The role of resource imbalances in the evolutionary ecology of tropical arboreal ants

In numbers and biomass, ants (Hymenoptera, Formicidae) often dominate arthropod faunas of tropical rainforest canopies. Extraordinary ant abundance is due principally to one or a few species able to tap the high productivity of canopy foliage by feeding on plant and homopteran exudates. Prior studies of nitrogen isotopic ratios show that exudate-feeders derive much of their nitrogen (N) by processing large quantities of N-poor, but carbohydrate (CHO)-rich, exudates. CHOs in excess of those that can be coupled with protein for growth and reproduction (postulated as the colony's first priorities) may be directed at little cost and some profit to functions that increase access to limiting protein. High dietary CHO:protein ratios for exudate-feeders appear to subsidize 'high tempo' foraging activity, defence of absolute (level III) territories, and production of N-free alarm/defence exocrine products that enhance ecological dominance in contests with other ants. Among organisms (e.g. plants and Lepidoptera) symbiotic with ants, CHO:protein ratios of ant rewards may control both the identities of ant associates and the quality of ant-rendered services. Dietary ratios of CHO:protein play an important and previously unrecognized role in the ecology and evolution of ants generally. Modifications of worker digestive systems in certain ant subfamilies and genera represent key innovations for handling and processing large volumes of liquid food. The supreme tropical dominants are species released from nest site limitation and able to place their nests in the vicinity of abundant exudate resources. Polydomy appears to be typical of these species and should produce energetic savings by taking colony fragments to the resource.     

A metacommmunity approach to co-occurrence patterns and the core-satellite hypothesis in a community of tropical arboreal ants

Inferring mechanisms of community assembly from co-occurrence patterns is difficult in systems where assembly processes occur at multiple spatial scales and among species with heterogeneous dispersal abilities. Here, we demonstrate that local scale analysis of co-occurrence patterns is inadequate to fully describe assembly mechanisms and instead utilize a metacommunity and core-satellite approach. We generated a co-occurrence and life-history data set for a community of twig-nesting ants on coffee plants across 36 sites within a tropical agroecosystem to test the following three hypotheses: (1) twig-nesting ant species compete for nest-sites, (2) they are structured as a metacommunity, and (3) core species show segregated patterns, while satellite species show random patterns of co-occurrence. Species were divided into four groups: core species that are well distributed regionally and dominant locally, regional dominants that are well distributed regionally but do not dominate locally, local dominants that are dominant locally but are not widely distributed, and satellites that are neither widely distributed nor dominant locally. Only the most abundant species in the community, Pseudomyrmex simplex, was classified as a core species. Regional dominants, local dominants, and satellite species show random patterns of co-occurrence. However, when P. simplex is included in the co-occurrence matrix, patterns become aggregated for all three species groupings. This suggests that P. simplex “assembles” the community by providing a core metapopulation that other species track. Analyzing co-occurrence patterns among candidate subsets of species, at multiple spatial scales, and linking them to species traits substantially improves the explanatory power of co-occurrence analyses in complex metacommunities.     

Novel interactions of non-pollinating ants with pollinators and fruit consumers in a tropical forest

The tropical ants Ectatomma ruidum and E. tuberculatum (Formicidae) regularly patrol leaves, flowers, and fruits of the understory shrub, Psychotria limonensis (Rubiaceae), on Barro Colorado Island, Panama. Ant and pollinator exclusion experiments elucidated both positive and negative effects of ant attendance on plant reproductive success, including pollination, fruit set, fruit loss, and fruit removal. Ants did not pollinate flowers but did contribute to higher pollination success, probably by increasing the relocation frequency of winged pollinators and thus the rate of flower visitation. Ants also prevented fruit loss to herbivorous insects which were common during the early stages of fruit development. Thus, ant attendance strongly improved both pollination and fruit set whereby plants with ants set more fruit per flower and also lost fewer fruits during fruit maturation. In contrast, ants had a negative effect on the removal of ripe fruits by avian frugivores. Thus, ant attendance has a non-trivial influence on plant reproduction, this interaction being beneficial at some stages of the plant reproductive cycle and carrying costs at another stage. A tight ecological or co-evolved relationship between these Ectatomma spp. and P. limonensis is unlikely given that ant attendance of plants is detrimental to fruit removal. Received: 18 May 1998 / Accepted: 1 March 1999     

