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.     

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.     

Disadvantages of living in a populous neighborhood for sit-and-wait predators: Competition for space reduces pit-trap size in antlion larvae

The study of how trap design responds to biotic and abiotic conditions can help to understand the selective forces affecting the foraging of trap-building organisms. We experimentally tested whether pit design can be modified by intraspecific competition for space in larvae of Myrmeleon crudelis, a common sit-and-wait predator that digs conical pit traps in the soil to capture walking arthropods. In a tropical forest in Costa Rica, we measured pit dimensions, larval body size, and the level of competition (i.e., density of neighboring traps) in 40 antlion larvae. These larvae were then taken to the laboratory and allowed to build new traps in individual containers. We measured within-individual changes in the size of traps in the field and in the laboratory, and related these to the level of competition experienced in the field. Larvae with relatively high levels of competition in the field showed a greater increase in the size of their pits in the laboratory. This change was independent of larval size. Larvae with none or few neighbors in the field showed little change in their pit sizes, whereas those with higher competition levels increased their diameter and depth up to 1,400% and 1,000%, respectively. Our results demonstrate that, at least in high-density aggregations, pit design is restricted by competition in addition to the constraints imposed by body size. This work suggests that biotic interactions can play a role in the design of extended phenotypes in sit-and-wait predators that live in dense aggregations.     

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

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

Tropical nematode diversity: vertical stratification of nematode communities in a Costa Rican humid lowland rainforest

Comparisons of nematode communities among ecosystems have indicated that, unlike many organisms, nematode communities have less diversity in the tropics than in temperate ecosystems. There are, however, few studies of tropical nematode diversity on which to base conclusions of global patterns of diversity. This study reports an attempt to estimate nematode diversity in the lowland tropical rainforest of La Selva Biological Research Station in Costa Rica. We suggest one reason that previous estimates of tropical nematode diversity were low is because habitats above the mineral soil are seldom sampled. As much as 62% of the overall genetic diversity, measured by an 18S ribosomal barcode, existed in litter and understorey habitats and not in soil. A maximum-likelihood tree of barcodes from 360 individual nematodes indicated most major terrestrial nematode lineages were represented in the samples. Estimated 'species' richness ranged from 464 to 502 within the four 40 × 40 m plots. Directed sampling of insects and their associated nematodes produced a second set of barcodes that were not recovered by habitat sampling, yet may constitute a major class of tropical nematode diversity. While the generation of novel nematode barcodes proved relatively easy, their identity remains obscure due to deficiencies in existing taxonomic databases. Specimens of Criconematina, a monophyletic group of soil-dwelling plant-parasitic nematodes were examined in detail to assess the steps necessary for associating barcodes with nominal species. Our results highlight the difficulties associated with studying poorly understood organisms in an understudied ecosystem using a destructive (i.e. barcode) sampling method.     

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.     

Host plant scents attract rolled-leaf beetles to Neotropical gingers in a Central American tropical rain forest

Leaf volatile chemicals are known to reduce herbivory rates by repelling or intoxicating insect herbivores and by attracting the predators and parasitoids of herbivores. However, leaf volatiles may also be used by insect herbivores as cues to locate their host plants. Leaf volatiles are suggested to be important host search cues for herbivores in structurally complex and diverse habitats, such as tropical rain forests. A group of insect herbivores, the rolled-leaf beetles (Coleoptera: Chrysomelidae: Hispinae), have maintained a highly specialized interaction with Neotropical gingers (Zingiberales) for ca. 60 million years. In this study, we explored chemical attraction to host plants under controlled laboratory conditions, using four sympatric rolled-leaf beetle species, Cephaloleia dorsalis Baly, Cephaloleia erichsonii Baly, Cephaloleia fenestrata Weise, and Cephaloleia placida Baly. For each beetle species, we investigated (i) whether it was repelled or attracted by leaf scents produced by four host and four non-host plant species, including Neotropical gingers in the families Marantaceae, Costaceae, and Zingiberaceae; and (ii) its ability to use scents to detect its host plant. We found that rolled-leaf beetles can detect and are attracted by leaf volatiles from both host and non-host gingers. Additionally, when beetles were simultaneously exposed to leaf volatiles from host and non-host plants, three rolled-leaf beetle species were significantly more attracted by volatiles from their host plants than from non-hosts. Only one of the beetle species was not able to discriminate between host and non-host scents.     

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.     

