must be looked for at the community level.     

Vertebrate Fruit Removal and Ant Seed Dispersal in the Neotropical Ginger Renealmia alpinia (Zingiberaceae)*

Plants frequently display fruit characteristics that support multiple seed‐dispersal syndromes. These ambiguous characteristics may reflect the fact that seed dispersal is usually a complex process involving multiple dispersers. This is the case for the Neotropical ginger Renealmia alpinia (Zingiberaceae). It was originally suggested that the aromatic fruits of R. alpinia located at the base of the plant are adapted for terrestrial mammal seed dispersal. However, the dark‐purple coloration of the fruits and bright orange aril surrounding the seeds suggest that birds may play a role in R. alpinia seed dispersal. At La Selva Biological Station, Costa Rica, we used camera traps to record vertebrate visits to infructescences of R. alpinia. Most visitors were toucans and aracaris (Ramphastidae). However fruits were also removed by terrestrial mammals (coatis and armadillos). In addition to vertebrate fruit removal, some of the fruits dehisce and the seeds that fall on the ground are dispersed by ants. Fruitfall traps showed that 77 percent of fruits are removed by vertebrates. However, 15 percent of fruits fall to the base of parent plants to be potentially dispersed by ants. Experiments using a laboratory ant colony showed that ants are effective seed dispersers of R. alpinia. Ant seed manipulation increased germination success and reduced time to germination. In conclusion, primary seed dispersal in the Neotropical ginger R. alpinia is mostly performed by birds, additionally ants are effective dispersers at short distances. Seed dispersal in R. alpinia is a complex process involving a diverse array of dispersal agents.     

Sodium‐specific foraging by leafcutter ant workers (Atta cephalotes,Hymenoptera: Formicidae)

1. Sodium is often a limiting nutrient for terrestrial animals, and may be especially sought by herbivores. Leafcutter ants are dominant herbivores in the Neotropics, and leafcutter foraging may be affected by nutritional demands of the colony and/or the demands of their symbiotic fungal mutualists. We hypothesized that leafcutter colonies are sodium limited, and that leafcutter ants will therefore forage specifically for sodium. 2. Previous studies demonstrated that leafcutter Atta cephalotes Linnaeus workers preferentially cut and remove paper baits treated with NaCl relative to water control baits. Atta cephalotes colonies in this study were presented with baits offering NaCl, Na₂SO₄, and KCl to test whether leafcutters forage specifically for sodium. Sucrose and water were used as positive and negative controls, respectively. 3. Atta foragers removed significantly more of the baits treated with NaCl and Na₂SO₄ than the KCl treatment, which did not differ from water. The NaCl and Na₂SO₄ treatments were collected at similar rates. We conclude A. cephalotes forage specifically for sodium rather than for anions (chloride) or solutes in general. This study supports the hypothesis that leafcutter ants are limited by, and preferentially forage for, sodium.     

Increasing Liana Abundance and Basal Area in a Tropical Forest: The Contribution of Long‐distance Clonal Colonization

Recent evidence suggests that liana abundance and biomass are increasing in Neotropical forests, representing a major structural change to tropical ecosystems. Explanations for these increases, however, remain largely untested. Over an 8‐yr period (1999–2007), we censused lianas in nine, 24 × 36 m permanent plots in old‐growth and selectively logged forest at La Selva Biological Station, Costa Rica to test whether: (1) liana abundance and basal area are increasing in this forest; (2) the increase is being driven by increased recruitment, decreased mortality, or both; and (3) long‐distance clonal colonization explains the increase in liana abundance and basal area. We defined long‐distance clonal colonization as lianas that entered and rooted in the plots as vegetative propagules of stems that originated from outside or above the plot, and were present in 2007, but not in 1999 or 2002. Our hypotheses were supported in the old‐growth forest: mean liana abundance and BA (≥1 cm diameter) increased 15 and 20 percent, respectively, and clonal colonization from outside of the plots contributed 19 and 60 percent (respectively) to these increases. Lianas colonized clonally by falling vertically from the forest canopy above or growing horizontally along the forest floor and re‐rooting—common forms of colonization for many liana species. In the selectively logged forest, liana abundance and BA did not change, and thus the pattern of increasing lianas may be restricted to old‐growth forests. In summary, our data support the hypothesis that lianas are increasing in old‐growth forests, and that long‐distance clonal colonization is a major contributor.     

Vascular Epiphyte Communities in the Inner-Crown of Hyeronima alchorneoides and Lecythis ampla at La Selva Biological Station, Costa Rica

The community composition and percent cover of vascular epiphytes were examined in relation to photosynthetic photon flux density (PFD), temperature, vapor pressure, and tree characteristics in the inner-crown of two emergent tree species, Hyeronima alchorneoides and Lecythis ampla, at La Selva Biological Station, Costa Rica. A total of 53 species were found in a total sampled branch area of 32 m² in eight trees—four trees per species. Community composition varied both among individuals of the same tree species and between tree species. However, percent cover patterns of vascular epiphytes were significantly different between the two tree species; Hyeronima had a significantly greater percent cover of epiphytes than Lecythis . The higher percent light transmittance as well as lower humidity in Lecythis are likely causes of its lower percent cover of epiphytes.     

