Host associations and beta diversity of fungal endophyte communities in New Guinea rainforest trees

Processes shaping the distribution of foliar fungal endophyte species remain poorly understood. Despite increasing evidence that these cryptic fungal symbionts of plants mediate interactions with pathogens and herbivores, there remain basic questions regarding the extent to which dispersal limitation and host specificity might shape fungal endophyte community composition in rainforests. To assess the relative importance of spatial pattern and host specificity, we isolated fungi from a sample of mapped trees in lowland Papua New Guinea. Sequences of the internal transcribed spacer (ITS) region were obtained for 2079 fungal endophytes from three sites and clustered into molecular operational taxonomic units (MOTUs) at 95% similarity. Multivariate analyses suggest that host affinity plays a significant role in structuring endophyte community composition whereas there was no evidence of endophyte spatial pattern at the scale of tens to hundreds of metres. Differences in endophyte communities between sampled trees were weakly correlated with variation in foliar traits but not with tree species relatedness. The dominance of relatively few generalist endophytes and the presence of a large number of rare MOTUs was a consistent observation at three sites separated by hundreds of kilometres and regional turnover was low. Host specificity appears to play a relatively weak but more important role than dispersal limitation in shaping the distribution of fungal endophyte communities in New Guinea forests. Our results suggest that in the absence of strong ecological gradients and host turnover, beta diversity of endophyte communities could be low in large areas of contiguous forest.     

Gall‐forming insects in a lowland tropical rainforest: low species diversity in an extremely specialised guild

1. Gall‐forming insects are a guild of endophages that exhibit a high level of fidelity to their host plants, however, their level of host specificity is seldom explicitly tested. 2. Gall‐forming insect taxa from 32 species of woody tropical plants with resolved phylogenetic relationships were collected and reared, representing 15 families from all the major clades of angiosperms, at three lowland rainforest locations in Madang, Papua New Guinea (PNG). 3. More than 8800 galled plant parts were collected from 78 gall morphospecies at an average of 2.4 per host plant. Total species richness at the sampling sites was estimated to be 83–89. All but one morphospecies were monophagous resulting in an effective specialisation of 0.98. 4. Specific leaf weight, foliar nitrogen, the presence of latex, and the successional preference of plant species all gave a phylogenetic signal, but only plant successional preference influenced the species richness of galls on analysis of phylogenetically independent contrasts. Gall species were distributed randomly among host plant species and showed no preference for any particular plant lineage. Furthermore, most gall‐forming taxa were evenly dispersed across the host plant phylogeny. 5. In the tropical rainforests of New Guinea, gall‐forming insects are ubiquitous but occur in species‐poor assemblages. Local species richness is closely tied to the diversity of angiosperms owing to very high host specificity. 6. Finally, galler species richness data from the literature across habitats and latitudes were compared and suggest that tropical rainforests may be richer in galls than previously acknowledged.     

Elevational turnover in the composition of leaf miners and their interactions with host plants in Australian subtropical rainforest

Leaf miners are specialist herbivorous insects that are potentially vulnerable to environmental change because of their dependency on particular host plants. Little, however, is known about how climate affects the distribution of leaf miner communities and their interactions with host plants. Elevational gradients are useful tools for understanding how ecological communities respond to local clines in climate. Given that plant communities are known to undergo elevational turnover in response to changes in climatic conditions, we expect that leaf miner species will also change with elevation. We repeatedly hand collected leaf miners along three elevational gradients in subtropical rainforest in eastern Australia. Individual leaf miners were counted and identified to species, and their host plants were recorded. We tested if leaf miner species richness and the number of unique interactions among leaf miner and host plant species were affected by elevation. We also tested if the composition of leaf miner species and the composition of interactions between leaf miners and host plants showed a relationship with elevation. The rarefied number of unique leaf miner–host plant interactions significantly decreased with elevation, with a slight peak at approx. 700 m a.s.l., while neither rarefied or observed species richness (species density) of leaf miners nor observed numbers of unique interactions (interaction density) were significantly affected by elevation. The composition of leaf miner species and the composition of leaf miner–host plant interactions (occurrence of pairwise interactions) were significantly related to elevation. Elevational turnover in leaf miner species composition indicated that different species varied in their response to changes in biotic and/or abiotic conditions imposed by increasing elevation. Through our analyses, we identified four leaf miner species that may be locally vulnerable to climate change, as a result of their restricted elevational distribution and level of host specificity.     

