Rainforest Understory Research Articles

Low light availability affects leaf gas exchange, growth and survival of Euterpe edulis seedlings transplanted into the understory of an anthropic tropical rainforest

Euterpe edulis Mart. (Arecaceae) is a threatened palm tree of the Brazilian Atlantic Rainforest understory with fundamental importance for the restoration of degraded forest environments. We assessed the leaf gas exchange, growth and survival of E. edulis seedlings transplanted at three different forest sites (S1, S2 and S3) in the same area in which cocoa trees had been cultivated in a rustic agroforestry system. Measurement was carried out during the first year after seedling transplantation. The sites were characterised according to canopy openness (CO) and total daily photosynthetic photon flux density (PPFD). Average CO and PPFD values were 13.3%, 8.0% and 6.7%, and 3.34, 2.79 and 0.62 mol m−2 d−1 for S1, S2 and S3, respectively. A progressive decline in seedling survival was observed in all sites throughout the experiment. At 387 d after planting, survival at S1, S2 and S3 was 57%, 44% and 37%, respectively. The gross light-saturated photosynthetic rate (Amax), leaf area and plant biomass were significantly higher (P < 0.05) in S1 and S2 when compared with S3. The values of dark respiration rate (R d) and photosynthetic compensation irradiance (I c) were sufficiently low for a positive carbon balance. Notwithstanding, the interpretation of results of microclimate variables together with leaf gas exchange and growth variables indicated that seedlings at all sites were in a suboptimal condition to achieve Amax, which is probably the main cause of the dramatic decline in the seedlings’ survival throughout the first year after transplantation. From a practical point of view, if the values of CO and PFD are lower than 10% and 3 mol m−2 d−1, respectively, it is suggested that the transplanting of E. edulis seedlings to the understory of abandoned agroforestry systems be accompanied by cultural practices, such as the thinning and pruning of tree tops.

Like a jungle sometimes: how leaves survive in the rainforest understory

Like a jungle sometimes: how leaves survive in the rainforest understory

Two new species of &lt;I&gt;Leandra&lt;/I&gt; s. str. (&lt;I&gt;Melastomataceae&lt;/I&gt;) from the Atlantic Forest in Espírito Santo, Brazil

Two species of Leandra that occur in the Atlantic Forest, in the state of Espirito Santo, eastern Brazil, are described and illustrated here. Leandra cristata has been found in the understory of montane rain forest, and can be recognized by the distinct nodal ridges on the young branches, by the leaves with decurrent bases and transversal nerves consistently perpendicular to the main nerve, by the triangular external teeth, and by the dorsal bump on the stamen connective. Leandra fontanae has been found in shrubby vegetation on inselbergs, and can be recognized by the small and cordate leaves with five main nerves plus a faint submarginal pair, by the apical and lateral inflorescences made up by single dichasia or a triad, by the flowers mostly 4-merous and by the 2-celled ovary.

Ottotettix, a New Katydid Genus and Species from the Rainforest of Southern Ecuador (Orthoptera, Tettigoniidae, Pseudophyllinae, Eucocconotini)

Ottotettix smaragdopoda gen. et sp. nov., a brachypterous katydid from the eastern Andean foothills of southern Ecuador, is described. It is moderately common in rainforest understory along creeks and in vales between 850 and 1300 m. The male's ultrasonic calling song, which is also described, is apparently performed only sporadically, and males seem also to use substrate-borne signals.

Ocellar adaptations for dim light vision in a nocturnal bee

Growing evidence indicates that insect ocelli are strongly adapted to meet the specific functional requirements in the environment in which that insect lives. We investigated how the ocelli of the nocturnal bee Megalopta genalis are adapted to life in the dim understory of a tropical rainforest. Using a combination of light microscopy and three-dimensional reconstruction, we found that the retinae contain bar-shaped rhabdoms loosely arranged in a radial pattern around multi-layered lenses, and that both lenses and retinae form complex non-spherical shapes reminiscent of those described in other ocelli. Intracellular electrophysiology revealed that the photoreceptors have high absolute sensitivity, but that the threshold location varied widely between 10(9) and 10(11) photons cm(-2) s(-1). Higher sensitivity and greater visual reliability may be obtained at the expense of temporal resolution: the corner frequencies of dark-adapted ocellar photoreceptors were just 4-11 Hz. Spectral sensitivity profiles consistently peaked at 500 nm. Unlike the ocelli of other flying insects, we did not detect UV-sensitive visual pigments in M. genalis, which may be attributable to a scarcity of UV photons under the rainforest canopy at night. In contrast to earlier predictions based on anatomy, the photoreceptors are not sensitive to the e-vector of polarised light. Megalopta genalis ocellar photoreceptors possess a number of unusual properties, including inherently high response variability and the ability to produce spike-like potentials. These properties bear similarities to photoreceptors in the compound eye of the cockroach, and we suggest that the two insects share physiological characteristics optimised for vision in dim light.

