Feeding guild structure of beetles on Australian tropical rainforest trees reflects microhabitat resource availability

1.?We tested the hypotheses that feeding guild structure of beetle assemblages changed with different arboreal microhabitats and that these differences were consistent across rainforest tree species. 2.?Hand collection and beating techniques were used from the gondola of the Australian Canopy Crane to collect beetles from five microhabitats (mature leaves, flush leaves, flowers, fruit and suspended dead wood) within the rainforest canopy. A simple randomization procedure was implemented to test whether the abundances of each feeding guild on each microhabitat were different from that expected based on a null hypothesis of random distribution of individuals across microhabitats. 3.?Beetles from different feeding guilds were not randomly distributed, but congregated on those microhabitats that are likely to provide the highest concentrations of their preferred food sources. Herbivorous beetles, in particular, were over-represented on flowers and flush foliage and under-represented on mature leaves and dead wood. Proportional numbers of species within each feeding guild were remarkably uniform across tree species for each microhabitat, but proportional abundances of feeding guilds were all significantly non-uniformly distributed between host tree species, regardless of microhabitat, confirming patterns previously found for arthropods in trees in temperate and tropical forests. 4.?These results show that the canopy beetle community is partitioned into discrete assemblages between microhabitats and that this partitioning arises because of differences in feeding guild structure as a function of the diversity and the temporal and spatial availability of resources found on each microhabitat.     

Body size variation among invertebrates inhabiting different canopy microhabitat: flower visitors are smaller

1. Factors such as reproductive fitness, climatic tolerance, predation pressure, energetic requirements and the quality and quantity of food sources all correlate with invertebrate body sizes. 2. This study examines body size variation between an invertebrate community inhabiting five different microhabitats (mature leaves, new leaves, flowers, fruit and suspended dead wood) that differ in quality, quantity, and availability in an Australian tropical rainforest canopy. 3. Mean body size varied significantly between invertebrate and beetle feeding guilds across microhabitats. Phylogenetically independent contrasts revealed that invertebrate taxonomic groups were significantly smaller on flowers than on mature and new leaves. Size differences between microhabitats were most pronounced among herbivorous taxa (Hemiptera, Lepidoptera). In particular, the immature stages or those groups that develop on flowers were significantly smaller on flowers and larger on leaves than expected. Taxonomic groups with many strong flying species, especially those that complete larval development on resources other than flowers, typically showed no differences in body size across microhabitats. 4. There are a number of potential hypotheses for the smaller body sizes of flower visitors, including: (i) differences in the physical sizes of the microhabitats; (ii) variation in time–dependent mortality risks that influence development times; and (iii) differences in the nutritional quality of the microhabitats, which can influence body size via metabolic pathways. 5. The findings of this study do not support hypothesis (i) (with the possible exception of one or two predatory groups). It is suggested that hypotheses (ii (time–dependent mortality factors) and particularly (iii) (nutritional variation) may be the best avenues for future study as the main drivers of body size differences between microhabitats.     

The Overlooked Biodiversity of Flower-Visiting Invertebrates

Estimates suggest that perhaps 40% of all invertebrate species are found in tropical rainforest canopies. Extrapolations of total diversity and food web analyses have been based almost exclusively on species inhabiting the foliage, under the assumption that foliage samples are representative of the entire canopy. We examined the validity of this assumption by comparing the density of invertebrates and the species richness of beetles across three canopy microhabitats (mature leaves, new leaves and flowers) on a one hectare plot in an Australian tropical rainforest. Specifically, we tested two hypotheses: 1) canopy invertebrate density and species richness are directly proportional to the amount of resource available; and 2) canopy microhabitats represent discrete resources that are utilised by their own specialised invertebrate communities. We show that flowers in the canopy support invertebrate densities that are ten to ten thousand times greater than on the nearby foliage when expressed on a per-unit resource biomass basis. Furthermore, species-level analyses of the beetle fauna revealed that flowers support a unique and remarkably rich fauna compared to foliage, with very little species overlap between microhabitats. We reject the hypothesis that the insect fauna on mature foliage is representative of the greater canopy community even though mature foliage comprises a very large proportion of canopy plant biomass. Although the significance of the evolutionary relationship between flowers and insects is well known with respect to plant reproduction, less is known about the importance of flowers as resources for tropical insects. Consequently, we suggest that this constitutes a more important piece of the ‘diversity jigsaw puzzle’ than has been previously recognised and could alter our understanding of the evolution of plant-herbivore interactions and food web dynamics, and provide a better foundation for accurately estimating global species richness.     