The decomposition of mangrove litter in a subtropical mangrove forest

Decomposition of Avicennia marina (Forsk.) Vierh. leaf and woody litter (twigs) was studied using litter bag experiments in a subtropical mangrove forest at two tidal levels (high and low) with different inundation regimes and during two seasons (summer and winter). Losses in dry weight were best described by a single exponential model which showed loss rates of both leaves and twigs were significantly higher low down on the shoreline (greater inundation) and in summer. The time (days) required for the loss of half of the initial dry weight (t ₅₀) was summer: high 59, low 44; winter: high 98, low 78. For twigs the values (days) were summer: high 383, low 179; winter; high 1327, low 1207. There is an exponential relationship between leaf litter t ₅₀'s and latitude which indicates the importance of temperature and therefore season, to the dynamics of organic cycling and export in mangrove systems.     

Behavioral diversity of predatory arboreal ants in coffee agroecosystems

Aspects of predator assemblages that alter predator effects on prey have received extensive recent attention. Among other mechanisms, differences in behavior or resource use within predator trophic levels may enhance predator effects on prey, especially if effects of each predator species differ with environmental conditions. We address whether three common ant species (Azteca instabilis F. Smith, Camponotus textor Forel, and Crematogaster spp.) are functionally unique in coffee agroecosystems, asking if each species differs in (1) cooperative foraging behavior, (2) responses to experimentally introduced herbivores, and (3) responses to pest outbreaks. Furthermore, we examined the behaviors and effects of each ant species under different conditions by varying herbivore species, herbivore size, and herbivore density and carrying out observations in different seasons. Ant species significantly differed in foraging behaviors, in effects on individual herbivores, and in responses to pest outbreaks in terms of both type and time of response to herbivores. The behaviors and effects of each ant species differed in the dry and wet seasons and for different herbivore species and sizes. Although A. instabilis generally removed more larvae and more quickly removed larvae, this was not the case under all conditions. The data presented thus support that common ant species in coffee agroecosystems are behaviorally diverse in their responses to herbivores under different conditions. We discuss the implications of these differences in ant behaviors for enhancement of predatory function in light of both multipredator effects and in terms of the potential importance of predator diversity.     

Assessing environmental requirements effects on forest fragmentation sensitivity in two arboreal rodents

The study of the effect of habitat fragmentation on species that inhabit residual patches requires the investigation of the relationship existing between species distribution and landscape components. To understand which components of landscape mosaics are more influential for species’ persistence, we compared the distribution of two arboreal rodents proved to be sensitive to habitat fragmentation, the hazel dormouse Muscardinus avellanarius and the red squirrel Sciurus vulgaris. Their occurrence in residual oak woods in central Italy was studied with nest-boxes and hair-tubes, developing a new method for hair analysis. Their distributions were analysed considering patch, matrix composition and configuration, and landscape vegetation variables. The two species showed a different degree of plasticity, with the squirrel being significantly more specialised at the landscape scale. The comparison of the two distribution patterns highlighted the influence of different ecological constraints and the existence of different strategies to cope with fragmentation. Patch size and patch attributes were generally weaker determinants of occurrence, compared to landscape metrics. The squirrel presence was significantly influenced by the presence of shared perimeter between hedgerows and woods and by the lack of isolation of the residual patches, suggesting the use of several fragments to compensate the low habitat quality. Conversely the hazel dormouse seemed to be more affected by the internal management of the woods, and in particular by the mean DBH. Our results highlight how the recognition of the extrinsic constraints and the influence of multi-scale habitat selection may help guiding land use management, to ensure species conservation in profoundly exploited landscapes.     