Elevational filtering and the evolution of planthoppers (Hemiptera,Fulgoromorpha) in Papua New Guinea

Along elevational gradients, phylogenetic relatedness patterns constitute a considerable source of information and may shed light on ecological processes that structure communities. This study focuses on community phylogenetic structure of planthoppers, specifically the species-rich and abundant Fulgoromorpha families (Hemiptera, Auchenorrhyncha), Cixiidae and Derbidae + Achilidae, along an elevational gradient on Mount Wilhelm (Papua New Guinea). In order to assess the factors driving planthoppers community composition, we recorded abundance data for planthoppers species at each elevation and we generated a molecular phylogeny of the local species, using Bayesian inference. We analyzed 168 individuals representing 59 local morphospecies. Using a fully resolved and well-supported phylogeny, we then investigated the phylogenetic structure of the communities by performing a Spatial Analysis of Community Diversity. We show that Cixiidae are phylogenetically clustered along the elevational gradient, whereas Derbidae + Achilidae harbor a random structure, suggesting that local adaptation to elevation shapes community structure of Cixiidae, but not that of Derbidae + Achilidae. Our findings highlight the importance of phylogenies in the study of tropical elevational gradients.     

Host specificity of ambrosia and bark beetles (Col., Curculionidae: Scolytinae and Platypodinae) in a New Guinea rainforest

Abstract.  1. Bark and ambrosia beetles are crucial for woody biomass decomposition in tropical forests worldwide. Despite that, quantitative data on their host specificity are scarce.
2. Bark and ambrosia beetles (Scolytinae and Platypodinae) were reared from 13 species of tropical trees representing 11 families from all major lineages of dicotyledonous plants. Standardised samples of beetle-infested twigs, branches, trunks, and roots were taken from three individuals of each tree species growing in a lowland tropical rainforest in Papua New Guinea.
3. A total of 81 742 beetles from 74 species were reared, 67 of them identified. Local species richness of bark and ambrosia beetles was estimated at 80–92 species.
4. Ambrosia beetles were broad generalists as 95% of species did not show any preference for a particular host species or clade. Similarity of ambrosia beetle communities from different tree species was not correlated with phylogenetic distances between tree species. Similarity of ambrosia beetle communities from individual conspecific trees was not higher than that from heterospecific trees and different parts of the trees hosted similar ambrosia beetle communities, as only a few species preferred particular tree parts.
5. In contrast, phloeophagous bark beetles showed strict specificity to host plant genus or family. However, this guild was poor in species (12 species) and restricted to only three plant families (Moraceae, Myristicaceae, Sapindaceae).
6. Local diversity of both bark and ambrosia beetles is not driven by the local diversity of trees in tropical forests, since ambrosia beetles display no host specificity and bark beetles are species poor and restricted to a few plant families.     

Multilocus population genetic analysis of the Southwest Pacific malaria vector Anopheles punctulatus

The population structure and history of the cryptic malaria vector species, Anopheles punctulatus (Doenitz), was investigated throughout Papua New Guinea and the Solomon Islands with the aim of detailing genetic subdivisions and the potential for movement through this biogeographically complex region. We obtained larval collections from over 80 sites and utilised a diverse array of molecular markers that evolve through different processes. Individuals were initially identified to species and genotyped using the ribosomal DNA second internal transcribed spacer. DNA sequencing of a single copy nuclear ribosomal protein S9 and the mitochondrial cytochrome oxidase I loci were then investigated and 12 nuclear microsatellite markers were developed and analysed. Our data revealed three genetically distinct populations – one in Papua New Guinea, the second on Buka Island (Bougainville Province, Papua New Guinea), and the third on Guadalcanal Island (Solomon Islands). Genetic differentiation within Papua New Guinea was much lower than that found in studies of other closely related species in the region. The data does suggest that A. punctulatus has undergone a population bottleneck followed by a recent population and range expansion in Papua New Guinea. Humans and regional economic growth may be facilitating this population expansion, as A. punctulatus is able to rapidly occupy human modified landscapes and traverse unsealed roads. We therefore anticipate extensive movement of this species through New Guinea – particularly into the highlands, with a potential increase in malaria frequency in a warming climate – as well as relatively unrestricted gene flow of advantageous alleles that may confound vector control efforts.     