Patterns of invertebrate food availability and the persistence of an avian insectivore on the brink

Globally, insectivorous birds are at high risk of decline. One explanation of this relates to changes in invertebrate resources due to anthropogenic pressures. The northern population of the eastern bristlebird (Dasyornis brachypterus) relies heavily on invertebrate food resources, and has experienced an 80% population reduction over the past 40 years. We investigated invertebrate abundance and nutritional quality across 23 currently and historically occupied northern bristlebird sites to determine whether extant territories were associated with more, or more nutritious, invertebrate resources. Pitfall and leaf‐litter invertebrate sampling were done in both breeding and non‐breeding seasons from 2014 to 2016. There was no difference in abundance, biomass or nutritional value of invertebrates between occupied and abandoned territories; however, within territories invertebrate abundance and nutritional value did correspond to the habitat characteristics with which bristlebirds are associated. Nutritional value of invertebrates increased with proximity to rainforest, while the abundance of macro‐invertebrates (>1 mm) was correlated with grass height. Bristlebird territories are often close to rainforest margins, and these ecotones may provide more nutritious mesic‐associated invertebrates. Higher abundances of large invertebrates in tall grasses may also contribute to the known association of bristlebirds with tall grasses. Maintenance of tall grass adjacent to rainforest through appropriate fire and grazing management is likely to be important for northern bristlebird recovery and long‐term persistence of the population.     

Specialization of rainforest canopy beetles to host trees and microhabitats: not all specialists are leaf‐feeding herbivores

Reliable estimates of host specificity in tropical rainforest beetles are central for an understanding of food web dynamics and biodiversity patterns. However, it is widely assumed that herbivores constitute the majority of host specific species, and that most herbivore species feed on leaves. We tested the generality of this assumption by comparing both plant host‐ and microhabitat‐specificity between beetle communities inhabiting the foliage (flush and mature), flowers, fruit, and suspended dead wood from 23 canopy plant species in a tropical rainforest in north Queensland, Australia. Independent of host tree identity, 76/77 of the most abundant beetle species (N ≥ 12 individuals) were aggregated on a particular microhabitat. Microhabitat specialization (measured by S_m and Lloyd's indices) was very high and did not differ between flower and foliage communities, suggesting that each newly‐sampled microhabitat has a large additive effect on total species richness. In accordance with previous studies, host specificity of foliage‐inhabiting beetles was most pronounced among herbivorous families (Curculionidae, Chrysomelidae). By contrast, host specificity among flower‐visitors was equally high among herbivorous and nonherbivorous families (e.g. Nitidulidae, Staphylinidae, Cleridae). Effective specialization (F_T) measures showed that traditional correction factors used to project total species richness in nonherbivorous groups fail to fully capture diversity in the flower‐visiting beetle fauna. These results demonstrate that host specialization is not concentrated within folivores as previously assumed. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 215–228.     

Terrestrial Invertebrate Community Structure as an Indicator of the Success of a Tropical Rainforest Restoration Project