Tree functional traits as predictors of microburst-associated treefalls in tropical wet forests

On 19 May 2018, a microburst caused 600 isolated forest gaps in a Costa Rican tropical forest. We surveyed fallen and standing trees within gaps to determine whether certain variables are associated with treefalls. Our results highlight considerations for future research to understand the impacts of microbursts in tropical forests.     

Density, Distribution, and Attributes of Tree Cavities in an Old-Growth Tropical Rain Forest

Tree cavities are a critical resource for many animals, especially as nesting sites for birds. Patterns of cavity distribution in temperate forests are well studied, yet little is known of cavities in tropical forests, despite a hypothesized decrease in cavity availability with decreasing latitude. We studied cavity density and distribution in a wet lowland tropical forest in Costa Rica and compared our results with estimates from forests around the world. Cavities at our site were common, occurred frequently in living trees, and were often formed by damage or decay rather than by woodpeckers. Most cavities had small openings, and woodpecker-created cavities were nonrandomly oriented. Contrary to prediction, cavity density appears to increase from the poles to the tropics. We suggest potential mechanisms to explain these patterns.     

Quantifying mortality of tropical rain forest trees using high-spatial-resolution satellite data

Assessment of forest responses to climate change is severely hampered by the limited information on tree death on short temporal and broad spatial scales, particularly in tropical forests. We used 1‐m resolution panchromatic IKONOS and 0.7‐m resolution QuickBird satellite data, acquired in 2000 and 2002, respectively, to evaluate tree death rates at the La Selva Biological Station in old‐growth Tropical Wet Forest in Costa Rica, Central America. Using a calibration factor derived from ground inspection of tree deaths predicted from the images, we calculated a landscape‐scale annual exponential death rate of 2.8%. This corresponds closely to data for all canopy‐level trees in 18 forest inventory plots, each of 0.5 ha, for a mostly‐overlapping 2‐year period (2.8% per year). This study shows that high‐spatial‐resolution satellite data can now be used to measure old‐growth tropical rain forest tree death rates, suggesting many new avenues for tropical forest ecology and global change research.     

A preliminary survey of the Chironomidae (Diptera) of Costa Rica,with emphasis on the lotic fauna

Extensive collections of Chironomidae were made in Costa Rica, Central America, during 1986 and 1987. Fifty-five genera and at least 148 species belonging to the subfamilies Podonominae, Tanypodinae, Orthocladiinae and Chironomidae were found. Chironominae and Orthocladiinae predominated. Only one species of Podonominae was collected. Tanypodinae was represented by many genera, but species richness was low.Cricotopus was the most widespread and diverse genus of Orthocladiinae. Among the Chironominae the generaPolypedilum, Pseudochironomus, Tanytarsus andRheotanytarsus showed high species richness. Several species were collected that could not be assigned to genus. A number of range extensions were recorded for taxa found in the Neotropical region for the first time and for Neotropical taxa recorded outside of South America for the first time. The Costa Rican chironomid fauna consists of cosmopolitan, holarctic and neotropical components. There is probably an endemic Central American chironomid fauna at the species level.     

Annual wood production in a tropical rain forest in NE Costa Rica linked to climatic variation but not to increasing CO2

Increased atmospheric [CO₂] could theoretically lead to increased forest productivity (‘CO₂ fertilization’). This mechanism was hypothesized as a possible explanation for biomass increases reported from tropical forests in the last 30+ years. We used unique long‐term records of annually measured stands (eighteen 0.5 ha plots, 10 years) and focal tree species (six species, 24 years) to assess the effects of rainfall, temperature, and atmospheric [CO₂] on annual wood production in a neotropical rain forest. Our study area was a meso‐scale section (600 ha) of old‐growth Tropical Wet Forest in NE Costa Rica. Using the repeated remeasurements we directly assessed the relative effects of interannual climatic variation and increasing atmospheric [CO₂] on wood production. A remarkably simple two‐factor model explained 91% of the interannual variance in stand‐level tree growth; the statistically independent factors were total dry season rainfall (positive effect, r²=0.85) and night‐time temperature (negative effect, r²=0.42). Stand‐level tree mortality increased significantly with night‐time temperature. After accounting for dry season rainfall and night‐time temperature, there was no effect of annual [CO₂] on tree growth in either the stand or focal species data. Tree growth in this Tropical Wet Forest was surprisingly sensitive to the current range of dry season conditions and to variations in mean annual night‐time temperature of 1–2°. Our results suggest that wood production in the lowland rainforests of NE Costa Rica (and by extension in other tropical regions) may be severely reduced in future climates that are only slightly drier and/or warmer.