Patterns in a species‐rich tropical understory plant community

Understory plants are an important component of the high plant species diversity characteristic of neotropical rain forests. Herbs, shrubs, understory trees, and saplings of canopy trees occupy a broadly uniform environment of abundant rainfall, low light levels, and high humidity. We asked whether this community at the La Selva Biological Station in the Caribbean lowlands of Costa Rica was structured by environmental filters such as soil origin, topographic position, and understory light availability. We used nested quadrats to assess effects of soil origin (recent alluvium, weathered alluvium, residual volcanic soil) and topographic position (ridges, mid‐slopes and flats) on species composition, density, and diversity and measured six edaphic and understory light parameters. Non‐metric multidimensional scaling ordinations were based on frequency of occurrence in 20 quadrats for 272 species in the shrub size class and 136 species in the small‐tree size class for 17 sites. Three axes were correlated with composite environmental variables produced by principal component analysis representing slope, extractable phosphorus, and light. NMS site positions also reflected soil origin, topographic position, and geographic location. The analyses illustrated a complex community structured by species responses to environmental filters at multiple, interdigitated spatial scales. We suggest that light availability affected by canopy dynamics and dispersal limitation provides additional sources of variation in species distributions, which interact with edaphic patterns in complex ways. Abstract in Spanish is available with online material.     

Diversity and composition of tropical soil nitrifiers across a plant diversity gradient and among land-use types

Ammonia‐oxidizing bacteria (AOB) perform the rate‐limiting step of nitrification, a key ecosystem process that in part determines the fate of nitrogen in ecosystems. However, little is known about the factors that determine soil AOB diversity or composition, especially in tropical systems. Using a set of study systems in Costa Rica, we examined whether plant diversity or land‐use influenced AOB diversity or composition and whether AOB diversity or composition were associated with nitrification rates. We characterized the molecular diversity and composition of AOB via polymerase chain reaction amplification, cloning, and sequencing of 16S rDNA. We found that AOB diversity or composition did not change significantly across plant diversity treatments. In contrast, AOB differed among land‐use types in some measures of diversity and in composition, and differences in AOB composition among land‐use types were correlated with potential rates of nitrification. Our results suggest that anthropogenic changes of ecosystems can alter microbial communities in ways that may affect the processes they mediate.     

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     

Interspecific Territoriality by a Rufous-Tailed Hummingbird (Amazilia tzacatl): Effects of Intruder Size and Resource Value1

I studied two aspects of interspecific territoriality in a Costa Rican nectarivorous bird, the rufous-tailed hummingbird (Amazilia tzacatl). First, I examined variation in the response of the territory holder to four species of intruding hummingbirds by quantifying the proportion of intruders chased from the territory. This measure of territory defense varied significantly among species of intruders and was negatively related to the intruding species’body mass, possibly due to potential costs associated with becoming involved in escalated contests with larger individuals. Second, I tested for an effect of resource manipulation on territory defense. I increased the resource value of the territory by injecting extant natural flowers with artificial sucrose solution. While the frequency of territorial intrusions did not change, the proportion of intruders chased by the territory holder increased. Apparently, nectar supplementation changed the territory holder's perception of resource value but had little impact on the intruders’perception.     

Do Collared Peccaries Negatively Impact Understory Insectivorous Rain Forest Birds Indirectly Via Lianas and Vines?

Tropical rain forest understory insectivorous birds are declining, even in large forest reserves, yet the mechanisms remain unclear. Abundant large mammals can reshape forest structure, which degrades foraging microhabitat. We used six sites in Nicaragua, Costa Rica, and Panama with varying collared peccary (Pecari tajacu) density to test three linked hypotheses: (1) locally declining understory insectivores forage preferentially in liana tangles; (2) vine and liana density, cover, and frequency of dense tangles are lower in the presence of abundant collared peccaries; and consequently (3) abundant collared peccaries are associated with reduced understory insectivorous bird abundance. Three insectivores that declined at La Selva preferentially foraged in liana tangles: Checker‐throated Antwren (Epinecrophylla fulviventris), Dot‐winged Antwren (Microrhopias quixensis), and Ruddy‐tailed Flycatcher (Terenotriccus erythrurus). Vine density, liana cover, liana tangle frequency, and forest cover were lower in the presence of collared peccaries relative to experimental mammal exclosures, with the greatest differences at La Selva Biological Station, Costa Rica. Across sites, five of seven vine and liana measures showed negative, curvilinear relationships with peccary densities. Vine and liana measures peaked at sites with intermediate peccary density, and were low at La Selva. Structural equation models suggest negative indirect effects of the collared peccaries on focal bird densities, mediated by vine and liana density, cover, or tangle frequency. Forest area and rainfall affected both lianas and birds, but collared peccaries also contributed to the reduced abundance of understory insectivores. Indirect effects such as that suggested here may occur even in large, protected forest reserves where large mammal communities are changing.