Influence of proximity to a geographical range limit on the physiology of a tropical bird

1. Species' geographical ranges can be limited by a variety of biotic and abiotic factors. Physiological challenge in response to unsuitable environmental conditions can establish limits to geographical ranges. 2. We studied the physiology of Song Wrens (Cyphorhinus phaeocephalus) across their geographical range on the isthmus of Panama, an area characterized by a strong rainfall gradient. Wrens are common on the Caribbean slope of the isthmus where annual rainfall is greatest, but wren abundance declines towards the south as annual rainfall declines. Song Wrens are completely absent from the driest third of the isthmus. 3. We searched for the existence of a physiologically induced distribution limit by measuring body condition (an integrative measure of energy balance), hematocrit (% packed red blood cells in a given blood sample), and corticosterone levels (CORT, a steroid hormone that regulates the availability of energy and the endocrine stress response) in males and females. We caught birds by luring them into nets when they responded to playback of conspecific song. 4. Wrens living in drier habitat near the geographical range edge were significantly more likely to have abnormally low hematocrit scores. Baseline CORT levels were negatively associated with rainfall in one of our three best-fit path models, indicating potential energetic challenge in some individuals. Maximum CORT levels during a 60-min period of restraint correlated significantly only with sex, being higher in females. Birds with the poorest body condition lived at the dry end of the gradient. Birds on the wet end of the gradient responded fastest to conspecific song. 5. Environmental conditions vary across geographical ranges and may influence the physiological conditions of organisms, thereby enforcing limits to species' distributions. Highly specialized species, such as birds of the rain forest understory, may be especially susceptible to environmental variation associated with changing climatic conditions.

Comparison of rainforest butterfly assemblages across three biogeographical regions using standardized protocols

Insects, like most other organisms, are more diverse in tropical than in temperate regions, but standardized comparisons of diversity among tropical regions are rare. Disentangling the effects of ecological, evolutionary, and biogeographic factors on community diversity requires standardized protocols and long-term studies. We compared the abundance and diversity of butterflies using standardised ‘Pollard walk’ transect counts in the understory of closed-canopy lowland rainforests in Panama (Barro Colorado Island, BCI), Thailand (Khao Chong, KHC) and Papua New Guinea (Wanang, WAN). We observed 1792, 1797 and 3331 butterflies representing 128, 131 and 134 species during 230, 231 and 120 transects at BCI, KHC and WAN, respectively. When corrected for length and duration of transects, butterfly abundance and species richness were highest at WAN and KHC, respectively. Although high butterfly abundance at WAN did not appear to result from methodological artefacts, the biological meaning of this observation remains obscure. The WAN site appeared as floristically diverse as KHC, but supported lower butterfly diversity. This emphasizes that factors other than plant diversity, such as biogeographic history, may be crucial for explaining butterfly diversity. The KHC butterfly fauna may be unusually species rich because the site is at a biogeographic crossroads between the Indochinese and Sundaland regions. In contrast, WAN is firmly within the Australian biogeographic region and relatively low species numbers may result from island biogeographic processes. The common species at each of the three sites shared several traits: fruit and nectar feeders were equally represented, more than half of common species fed on either epiphytes or lianas as larvae, and their range in wing sizes was similar. These observations suggest that Pollard walks in different tropical rainforests target similar assemblages of common species, and, hence, represent a useful tool for long-term monitoring of rainforest butterfly assemblages.