The insect‐focused classification of fruit syndromes in tropical rain forests: An inter‐continental comparison

We propose a new classification of rain forest plants into eight fruit syndromes, based on fruit morphology and other traits relevant to fruit‐feeding insects. This classification is compared with other systems based on plant morphology or traits relevant to vertebrate fruit dispersers. Our syndromes are based on fruits sampled from 1,192 plant species at three Forest Global Earth Observatory plots: Barro Colorado Island (Panama), Khao Chong (Thailand), and Wanang (Papua New Guinea). The three plots differed widely in fruit syndrome composition. Plant species with fleshy, indehiscent fruits containing multiple seeds were important at all three sites. However, in Panama, a high proportion of species had dry fruits, while in New Guinea and Thailand, species with fleshy drupes and thin mesocarps were dominant. Species with dry, winged seeds that do not develop as capsules were important in Thailand, reflecting the local importance of Dipterocarpaceae. These differences can also determine differences among frugivorous insect communities. Fruit syndromes and colors were phylogenetically flexible traits at the scale studied, as only three of the eight seed syndromes, and one of the 10 colors, showed significant phylogenetic clustering at either genus or family levels. Plant phylogeny was, however, the most important factor explaining differences in overall fruit syndrome composition among individual plant families or genera across the three study sites. Abstract in Melanesian is available with online material.     

Inter-specific aggression generates ant mosaics in canopies of primary tropical rainforest

The ant mosaic is a concept of the non-random spatial distribution of individual ant species in trees built upon the assumption of interspecific behavioural associations. However, colony identity and environmental variance may also play a role in species distribution. Here we assess the presence of ant mosaics in a primary forest ecosystem and whether they are structured by species' aggressive behaviours or by habitat filtering. We sampled arboreal ants from vertically stratified baits exposed in 225 canopy trees in a 9-ha plot of primary lowland forest in Papua New Guinea, the largest forest area surveyed to detect ant mosaics. We performed behavioural tests on conspecific ants from adjacent trees to determine the territories of individual colonies. We explored the environmental effects on the ant communities using information on the plot vegetation structure and topography. Furthermore, we created a novel statistical method to test for the community non-random spatial structure across the plot via spatial randomisation of individual colony territories. Finally, we linked spatial segregation among the four most common species to experimentally assessed rates of interspecies aggression. The ant communities comprised 57 species of highly variable abundance and vertical stratification. Ant community composition was spatially dependent, but it was not affected by tree species composition or canopy connectivity. Only local elevation had a significant but rather small effect. Individual colony territories ranged from one tree to 0.7 ha. Species were significantly over-dispersed, with their territory overlap significantly reduced. The level of aggression between pairs of the four most common species was positively correlated with their spatial segregation. Our study demonstrates the presence of ant mosaics in tropical pristine forest, which are maintained by interspecific aggression rather than habitat filtering, with vegetation structure having a rather small and indirect effect, probably linked to microclimate variability.     

Quantifying the Human Impacts on Papua New Guinea Reef Fish Communities across Space and Time

Describing the drivers of species loss and of community change are important goals in both conservation and ecology. However, it is difficult to determine whether exploited species decline due to direct effects of harvesting or due to other environmental perturbations brought about by proximity to human populations. Here we quantify differences in species richness of coral reef fish communities along a human population gradient in Papua New Guinea to understand the relative impacts of fishing and environmental perturbation. Using data from published species lists we categorize the reef fishes as either fished or non-fished based on their body size and reports from the published literature. Species diversity for both fished and non-fished groups decreases as the size of the local human population increases, and this relationship is stronger in species that are fished. Additionally, comparison of modern and museum collections show that modern reef communities have proportionally fewer fished species relative to 19^th century ones. Together these findings show that the reef fish communities of Papua New Guinea experience multiple anthropogenic stressors and that even at low human population levels targeted species experience population declines across both time and space.     