Edge effects in a tropical montane forest mosaic: experimental tests of post-dispersal acorn removal

Forest edges created by scattered-patch clear-cutting have become common in tropical montane cloud forests in the highlands of Chiapas, Mexico. It was hypothesised that forest edges may influence regeneration of oak species, which are canopy dominants in these forests, by affecting the activities of small mammal species. Acorns of different oak species varying in germination timing were offered to predators and/or dispersers at different positions along replicated forest edges during 2 consecutive years. We investigated the effects of (1) edge type (hard and soft), (2) distance from the edge (0, 15, 30, 45 and 60 m inside forest fragments) and (3) oak species, on the rate of acorn removal mainly by small mammals. During a non-masting year, acorn removal was affected by the interaction of edge type and distance from the edge (P<0.05), with acorn removal being highest near hard edges compared to adjacent forest interiors. As predicted, acorn removal was greater along soft (100%) than along hard edges (82%), but this pattern was recorded only during the non-masting year. This study partly supports previous studies of rodents preferentially consuming acorns with early germination rather than acorns exhibiting dormancy, however these patterns may change with variation in acorn abundance. These results suggest that patch clear-cutting affects regeneration processes within forest fragments by influencing the activities of small mammals, but the nature of this effect also depends on acorn abundance and the characteristics of the forest edge created.     

Effects of selective logging on the arboreal ants of a Bornean rainforest

We examined the effect of selective logging and corresponding forest canopy loss on arboreal ant diversity in a tropical rainforest. Arboreal ants were collected from an unlogged forest plot and from forest plots selectively logged 14 years and 24 years earlier in Danum Valley, Sabah, Malaysia, using a canopy fogging method. Selective logging was associated with a significant decrease in canopy cover and an increase in understory vegetation density relative to unlogged forest. Our study showed that selective logging in primary forest might not dramatically decrease total species number and overall abundance of arboreal ants; however, it may influence the species composition and dominance structure of the ant community, accompanied by an increase in abundance of shrub‐layer species and trophobiotic species. In view of the results of this study, management techniques that minimize logging impact on understory vegetation structure are likely to help maintain the conservation value of logged forests for arboreal ants. Our results also suggest that accurate assessment of the impacts on biodiversity should not be based only on measurement of species number and overall abundance, but also on analysis of species composition and community structure.     

Evolution and ecology of directed aerial descent in arboreal ants

Directed aerial descent (DAD) is used by a variety of arboreal animals to escape predators, to remain in the canopy, and to access resources. Here, we build upon the discovery of DAD in ants of tropical canopies by summarizing its known phylogenetic distribution among ant genera, and within both the subfamily Pseudomyrmecinae and the genus Cephalotes. DAD has multiple evolutionary origins in ants, occurring independently in numerous genera in the subfamilies Myrmicinae, Formicinae, and Pseudomyrmecinae. Ablation experiments and video recordings of ants in a vertical wind tunnel showed that DAD in Cephalotes atratus is achieved via postural changes, specifically orientation of the legs and gaster. The occurrence of DAD in Formicinae indicates that the presence of a postpetiole is not essential for the behavior. Evidence to date indicates that gliding behavior is accomplished by visual targeting mediated by the compound eyes, and is restricted to diurnally active ants that nest in trees. Occlusion of ocelli in Pseudomyrmex gracilis workers had no effect on their success or performance in gliding. Experimental assessment of the fate of ants that fall to the understory showed that ants landing in water are 15 times more likely to suffer lethal attacks than are ants landing in leaf litter. Variation in both the aerodynamic mechanisms and selective advantages of DAD merits further study given the broad taxonomic diversity of arboreal ants that engage in this intriguing form of flight.     