Reliable detection of predator cues in afferent spike trains of a katydid under high background noise levels

Katydid receivers face the problem of detecting behaviourally relevant predatory cues from echolocating bats in the same frequency domain as their own conspecific mating signals. We therefore tested the hypothesis that katydids are able to detect the presence of insectivorous bats in spike discharges at early stages of nervous processing in the auditory pathway by using the temporal details characteristic for responses to echolocation sequences. Spike activity was recorded from an identified nerve cell (omega neuron) under both laboratory and field conditions. In the laboratory, the preparation was stimulated with sequences of bat calls at different repetition rates typical for the guild of insectivorous bats, in the presence of background noise. The omega cell fired brief high-frequency bursts of action potentials in response to each bat sound pulse. Repetition rates of 18 and 24 Hz of these pulses resulted in a suppression of activity resulting from background noise, thus facilitating the detection of bat calls. The spike activity typical for responses to bat echolocation contrasts to responses to background noise, producing different distributions of inter-spike intervals. This allowed development of a 'neuronal bat detector' algorithm, optimized to detect responses to bats in afferent spike trains. The algorithm was applied to more than 24 hours of outdoor omega-recordings performed either at a rainforest clearing with high bat activity or in rainforest understory, where bat activity was low. In 95% of cases, the algorithm detected a bat reliably, even under high background noise, and correctly rejected responses when an electronic bat detector showed no response.

Structure and composition of the understory treelets in a non-dipterocarp forest of tropical Asia

Species richness, floristic composition, and structure of understory treelets were investigated in three 1-ha plots, distributed in the undisturbed tropical seasonal rain forest of Xishuangbanna, SW China. We investigated all the woody species except lianas in these plots, compared the differences of species diversity of trees and treelets in these plots and the differences of understory structure between this forest and the typical dipterocarp forests in SE Asia. We found 5089 individuals belonging to 356 species, 189 genera, and 63 families in the three plots. The five most important families were Rubiaceae, Euphorbiaceae, Lauraceae, Meliaceae, and Annonacae. The treelet layer was much more diverse than the tree layer, indicating that result based only on trees may be not an appropriate representation of the diversity status of a particular tropical forest type. The three plots were more similar when the treelet layer was considered than when tree layer was considered. In contrast with the dipterocarp forests in SE Asia, the understory of tropical seasonal rain forest was mainly composed of the trees with small and middle stature at maturity classes. These results indicate that the structure of tropical forest understory may differ not only among continents but also within continent.

Diversity and composition of understory vegetation in the tropical seasonal rain forest of Xishuangbanna, SW China

Tropical forests vegetation and community research have tended to focus on the tree component, and limited attention has been paid to understory vegetation. Species diversity and composition of the understory of tropical seasonal rain forest were inventoried in a 625 m2 area (for sapling layer) and a 100 m2 area (for herb/seedling layer) in three 1 ha plots. We found 3068 individuals belonging to 309 species, 192 genera and 89 families. The most important family as determined by the Family Importance Value (FIV) was Rubiaceae in both sapling and herb/seedling layers. In terms of Importance Value Index (IVI), the shrub Mycetia gracilis (Rubiaceae) was the most important species in the sapling layer and the pteridophyte Selaginella delicatula (Selaginellaceae) was the most ecological significant species in the herb/seedling layer. Much more vascular plant species were registered in the understory than in the tree layer totaled among the three plots. The species diversity did not differ significantly among the tree layer, sapling layer and herb/seedling layer. Given that we still know little about the understory plant community for growth forms other than trees, the results from the present study indicate that more attention should be paid to the understory vegetation during the decision-making process for biodiversity conservation in the tropical forests.

Traits, Habitats, and Clades: Identifying Traits of Potential Importance to Environmental Filtering

Environmental filtering is a fundamental process in the ecological assembly of communities. Recently developed phylogenetic tools identify patterns associated with environmental filtering across whole communities. Here we introduce a novel method that allows the detection of traits involved in the environmental filtering of species from specific clades in specific habitat types. Our approach identifies nonindependent trait/habitat/clade (THC) associations and also provides a framework for detecting clearly defined two-way trait/clade, trait/habitat, and clade/habitat associations. The THC method relies on exact binomial tests and differentiates THC associations resulting from a three-way interaction from those that are generated by one or more underlying significant two-way interactions. It can also detect THC associations for which there are no significant two-way associations (trait/habitat, trait/clade, clade/habitat). To illustrate the THC method, we examine plant pollination and dispersal traits from six habitat types in a fragmented Costa Rican landscape. Results suggest that these traits are not widely important for the environmental filtering of most clades in this landscape, but animal dispersal and insect pollination are involved in the filtering of monocots and the Piperaceae in rain forest understory.

Two new species of shrub frogs (Rhacophoridae: Philautus) from the lowlands of Sri Lanka

Two new species of Sri Lankan frogs of the genus Philautus are described. Species diagnoses are based on morphology, morphometrics and mitochondrial DNA sequence data. Philautus tanu sp. nov. inhabits shrubs in open areas of the lowland wet zone, while P. singu sp. nov. is found on shrubs in the understory of lowland and mid-elevation rainforests. These descriptions bring the total number of valid Sri Lankan Philautus to 65 species, of which 46 are extant.