Effects of phylogeny,leaf traits,and the altitudinal distribution of host plants on herbivore assemblages on congeneric Acer species

Historical, niche-based, and stochastic processes have been proposed as the mechanisms that drive community assembly. In plant–herbivore systems, these processes can correspond to phylogeny, leaf traits, and the distribution of host plants, respectively. Although patterns of herbivore assemblages among plant species have been repeatedly examined, the effects of these factors among co-occurring congeneric host plant species have rarely been studied. Our aim was to reveal the process of community assembly for herbivores by investigating the effects of phylogeny, leaf traits, and the altitudinal distribution of closely related host plants of the genus Acer. We sampled leaf functional traits for 30 Acer species in Japan. Using a newly constructed phylogeny, we determined that three of the six measured leaf traits (leaf thickness, C/N ratio, and condensed tannin content) showed a phylogenetic signal. In a field study, we sampled herbivore communities on 14 Acer species within an elevation gradient and examined relationships between herbivore assemblages and host plants. We found that herbivore assemblages were significantly correlated with phylogeny, leaf traits, phylogenetic signals, and the altitudinal distribution of host plants. Our results indicate that the interaction between historical and current ecological processes shapes herbivore community assemblages.     

Plant‐driven changes in soil microbial communities influence seed germination through negative feedbacks

相似文献   

Host specificity and interaction networks of insects feeding on seeds and fruits in tropical rainforests

In the tropics, antagonistic seed predation networks may have different properties than mutualistic pollination and seed dispersal networks, but the former have been considerably less studied. We tested whether the structure of antagonistic tripartite networks composed of host plants, insects developing within seeds and fruits, and their insect parasitoids could be predicted from plant phylogenetic distance and plant traits. We considered subsets of the networks (‘subnetworks') at three rainforest locations (Panama, Thailand, Papua New Guinea), based on insect families, plant families or plant functional groups. We recorded 3197 interactions and observed a low percentage of realized interactions, especially in Panama, where insect host specificity was higher than in Thailand or New Guinea. Several factors may explain this, including insect faunal composition, incidence of dry fruits, high fruit production and high occurrence of Fabaceae at the Panamanian site. Host specificity was greater among seed-eaters than pulp-eaters and for insects feeding on dry fruits as opposed to insects feeding on fleshy fruits. Plant species richness within plant families did not influence insect host specificity, but site characteristics may be important in this regard. Most subnetworks were extremely specialized, such as those including Tortricidae and Bruchinae in Panama. Plant phylogenetic distance, plant basal area and plant traits (fruit length, number of seeds per fruit) had important effects on several network statistics in regressions weighted by sampling effort. A path analysis revealed a weak direct influence of plant phylogenetic distance on parasitoid richness, indicating limited support for the ‘nasty host hypothesis'. Our study emphasizes the duality between seed dispersal and seed predation networks in the tropics, as key plant species differ and host specificity tends to be low in the former and higher in the latter. This underlines the need to study both types of networks for sound practices of forest regeneration and conservation.     

How specialised must natural enemies be to facilitate coexistence among plants?

The Janzen‐Connell hypothesis proposes that plant interactions with host‐specific antagonists can impair the fitness of locally abundant species and thereby facilitate coexistence. However, insects and pathogens that associate with multiple hosts may mediate exclusion rather than coexistence. We employ a simulation model to examine the effect of enemy host breadth on plant species richness and defence community structure, and to assess expected diversity maintenance in example systems. Only models in which plant enemy similarity declines rapidly with defence similarity support greater species richness than models of neutral drift. In contrast, a wide range of enemy host breadths result in spatial dispersion of defence traits, at both landscape and local scales, indicating that enemy‐mediated competition may increase defence‐trait diversity without enhancing species richness. Nevertheless, insect and pathogen host associations in Panama and Papua New Guinea demonstrate a potential to enhance plant species richness and defence‐trait diversity comparable to strictly specialised enemies.     