Rainforest restoration is a relatively new endeavor, and few attempts have been made to assess the success of such restoration efforts in terms of the reestablishment of an ecosystem. Small plantings of rainforest tree species have been carried out adjacent to mature rainforest at Lake Barrine National Park in North Queensland, Australia, since 1988. The aim of this project was to assess the leaf litter invertebrate fauna of these plantings as indicators of the success of the restoration process. Plots planted in 1988, 1989, and 1990, as well as adjacent mature rainforest, were sampled in the wet and dry seasons of 1994. Invertebrates were extracted from leaf litter samples with Berlese Funnels and sorted to order. Diversity, the abundance of different size classes and orders of invertebrates, and the abundance of different functional groups were examined. In most respects the 1988 plot was found to differ little from the mature rainforest plots, whereas the 1990 plot lacked small and predatory invertebrates, especially in the dry season. The 1989 plot was intermediate in invertebrate abundance and diversity. The use of partially deciduous trees in the 1989 and 1990 plots, resulting in lower canopy cover at the driest time of the year, may have contributed significantly to the differences found between the early and later plantings. It is recommended that trees that provide good canopy cover year-round be used as dominant species in plantings to facilitate the development of a leaf litter ecosystem that can be sustained throughout the year.     

Substrate requirements of red-listed saproxylic invertebrates in Sweden

To evaluate which qualities of dead wood have the highest conservational value for invertebrates, substrate requirements were recorded for all 542 saproxylic (wood-living) red-listed invertebrates in Sweden. Preferred qualities of dead trees were described for each species in seven category variables: tree species, stage of decay, type of main stem, coarseness of stem, part of tree, light preference and microhabitat. For each quality we quantified the number of red-listed insect species using it and the number that are specific to the quality. The number of species associated with different tree genera ranged from five to 202. Species-rich genera had a higher proportion of monophagous species. To what extent different tree genera have faunas complementary to each other was also analysed. As the wood decays, the host range broadens and the flora of decay fungi will probably take over as the most important factor determining the saproxylic fauna. Because there are many different types of rot required by different invertebrates, an abundance of similar looking dead trees, logs and snags are probably needed. Some 59% of the invertebrate species can live in sun-exposed sites. Most of these are favoured when the forest burns and at least 29 of them are more strongly associated with fires. For the former category, leaving dead wood on clear cuts should be an effective way to increase the amount of breeding substrate. However, some species, especially those living in the last successional stages, are dependent on shaded sites. Hollow tree trunks are another important microhabitat; 64 of the 107 species living there are specialists. To maintain all saproxylic species we need a diversity of substrate types and management methods. Our data can help to decide how to optimize the conservation measures taken during forest operations.     

Diet selection in the green ringtail possum (Pseudochirops archeri): A specialist folivore in a diverse forest

Abstract This study investigated the feeding ecology of the green ringtail possum, Pseudochirops archeri (Pseudocheiridae) in a tropical rainforest with 94 plant species in the canopy. Over 50% of tree use was from only four tree species, Aleurites rockinghamensis, Ficus fraseri, Arytera divaricata and Ficus copiosa. These species were used significantly more frequently than would be expected if tree species were selected randomly in proportion to their relative abundance in the forest. Conversely, 88 other tree species present were used less frequently than expected. Possums also favoured particular individual trees within some of the preferred tree species. In 91% of feeding observations, possums consumed mature leaves only. The availability of young leaves, flowers and fruit varied throughout the year, with a peak in availability of these resources during the early wet season. By primarily selecting mature leaves, green ringtail possums reduce their dependence on seasonally variable resources. We suggest that green ringtail possums should be considered as specialist folivores, focusing on only a few of the tree species available, possibly due to advantages associated with limiting the number of plant secondary metabolites in their diet. Furthermore, they favour certain individual trees within species, perhaps due to intraspecific variation in plant secondary metabolites or nutrient content, behaviour that has been well established in eucalypt folivores. We conclude that green ringtail possums are highly specialized in their feeding ecology, limiting their diet to a small number of continuously available food items.     

Species richness of phytophagous beetles in the tropical tree Brosimum utile (Moraceae): the effects of sampling strategy and the problem of tourists