Graveyards on the Move: The Spatio-Temporal Distribution of Dead Ophiocordyceps-Infected Ants

Parasites are likely to play an important role in structuring host populations. Many adaptively manipulate host behaviour, so that the extended phenotypes of these parasites and their distributions in space and time are potentially important ecological variables. The fungus Ophiocordyceps unilateralis, which is pan-tropical in distribution, causes infected worker ants to leave their nest and die under leaves in the understory of tropical rainforests. Working in a forest dynamic plot in Southern Thailand we mapped the occurrence of these dead ants by examining every leaf in 1,360 m2 of primary rainforest. We established that high density aggregations exist (up to 26 dead ants/m2), which we coined graveyards. We further established that graveyards are patchily distributed in a landscape with no or very few O. unilateralis-killed ants. At some, but not all, spatial scales of analysis the density of dead ants correlated with temperature, humidity and vegetation cover. Remarkably, having found 2243 dead ants inside graveyards we only found 2 live ants of the principal host, ant Camponotus leonardi, suggesting that foraging host ants actively avoid graveyards. We discovered that the principal host ant builds nests in high canopy and its trails only occasionally descend to the forest floor where infection occurs. We advance the hypothesis that rare descents may be a function of limited canopy access to tree crowns and that resource profitability of such trees is potentially traded off against the risk of losing workers due to infection when forest floor trails are the only access routes. Our work underscores the need for an integrative approach that recognises multiple facets of parasitism, such as their extended phenotypes.

A New Rare Species of Lobelia (Campanulaceae, Lobelioideae) from Mexican Lowland Rainforest

Lobelia lithophila Senterre & Castillo-Campos (Campanulaceae, Lobelioideae) is described as a new species from the lowland tropical evergreen rainforest of the region of Uxpanapa, in southern Veracruz, Mexico. This species is only known from the type locality and probably represents a microendemic. The species described is most closely related to L. orientalis Rzedowski & Calderón, another local endemic from Querétaro, Mexico. Both taxa combine the herbaceous life form with leaves more or less in a rosette at ground level and a completely superior ovary, and are from specialized habitats. The new species is easily distinguished by its longer leaf blade and smaller petiole, and by its reduced leaf pubescence. Its habitat is also unusual, as it is restricted to limestone rocks in the understory of lowland tropical rainforest. Lobelia lithophila belongs to subsection Leiospermae E. Wimmer within the section Lobelia, subgenus Lobelia L.

Inter- and Intraspecific Comparisons of Antiherbivore Defenses in Three Species of Rainforest Understory Shrubs

Plants defend themselves against herbivores and pathogens with a suite of morphological, phenological, biochemical, and biotic defenses, each of which is presumably costly. The best studied are allocation costs that involve trade-offs in investment of resources to defense versus other plant functions. Decreases in growth or reproductive effort are the costs most often associated with antiherbivore defenses, but trade-offs among different defenses may also occur within a single plant species. We examined trade-offs among defenses in closely related tropical rain forest shrubs (Piper cenocladum, P. imperiale, and P. melanocladum) that possess different combinations of three types of defense: ant mutualists, secondary compounds, and leaf toughness. We also examined the effectiveness of different defenses and suites of defenses against the most abundant generalist and specialist Piper herbivores. For all species examined, leaf toughness was the most effective defense, with the toughest species, P. melanocladum, receiving the lowest incidence of total herbivory, and the least tough species, P. imperiale, receiving the highest incidence. Although variation in toughness within each species was substantial, there were no intraspecific relationships between toughness and herbivory. In other Piper studies, chemical and biotic defenses had strong intraspecific negative correlations with herbivory. A wide variety of defensive mechanisms was quantified in the three Piper species studied, ranging from low concentrations of chemical defenses in P. imperiale to a complex suite of defenses in P. cenocladum that includes ant mutualists, secondary metabolites, and moderate toughness. Ecological costs were evident for the array of defensive mechanisms within these Piper species, and the differences in defensive strategies among species may represent evolutionary trade-offs between costly defenses.

Graminofolium Nickle: a new genus of katydid with two species from northern South America

Abstract Two new species of a new genus of katydids collected in the understory of rainforests of northern Peru are described and figured. Observations on one species suggests that both may be predaceous or, at least, opportunistic predators. Placement of this genus into either Copiphorinae or Agraeciinae has not been resolved and demonstrates the need for studies to refine subfamilial and tribal relationships among the tettigoniids.