Higher host plant specialization of root‐associated endophytes than mycorrhizal fungi along an arctic elevational gradient

How community‐level specialization differs among groups of organisms, and changes along environmental gradients, is fundamental to understanding the mechanisms influencing ecological communities. In this paper, we investigate the specialization of root‐associated fungi for plant species, asking whether the level of specialization varies with elevation. For this, we applied DNA barcoding based on the ITS region to root samples of five plant species equivalently sampled along an elevational gradient at a high arctic site. To assess whether the level of specialization changed with elevation and whether the observed patterns varied between mycorrhizal and endophytic fungi, we applied a joint species distribution modeling approach. Our results show that host plant specialization is not environmentally constrained in arctic root‐associated fungal communities, since there was no evidence for changing specialization with elevation, even if the composition of root‐associated fungal communities changed substantially. However, the level of specialization for particular plant species differed among fungal groups, root‐associated endophytic fungal communities being highly specialized on particular host species, and mycorrhizal fungi showing almost no signs of specialization. Our results suggest that plant identity affects associated mycorrhizal and endophytic fungi differently, highlighting the need of considering both endophytic and mycorrhizal fungi when studying specialization in root‐associated fungal communities.     

An alien in an archipelago: Spathodea campanulata and the geographic variability of its moth (Lepidoptera) communities in the New Guinea and Bismarck Islands

Aim This analysis of moth (Lepidoptera) communities colonizing an alien tree invading secondary rain forest vegetation in Melanesia examines the predictability of insect herbivorous communities across distances of tens to thousands of km and the effect of dispersal barriers on community composition in the tropics. Location Six secondary rain forest sites were studied within four equidistant yet distinct geographic areas of the New Guinea mainland and the Bismarck Archipelago, including two watershed areas (Madang and Sepik) on mainland New Guinea and the adjacent large island of New Britain and small island of Unea. Methods The analysis is based on feeding records obtained by quantitative sampling and rearing of caterpillars from the alien host Spathodea campanulata (Bignoniaceae). It examines the variation in Lepidoptera community composition at six study sites distributed on three adjacent islands ranging in size from 30 to 865,000 km². Results Spathodea campanulata was colonized by 54 folivorous species of Lepidoptera. Most of them were generalists, feeding on > 1 native plant family. However, the three most abundant species representing 83% of all individuals (Acherontia lachesis, Hyblaea puera complex and Psilogramma menephron) were relatively host specific, feeding predominantly on a single native family that is not the Bignoniaceae. Most of the 23 species analysed in detail had a wide geographic distribution, including 13 species spanning the entire 1000‐km study transect. While the Lepidoptera in two New Guinea areas 280 km apart were similar to each other, there was a discontinuity in species composition between New Guinea and the smaller islands. However, no negative effect of small islands on species richness was detected. Main conclusions Spathodea campanulata was rapidly colonized by folivorous Lepidoptera communities with species richness and dominance structure indistinguishable from the assemblages feeding on native hosts, despite its phylogenetic isolation from the native vegetation. Although most species were generalists, the highest population densities were reached by relatively specialized species, similar to the communities on native hosts. The species turnover across distances from 10 to 1000 km was relatively low as most of the species had wide geographic ranges.     

Host functional and phylogenetic composition rather than host diversity structure plant–herbivore networks

Declining plant diversity alters ecological networks, such as plant–herbivore interactions. However, our knowledge of the potential mechanisms underlying effects of plant species loss on plant–herbivore network structure is still limited. We used DNA barcoding to identify herbivore–host plant associations along declining levels of tree diversity in a large‐scale, subtropical biodiversity experiment. We tested for effects of tree species richness, host functional and phylogenetic diversity, and host functional (leaf trait) and phylogenetic composition on species, phylogenetic and network composition of herbivore communities. We found that phylogenetic host composition and related palatability/defence traits but not tree species richness significantly affected herbivore communities and interaction network complexity at both the species and community levels. Our study indicates that evolutionary dependencies and functional traits of host plants determine the composition of higher trophic levels and corresponding interaction networks in species‐rich ecosystems. Our findings highlight that characteristics of the species lost have effects on ecosystem structure and functioning across trophic levels that cannot be predicted from mere reductions in species richness.     