Seasonal dynamics of arboreal spider diversity in a temperate forest

Measuring and estimating biodiversity patterns is a fundamental task of the scientist working to support conservation and inform management decisions. Most biodiversity studies in temperate regions were often carried out over a very short period of time (e.g., a single season) and it is often-at least tacitly-assumed that these short-term findings are representative of long-term general patterns. However, should the studied biodiversity pattern in fact contain significant temporal dynamics, perhaps leading to contradictory conclusions. Here, we studied the seasonal diversity dynamics of arboreal spider communities dwelling in 216 European beeches (Fagus sylvatica L.) to assess the spider community composition in the following seasons: two cold seasons (I: November 2005-January 2006; II: February-April) and two warm seasons (III: May-July; IV: August-October). We show that the usually measured diversity of the warm season community (IV: 58 estimated species) alone did not deliver a reliable image of the overall diversity present in these trees, and therefore, we recommend it should not be used for sampling protocols aimed at providing a full picture of a forest's biodiversity in the temperate zones. In particular, when the additional samplings of other seasons (I, II, III) were included, the estimated species richness nearly doubled (108). Community I possessed the lowest diversity and evenness due to the harsh winter conditions: this community was comprised of one dominant species together with several species low in abundance. Similarity was lowest (38.6%) between seasonal communities I and III, indicating a significant species turnover due to recolonization, so that community III had the highest diversity. Finally, using nonparametric estimators, we found that further sampling in late winter (February-April) is most needed to complete our inventory. Our study clearly demonstrates that seasonal dynamics of communities should be taken into account when studying biodiversity patterns of spiders, and probably forest arthropods in general.     

Photosynthetic performance and resource utilization of two mangrove species coexisting in a hypersaline scrub forest

In a hypersaline mangrove scrub forest in northern Florida, coexisting trees of Laguncularia racemosa and Avicennia germinans were either fertilized with nitrogen or phosphorus, or not fertilized (controls). We aimed to test whether nutrient additions differentially altered photosynthetic performance and resource utilization in these two species. In control trees, photosynthetic rates were higher in L. racemosa than A. germinans. However, leaf nitrogen concentrations were higher in A. germinans than L. racemosa. Avicennia germinans responded to fertilization with nitrogen by increasing leaf nitrogen concentrations and rates of photosynthesis such that they were equivalent to photosynthesis in L. racemosa. Laguncularia racemosa did not show a response to nitrogen additions. Neither species showed strong responses to phosphorus fertilization. Avicennia germinans had high photosynthetic water-use efficiency (photosynthesis/transpiration), but low photosynthetic nitrogen-use efficiency (photosynthesis/leaf nitrogen). In contrast, L. racemosa had comparatively low photosynthetic water use efficiency and high photosynthetic nitrogen use efficiency. Leaf level characteristics lead us to hypothesize that coexistence of A. germinans and L. racemosa should occur where nitrogen levels are low and salinity is moderate, or at least moderate for some period of the year.     

The importance of crevices to the intertidal snail Littoraria articulata (Philippi) in a tropical mangrove forest

The intertidal gastropod, Littoraria articulata (Philippi), is common on the trunks of mangroves at Coorooman Creek, Central Queensland. Individuals of shell length less than 3 mm were only found in empty tests of the barnacle Hexaminius popeiana (Foster). Larger snails were exposed on the trunk or occasionally in crevices such as knot holes. The relationship between shell length and shell strength differed between barnacle dwelling and larger L. articulata: in both cases it was linear, but the rate of increase of strength with increasing length was significantly greater for barnacle dwelling snails than for exposed ones. In contrast, there was an exponential relationship between shell strength and length for the weaker shelled Littoraria filosa (Sowerby), which occurs higher on the tree and does not inhabit barnacles. The survival of L. articulata was affected by the presence of crevices: significantly fewer disappeared from posts with artificial crivices than from those which lacked them, and some snails testhered to posts appeared to have been eaten by fish. The importance of crevices is discussed in relation to predation pressure, growth and habitat selection by larval L. articulata. This revised version was published online in July 2006 with corrections to the Cover Date.     

The shared preference niche of sympatric Asiatic black bears and sun bears in a tropical forest mosaic

Background

Ecologically similar species often coexist by partitioning use of habitats or resources. Such partitioning can occur through divergent or shared niches. We investigated overlap in habitat use and spatial co-occurrence by sympatric Asiatic black bears and sun bears in three habitats in Thailand, and thereby assessed which niche model best accounts for their coexistence.