Abstract.  1. Phytophagous beetles on six mature living trees and two dead trees of Brosimum utile (Moraceae) were surveyed during 1 year in a tropical wet forest in Panama. The dead trees were surveyed both as suspended in the canopy and after falling down to the understorey.
2. Canopy access was provided by a construction crane and sampling was performed by beating and hand-collecting. The same amount of time was spent on each tree in order to standardise sampling effort. A list of all species associated with the tree is presented.
3. A total of 3009 individuals representing 364 species were collected. Tourists were excluded from the analyses by recording host associations directly and by probability assessments of host associations based on abundance categories. A total of 2603 individuals and 244 species were associated with the tree. The proportion of tourists in the trees increased with sample size.
4. A single mature living tree had on average 58.5 ± 6.5 species. The local species richness of B. utile was estimated as 2.5 times higher than in a single mature tree; however, a substantial increase in species richness was attained when dead wood habitats were included. Saproxylic species made up 82% of the total material.
5. The investigated habitat types of B. utile constituted distinct, complementary species assemblages. Similarity between saproxylic species of dead suspended wood and dead understorey wood of the same tree was 0.2 (Morisita–Horn index), confirming a prominent vertical stratification among this guild.     

Detecting impacts of non‐native species on associated invertebrate assemblages depends on microhabitat

Invasive plants that displace native floral communities can cause changes to associated invertebrate species assemblages. Using a mini‐review of the literature and our own data we add to the still considerable debate about the most effective methods for testing community‐level impacts by invasive species. In endangered saltmarshes of southeast Australia, the non‐native rush Juncus acutus L. is displacing its native congener J. kraussii Hochst., with concurrent changes to floral and faunal assemblages. In two coastal saltmarshes, we tested the hypothesis that the ability to detect differences in the invertebrate assemblage associated with these congeneric rushes depends on the microhabitat of the plant sampled. We used three sampling methods, each targeting specific microhabitats: sweep netting of the plant stems, vacuum sampling of the plant tussock, and vacuum sampling of the ground directly below the plants. Over 3800 individuals and 92 morphospecies were collected across four main taxa: gastropods, crustaceans, hexapods and arachnids. Detection of differences in invertebrate density, richness and composition associated with native compared with non‐native rushes was dependent on the microhabitat sampled and these differences were spatially variable. For example, at one saltmarsh the stems and tussock of J. acutus had a lower density and richness of total invertebrates and hexapods than those of the native J. kraussii. In contrast, crustaceans on the ground were in greater abundance below J. acutus than J. kraussii. This study demonstrates that on occasions where overall differences in the assemblage are not detected between species, differences may become apparent when targeting different microhabitats of the plant. In addition, separately targeting multiple microhabitats likely leads to a greater probability of detecting impacts of invasion. Comparing the invertebrate assemblage without differentiating between or sampling an array of microhabitats can fail to determine the impact of invasive species. These results highlight that a combination of methods targeting different microhabitats is important for detecting differences within the invertebrate community, even for phylogenetically related species.     

Genetic effects of tank-forming bromeliads on the associated invertebrate community in a tropical forest ecosystem

Within the area of community genetics there is an expanding volume of literature demonstrating how within-species genetic variation in temperate trees can have important effects on structuring animal and plant communities. The influence of intraspecific genetic variation on associated communities in relatively more complex ecosystems is only starting to be appreciated. Within tropical forests, epiphytic bromeliad plants often grow high in the canopy and create unique nutrient-rich microhabitats on which many invertebrate and vertebrate species depend. We investigated the influence of within-species genetic variation in the bromeliad Aechmea bracteata on the invertebrate microhabitat community. We found that more genetically similar bromeliad plants were host to more similar communities of juvenile-stage invertebrates, but not adult invertebrates. We discuss possible mechanisms for this, including differential survival and active female oviposition choice. Our work shows that the impact of within-species genetic variation on associated ecological communities may be more general than previously considered. These results agree with recent research suggesting that within-species genetic variation may perform a supporting ecosystem service for maintaining community and ecological processes.     

Relationship between invertebrate fauna and bromeliad size.

Several bromeliads species store water and organic substrates, allowing the establishment of phytotelmata and associated fauna on their leaves. In this study, we sampled 70 individuals of Vriesea sp. (Carrière) (Bromeliaceae), in rupestrian fields in the Serra de Ouro Branco-MG, Brazil. The relationships between invertebrate species richness and abundance and size of bromeliads were tested using multiple regression. We found 19 species associated with bromeliads, mainly Diptera larvae. The abundance of the phytotelmate fauna increased principally in relation to the volume of water in the bromeliad reservoir. Phytotelmata richness was affected principally by diameter of the reservoir. There was a significant relationship between the abundance and richness of invertebrates associated with leaves with diameter and height of the plant. Invertebrate richness was better explained by abundance of individuals. These results suggest that the increase of richness was attended by higher numbers of microhabitats and more space for colonization of bigger bromeliads. Additionally, there was more chance of sampling different species in locales with greater abundance of individuals.     