Determinants of Lichen Diversity in a Rain Forest Understory

ABSTRACTChange in lichen diversity is often used as a bioindicator to estimate effects of atmospheric pollution, but natural variation in lichen cover and species richness can be very high. We examined the top‐down effects of spore‐consuming ants and the bottom‐up effects of nutrient and light availability on lichen diversity associated with the leaf surface of the rain forest understory plant, Piper cenocladum. Plots containing P. cenocladum were randomly assigned to treatments in factorial experiments that included high and low light levels, nutrient enrichment, and presence and absence of the ant mutualist, Pheidole bicornis. At the conclusion of the experiments, plants were harvested and size of leaves, secondary metabolite content (amides), epiphyll cover, and the species richness of the lichens (which comprised 85% of the epiphyll community) were quantified. Epiphyll cover (mosses, liverworts, and lichens) was greater on plants that had ant‐mutualists and balanced resources. Lichen species richness was greater for plants with balanced resources, particularly for those with high light availability. Relationships between toxins and lichen cover and richness were weak and unclear. In this system, natural sources of variation were reliable determinants of lichen diversity and both biotic and abiotic influences were important.

Edge Effects of Linear Canopy Openings on Tropical Rain Forest Understory Microclimate

ABSTRACTWe investigated microclimatic edge gradients associated with grassy powerlines, paved highways and perennial creeks in wet tropical forest in northeastern Australia during wet and dry seasons. Photosynthetically active radiation, air temperature and vapor pressure deficit, soil temperature, canopy temperature, soil moisture, and air speed in the rain forest understory were measured during traverses perpendicular to the forest edge. Light intensity was elevated near the edges of powerlines, highways, and creeks, but this effect was strongest for creek edges. Air temperature and vapor pressure deficit were elevated near powerline edges in the dry season and highway edges in both wet and dry seasons but were not elevated near creek edges in either season. In contrast, soil moisture was lowered near creek edges but not near either powerline or highway edges. No edge gradients were detected for air speed. Canopy temperature was elevated near highway edges and lowered near powerline edges in the wet season but no edge gradients in canopy temperature were detected near creek edges in either the wet or the dry season. We suggest that these different edge gradients may be largely the result of differences in the fluxes of latent and sensible heat within each type of linear canopy opening, with periodic flood disturbance assisting by maintaining a more open canopy near creek edges. Our data indicate that the nature of the linear canopy opening is at least as important as the width in determining the nature and severity of microclimatic edge effects, analogous to the “matrix effect” of traditional fragmentation studies.

Functional characteristics of corticolous lichens in the understory of a tropical lowland rain forest

In tropical lowland forests, corticolous crustose green algal lichens are abundant and highly diverse. This may be related to adaptation to prevailing microenvironmental conditions including, for example, high precipitation and low light intensities. In the understory of a tropical lowland rain forest in French Guiana, we studied the morphology of crustose green algal lichens and measured gas exchange and chlorophyll a fluorescence. We found that (i) periods of thallus suprasaturation with water were reduced by the presence of water-repelling surface structures of mycobiont hyphae at the thallus surface and the medulla; (ii) photosynthesis was adapted to the low light intensities present in the understory; and (iii) photosynthesis was rapidly activated in fluctuating light. The combination of these three mechanisms enables corticolous lichens to implement specific morphological and physiological strategies, which may favour growth in the limiting understory habitat of tropical lowland rain forests.

The impact of Amomum villosum cultivation on seasonal rainforest in Xishuangbanna, Southwest China

Integrating conservation goals with the sustainable use of tropical rain forests has received much attention in recent decades. Amomum villosum, a traditional Chinese medicinal herb, has been cultivated in the understory of seasonal rain forest for 40 years in Xishuangbanna, Southwest China. Cultivated area has reached 58.11 km2 much of which is distributed within nature reserves. This practice has caused controversy on whether A. villosum cultivation would alter the structure and function of the primary forest. This study examined the effects of medicinal plant cultivation on seasonal rain forest by comparing plant diversity, biomass, litterfall and soil nutrients of primary rain forest with disturbed areas where A. villosum is cultivated. The results indicate that plant diversity, tree biomass, litter production and soil nutrients are significantly lower in the disturbed than in the primary rain forest. These results suggest that the cultivation of A. villosum affects the structure and function of the seasonal rain forest in Xishuangbanna.