Methods/Principal Findings

We used density of species-specific signs to assess habitat use. Signs of both bear species occurred in all three habitats, and on >60% of sampling transects. Both species fed mostly on fruit; insect feeding signs were uncommon, and were mostly from sun bears. Significant differences in habitat use occurred only in montane forest, the habitat in which fruit was most abundant; incidence of black bear sign there was six times higher than that of sun bears. Habitat use was similar between the two species in the other habitats, which comprised 85% of the area. Of 10 habitat attributes examined, fruiting tree density was the best predictor of occurrence for both species. Models that included interspecific competition (fresh foraging activity of the other species) were less supported than the top models without competition.

Conclusions/Significance

Bear species co-occurrence at both coarse and fine spatial scales and use of the same resources (fruit trees) indicated common niche preferences. However, their habitat use differed in ways expected from their physical differences: larger black bears dominated in the most fruit-rich habitat, and smaller sun bears used less-preferred insects. These results indicate broadly overlapping fundamental niches combined with asymmetric competition—features consistent with the concept of shared preference niches. This model of the niche has received little attention in ecology, but appears to be relatively common in nature.     

Cyanobacterial diversity in the phyllosphere of a mangrove forest

The cyanobacterial community colonizing phyllosphere in a well-preserved Brazilian mangrove ecosystem was assessed using cultivation-independent molecular approaches. Leaves of trees that occupy this environment (Rhizophora mangle,Avicennia schaueriana and Laguncularia racemosa) were collected along a transect beginning at the margin of the bay and extending upland. The results demonstrated that the phyllosphere of R. mangle and L. racemosa harbor similar assemblages of cyanobacteria at each point along the transect. A. schaueriana, found only in the coastal portions of the transect, was colonized by assemblages with lower richness than the other trees. However, the results indicated that spatial location was a stronger driver of cyanobacterial community composition than plant species. Distinct cyanobacterial communities were observed at each location along the coast-to-upland transect. Clone library analysis allowed identification of 19 genera of cyanobacteria and demonstrated the presence of several uncultivated taxa. A predominance of sequences affiliated with the orders Nostocales and Oscillatoriales was observed, with a remarkable number of sequences similar to genera Symphyonemopsis/Brasilonema (order Nostocales). The results demonstrated that phyllosphere cyanobacteria in this mangrove forest ecosystem are influenced by environmental conditions as the primary driver at the ecosystem scale, with tree species exerting some effect on community structure at the local scale.     

Behavior of the groundwater in a riverine-type mangrove forest

The behavior of groundwater and physical properties of bottom sediment in a riverine-type mangrove forest which is composed of a tidal creek and fringing mangrove swamps were investigated through field observations at Iriomote Island, Japan. After the tidal water ebbed from the swamp surface to the creek, groundwater levels at swamp sites near the creek fell by up to 15 cm by the next flood tide, although the fall was negligible at sites far from the creek and at the open coast outside the mangrove forest. The amount of groundwater discharged to the creek from the swamp depended strongly both on the tidal range and the presence of the steep bank which separates the tidal creek from the fringing mangrove swamp. Based on the fall of groundwater level, the bulk hydraulic conductivity of the swamp was estimated to be 1.5×10⁻² cm/s. This value is two to three orders of magnitude larger than that measured in a laboratory using small scale sediment core samples collected in the swamp. These results suggest that the presence of crowded, intricate and large animal burrows as well as sediment layers rich in mangrove humus increases permeability in the mangrove swamp. Further, it is suggested that the mangrove topography with the steep bank of the tidal creek plays an important role which enhances material exchanges through groundwater between the mangrove swamp and the adjacent offshore waters.     

Microclimate of daytime den sites in a tropical possum: implications for the conservation of tropical arboreal marsupials

Tree cavities are an important shelter site for a variety of vertebrate species, including birds, reptiles and mammals. Studies indicate that in most taxa favoured tree hollows are those in larger trees and higher from the ground, generally thought to be related to decreased predation risk and a more optimal thermal environment. However, neither of these ideas has been tested definitively. Here, we investigate the microclimate of daytime den sites in tree hollows of common brushtail possums in tropical northern Australia. We compare tree and hollow characteristics of dens known to be used by possums, and those not known to be used, to determine whether possums choose trees with microhabitats with a more favourable daytime microclimate. Possums chose to den in tree hollows which were on average 1.6 °C cooler during the day, and were more buffered from temperature extremes, than other potential den locations. Important factors explaining daytime temperatures between hollows included height of the hollow, entrance width and tree diameter. Tropical arboreal marsupials have been identified as being particularly vulnerable to climate change and there are calls to identify and preserve natural refuges, such as tree hollows, which could buffer them from extreme temperatures. Our results highlight the value of older, larger hollow-bearing trees as refuges from extreme temperature, the importance of which may become critical for some temperature-sensitive species under the combined effects of continuing habitat loss and climate change.     