Vertical stratification of feeding guilds and body size in beetle assemblages from an Australian tropical rainforest

Abstract The vertical stratification of insect species assemblages inhabiting tropical rainforests is well established but few have examined whether these patterns are reflected in vertical stratification of body size or feeding guilds. We used Malaise and Flight Interception Traps to sample beetle assemblages from five locations, at both canopy and ground zones of a tropical lowland rainforest site near Cape Tribulation, Australia. Beetles from 4 years of sampling were sorted to Family and morphospecies, and allocated to one of five feeding guilds. Within feeding guilds the number of species and individuals, from canopy‐ and ground‐caught traps were compared. The body lengths of species were measure and compared within feeding guilds and families. Herbivores was the dominant guild but was not the majority of all species or individuals. Most beetle species (69%) were less than 5 mm in length and the mean size of canopy‐caught species was greater than that for ground‐caught species. This was probably due to slightly more species of plant feeders (herbivores and xylophages) present in the canopy, which were significantly larger than saprophages, fungivores and predators. Among feeding guilds, there were few overall canopy–ground differences. These results contrast with species composition results presented elsewhere where strong differences between the canopy and the ground were evident. We suggest that our guild groupings may have been too coarse to detect fine‐scale differences and that resource partitioning may have also masked faunal stratification. We propose that fine‐scale differences in resources between the canopy and the ground, together with strong microclimate gradients, are likely to be important in structuring the vertical stratification of insect assemblages at the level of species, but not with respect to functional groups.     

Survival of Flower-visiting Chrysomelids during Non General-flowering Periods in Bornean Dipterocarp Forests

In SE Asian rain forests, general flowering, a community-wide synchronous flowering, occurs at irregular and supra-annual intervals. During general flowering periods (GFP), most Dipterocarpaceae and many other trees flower profusely, while flowering plants are scant between GFP. During flowerless periods, anthophilous animals that depend on floral resources for food may suffer food shortages and subsequently decrease in abundance. Flower-visiting chrysomelid adults are major pollinators for some canopy tree species that flower during GFP. Although such chrysomelids feed on flower petals, the means by which they survive flowerless periods remains unknown. We determined the abundance of flower-visiting chrysomelids in GFP and non-GFP through light trap samples and examined the effects of the presence of young leaves and flowers of dipterocarps on local abundance, and feeding preferences of flower-visiting chrysomelids. We found no clear tendency that the chrysomelid species number and the abundance during GFP were consistently higher than those during non-GFP. Chrysomelid adults were more abundant on trees with many young leaves or flowers than on trees lacking young leaves and flowers. At least a few flower-visiting chrysomelids were observed feeding on young dipterocarp leaves and visiting young leaves and flowers of non-dipterocarps in the canopy during non-GFP. All results consistently suggest that chrysomelids are able to survive flowerless periods by feeding on the young leaves of dipterocarps and on the young leaves and flowers of non-dipterocarps; through this alternate feeding, chrysomelid populations are maintained at sufficient levels to function as effective pollinators of trees that flower during GFP.     