High specialization and limited structural change in plant‐herbivore networks along a successional chronosequence in tropical montane forest

Secondary succession is well‐understood, to the point of being predictable for plant communities, but the successional changes in plant‐herbivore interactions remains poorly explored. This is particularly true for tropical forests despite the increasing importance of early successional stages in tropical landscapes. Deriving expectations from successional theory, we examine properties of plant‐herbivore interaction networks while accounting for host phylogenetic structure along a succession chronosequence in montane rainforest in Papua New Guinea. We present one of the most comprehensive successional investigations of interaction networks, equating to > 40 person years of field sampling, and one of the few focused on montane tropical forests. We use a series of nine 0.2 ha forest plots across young secondary, mature secondary and primary montane forest, sampled almost completely for woody plants and larval leaf chewers (Lepidoptera) using forest felling. These networks comprised of 12 357 plant‐herbivore interactions and were analysed using quantitative network metrics, a phylogenetically controlled host‐use index and a qualitative network beta diversity measure. Network structural changes were low and specialisation metrics surprisingly similar throughout succession, despite high network beta diversity. Herbivore abundance was greatest in the earliest stages, and hosts here had more species‐rich herbivore assemblages, presumably reflecting higher palatability due to lower defensive investment. All herbivore communities were highly specialised, using a phylogenetically narrow set of hosts, while host phylogenetic diversity itself decreased throughout the chronosequence. Relatively high phylogenetic diversity, and thus high diversity of plant defenses, in early succession forest may result in herbivores feeding on fewer hosts than expected. Successional theory, derived primarily from temperate systems, is limited in predicting tropical host‐herbivore interactions. All succession stages harbour diverse and unique interaction networks, which together with largely similar network structures and consistent host use patterns, suggests general rules of assembly may apply to these systems.     

Trophic cascades in tropical rainforests: Effects of vertebrate predator exclusion on arthropods and plants in Papua New Guinea

Insect herbivores have the potential to consume large amounts of plant tissue in tropical forests, but insectivorous vertebrates effectively control their abundances, indirectly increasing plant fitness accordingly. Despite several studies already sought understanding of the top-down effects on arthropod community structure and herbivory, such studies of trophic cascades in old tropics are underrepresented, and little attention was paid to top-down forces in various habitats. Therefore, we examine how flying insectivorous vertebrates (birds and bats) impact arthropods and, consequently, affect herbivore damage of leaves in forest habitats in Papua New Guinea. In a 3-month long predator exclosure experiment conducted at four study sites across varying elevation and successional stage, we found that vertebrate predators reduced arthropod density by ∼52%. In addition, vertebrate predators decreased the mean body size of arthropods by 26% in leaf chewers and 47% in non-herbivorous arthropods but had only a small effect on mesopredators and sap suckers. Overall, the exclusion of vertebrate predators resulted in a ~ 41% increase in leaf damage. Our results, across different types of tropical forests in Papua New Guinea, demonstrate that flying vertebrate insectivores have a crucial impact on plant biomass, create a selective pressure on larger and non-predatory prey individuals and they prey partition with mesopredators.     

A large trophic quilt

Plants and invertebrate herbivores are major constituents of terrestrial food webs. Identifying component species and tracing their interactions in highly diverse communities are a monumental task. Novotny et al. 2010 present the first broad conspectus of herbivore–plant interactions in a forest in Papua New Guinea. In more than 15 years, nearly 7000 feeding links were traced between about 200 plants and 1500 insect herbivores. Although staggering, these figures might represent only 15% of the total herbivore richness and interaction diversity in that lowland forest. Standardized comparisons also showed distinctive specialization and diversification patterns in different feeding guilds, restricting the possibility of using any single guild as surrogate for the entire assemblage.     

Specificity among the Casuarinaceae in root nodulation byFrankia

Pure cultured isolates ofFrankia made from root nodules of plant species from among three genera of the host family Casuarinaceae were used in inoculation trials of seedlings grown in water culture. A large number of host species among the genera Allocasuarina, Casuarina and Gymnostoma from Australia, Papua New Guinea and other South Pacific Islands were tested. The most widely infectiveFrankia strains were CcI3 and AllI1; theFrankia strains with the narrowest host range within the Casuarinaceae were CcI2 and GpI1. Intrafamily cross-inoculations were uncommon. The most broadly receptive host species wasG. papuanum. For many species ofAllocasuarina tested, no infection by anyFrankia available for testing could be observed.