Ammonium adsorption in sediments of a tropical mangrove forest (Thailand) and a temperate Wadden Sea area (Denmark)

Dimensionless ammonium adsorption coefficients (K) were determined for tropical mangrove forest sediments and temperate Wadden Sea sediments. The K values were obtained from ammonium adsorption isotherms of KCl (2 M) extractable ammonium versus dissolved ammonium in the porewater; relationships that can be described by a linear model. Linearity was evident even at low porewater concentrations for mangrove sediment, according to isotherms based on KCl extractions on untreated sediment profiles. K-values were low in mangrove forest sediments (0.04–0.10), and higher in Wadden Sea sediments (0.17–1.12). The vertical range in K-values was larger at the vegetated sites, with highest values in subsurface sediments, which suggests differences in quantity and quality of the fine organic and inorganic fractions. The low ammonium adsorption in the mangrove sediments can be explained by a higher concentration of competitive cations, especially iron, in this iron-rich environment. The low adsorption of ammonium in mangrove sediments and vegetated surface sediment of the Wadden Sea was negatively related to the organic content of the sediments, which is in contrast to findings for other marine sediments. We suggest that organic material may have a diluting effect on the exchange capacity in fine-grained sediments, and that organic coatings may block ion exchange sites on clay surfaces. Thus, there may be a organic-rich ‘wetland’ versus organic-poor ‘sediment’ effect. This revised version was published online in July 2006 with corrections to the Cover Date.     

Resilience of arboreal folivores to habitat damage by a severe tropical cyclone

Abstract Severe tropical cyclones greatly modify habitat of arboreal folivores by destroying forest canopy, reducing structure and complexity and defoliating remaining trees. We hypothesized that forest modification following severe Cyclone Larry would stress arboreal folivores of the Family Pseudocheiridae and be reflected in increased home ranges and a decrease in body condition. We conducted 19 pre‐cyclone and 24 post‐cyclone spotlighting surveys at a site with severe cyclone damage, and 18 post‐cyclone surveys at a site with minor damage. We detected a greater number of lemuroid, Hemibelideus lemuroides and green, Pseudochirops archeri, ringtail possums as these possums remained in the severely damaged canopy and forest edge. In contrast, Herbert River ringtail possums, Pseudochirulus herbertensis, were detected in smaller numbers. We radio‐tracked eight P. herbertensis before the cyclone, following two of these and nine new animals after the category 4 cyclone. No significant post‐cyclone alteration in home range area or span was recorded in data pooled across the two sites or in limited post‐cyclone data at the severely disturbed site, but a greater variability in home range was observed after cyclone (pooled across sites: 1.72 ± 0.77 ha; 197 ± 47 m) than before the cyclone (1.35 ± 0.30 ha; 196 ± 23 m). In contrast, pooled pre‐ and post‐cyclone home range areas and spans were larger at the severely‐disturbed site (2.08 ± 0.56 ha; 231 ± 32 m) than at the site with minor damage (0.68 ± 0.11 ha; 114 ± 25 m), suggesting resources were more widely spread at the former site. Post‐cyclone home ranges were also larger at the severely damaged site (severe: 3.33 ± 1.36 ha, n = 3; minor: 0.52 ± 0.07 ha, n = 4). Condition of P. herbertensis (mass/tail length) did not differ significantly pre‐ and post‐cyclone or between less and severely disturbed sites. These results and observations of breeding after cyclone suggest that possum populations may be resilient to severe cyclone damage under the relatively wet conditions experienced post‐Cyclone Larry.