The invertebrate fauna of epiphyte mats in the canopy of northern rata (Myrtaceae: Metrosideros robusta A. Cunn.) on the West Coast of the South Island,New Zealand

Abstract

The range of microhabitats and microclimatic conditions provided by epiphytes has been linked to the high diversity of invertebrates in many forest canopies worldwide, but comparably little is known about the invertebrate fauna in this habitat in New Zealand. This study compiled an inventory of the invertebrate fauna of epiphyte mats in the canopy of northern rata (Myrtaceae: Metrosideros robusta A. Cunn.) at two study sites on the West Coast of the South Island. A total of 242 069 invertebrate specimens was collected over one year, representing 4 phyla, 9 classes and more than 160 families, 225 genera and 446 species. At least 10 new species and 3 new genera were identified, while 5 species were recorded outside their known geographical range. Epiphyte mats provided habitat for an invertebrate fauna, highly diverse and abundant both taxonomically and functionally, dominated in terms of abundance by Acari, Collembola and Hymenoptera (largely ants), and in terms of feeding guilds by epiphyte grazers and ants. As the first inventory of this taxonomic depth and breadth compiled for New Zealand forest epiphyte habitats, this study provides important baseline data for the conservation of biodiversity in New Zealand's indigenous forests.     

The microhabitat distribution of two Dactylogyrus species parasitizing the gills of the barbel, Barbus barbus

The microhabitat distribution of two congeneric species Dactylogyrus carpathicus and D. malleus (Monogenea) parasitizing the gills of the barbel (Barbus barbus L.) was investigated. We tested whether congeneric species exhibited microhabitat preference and whether interspecific interactions could be attributed to the microhabitat segregation of congeners. The outlying mean index method was used to evaluate species microhabitats. Gill variables (different microhabitats within gills) were used as environmental factors characterizing the gills. When abundances of both species were highest, and no significant difference was found between the abundance of the two species, the gill segments and gill areas were the most important factors segregating the Dactylogyrus species on the gills. Niche overlap was low within each of the four gill arches, and parasites were segregated in the same microhabitats within each gill arch. When abundances of both species were low, each monogenean species was segregated at the level of the gill arches. When abundances of both species increased, the niche and overlap between species increased. The distribution of both congeneric species confirmed microhabitat preference within the gills. The results suggest that microhabitat preference is dependent on species abundances, species being segregated in the case of low abundance, possibly to increasing mating opportunities. Both niche and overlap between species increased with species abundance. In the case of the high abundance of both species, microhabitat preference seems to be related to interspecific interactions between monogenean species, as previously found for endoparasitic species.     

Aquatic macrophytes, macroinvertebrate associations and water levels in a lowland Tasmanian river

Aquatic macrophytes are a common habitat for macroinvertebrates and may occupy depth zones in the littoral region of lowland rivers. Studies have indicated that different species of macrophyte typically support different assemblages, abundances and numbers of species of macroinvertebrates. This has often been attributed to differences in the dissectedness of stems and leaves of the macrophytes, resulting in differences in the surface area and/or the number of microhabitats available to invertebrates. I set out to measure the abundance and taxonomic richness and to describe the macroinvertebrate assemblages associated with three species of aquatic macrophyte in a pool in the Macquarie River, Tasmania and to examine responses of these variables to changes in water levels over summer. The macrophyte species sampled wereMyriophyllum simulans/variifolium, Triglochin procera} and Eleocharis sphacelata, each one differing in the dissectedness of its stems and leaves and its location in the littoral zone. Whereas the greatest abundance of macroinvertebrates was found associated in all months (i.e. at all water levels) with the structurally complex and shallowest macrophyte species, Myriophyllum, the number of taxa associated with this species was in several cases lower than for the structurally simpler and deeper water Triglochin and Eleocharis. While water depth and total plant biomass of samples were often correlated with invertebrate abundance and richness, these relationships were different for each macrophyte species. Of the nine most common invertebrate taxa collected from all samples, the abundances of more than half showed consistent differences among macrophyte species across months, two showed differences among macrophytes, but with an interaction with month and two showed no differences among macrophytes. There were major differences in the invertebrate assemblages associated with each macrophyte species in any one month, however, there was also a large turnover of taxa associated with the species of macrophytes from one month to the next. Changes in water level and concomitant changes in environmental variables are suggested as factors influencing the invertebrate fauna in the littoral zone of the pool of the Macquarie River. It is thus important for river managers to be aware that species of macroinvertebrates are not evenly distributed across species of macrophyte and that water levels and their influence on macrophytes as invertebrate habitat may play an integral part in determining the abundance, richness and assemblage of invertebrates in rivers.