The specialization and structure of antagonistic and mutualistic networks of beetles on rainforest canopy trees

Different kinds of species interactions can lead to different structures within ecological networks. Antagonistic interactions (such as between herbivores and host plants) often promote increasing host specificity within a compartmentalized network structure, whereas mutualistic networks (such as pollination networks) are associated with higher levels of generalization and form nested network structures. However, we recently showed that the host specificity of flower‐visiting beetles from three different feeding guilds (herbivores, fungivores, and predators) in an Australian rainforest canopy was equal to that of herbivores on leaves, suggesting that antagonistic herbivores on leaves are no more specialized than flower‐visitors. We therefore set out to test whether similarities in the host specificity of these different assemblages reflect similarities in underlying network structures. As shown before at the species level, mutualistic communities on flowers showed levels of specialization at the network scale similar to those of the antagonistic herbivore community on leaves. However, the network structure differed, with flower‐visiting assemblages displaying a significantly more nested structure than folivores, and folivores displaying a significantly more compartmentalized structure than flower‐visitors. These results, which need further testing in other forest systems, demonstrate that both antagonistic and mutualistic interactions can result in equally high levels of host specialization among beetle assemblages in tropical rainforests. If this is a widespread phenomenon, it may alter our current perceptions of food web dynamics, species diversity patterns, and co‐evolution in tropical rainforests. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 287–295.     

The induction of dominant lethal mutations in Tilapia mossambica by alkane sulphonic esters.

A serial mating system has been used to enable study of the effect of chemical mutagens in Tilapia mossambica. Embryopathies incompatible with survival are attributed to dominant lethal mutations induced in developing male germ cells by methyl methanesulphonate and the isomeric forms of dimethyl-myleran.     

The spatial and temporal distributions of arthropods in forest canopies: uniting disparate patterns with hypotheses for specialisation

Arguably the majority of species on Earth utilise tropical rainforest canopies, and much progress has been made in describing arboreal assemblages, especially for arthropods. The most commonly described patterns for tropical rainforest insect communities are host specificity, spatial specialisation (predominantly vertical stratification), and temporal changes in abundance (seasonality and circadian rhythms). Here I review the recurrent results with respect to each of these patterns and discuss the evolutionary selective forces that have generated them in an attempt to unite these patterns in a holistic evolutionary framework. I propose that species can be quantified along a generalist–specialist scale not only with respect to host specificity, but also other spatial and temporal distribution patterns, where specialisation is a function of the extent of activity across space and time for particular species. When all of these distribution patterns are viewed through the paradigm of specialisation, hypotheses that have been proposed to explain the evolution of host specificity can also be applied to explain the generation and maintenance of other spatial and temporal distribution patterns. The main driver for most spatial and temporal distribution patterns is resource availability. Generally, the distribution of insects follows that of the resources they exploit, which are spatially stratified and vary temporally in availability. Physiological adaptations are primarily important for host specificity, where nutritional and chemical variation among host plants in particular, but also certain prey species and fungi, influence host range. Physiological tolerances of abiotic conditions are also important for explaining the spatial and temporal distributions of some insect species, especially in drier forest environments where desiccation is an ever‐present threat. However, it is likely that for most species in moist tropical rainforests, abiotic conditions are valuable indicators of resource availability, rather than physiologically limiting factors. Overall, each distribution pattern is influenced by the same evolutionary forces, but at differing intensities. Consequently, each pattern is linked and not mutually exclusive of the other distribution patterns. Most studies have examined each of these patterns in isolation. Future work should focus on examining the evolutionary drivers of these patterns in concert. Only then can the relative strength of resource availability and distribution, host defensive phenotypes, and biotic and abiotic interactions on insect distribution patterns be determined.     

The relationship between fecundity and oocyte resorption in field populations of Lucilia cuprina

In the Australian sheep blowfly, Lucilia cuprina, the number of ovarioles, and hence the maximum number of eggs developed in each ovarian cycle, is a linear function of adult size (headwidth). Field females have a mean headwidth of 3.0 mm (range=2.30–3.50 mm) and a mean potential fecundity/cycle of 232 eggs (range=111–318 eggs). Realised fecundity is influenced by the quality and amount of protein in the adult diet. All field females appear to have the same minimum protein requirement for egg maturation, i.e., whatever their size (potential fecundity) females must obtain sufficient protein to mature at least 110 eggs in order to become gravid. Females whose protein intake exceeds this minimum but falls short of the amount needed for maturation of their full egg complements resorb some of their oocytes and mature the remainder. Since potential fecundity increases with fly size, large females resorb more oocytes than small females under suboptimal protein regimes, i.e., the proportion of oocytes resorbed increases with fly size. Under field conditions, females rarely matured full complements of eggs, which indicates a general shortage of protein-rich material. The mean reduction in fecundity of field females during these studies was 53.4 eggs/cycle (23.2%).

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Résumé Chez Lucilia cuprina Wied., le nombre d'ovarioles, et ainsi le nombre maximum d'oeufs développés au cours de chaque cycle ovarien, est une fonction linéaire de la taille de l'adulte (largeur de la tête). La largeur moyenne de la tête des femelles de la nature est 3 mm (2,3 à 3,5) et la fécondité potentielle moyenne par cycle de 232 oeufs (111 à 318).La fécondité réelle est influencée par la qualité et la quantité de protéines dans le régime alimentaire. Toutes les femelles de la nature semblent avoir les mêmes exigences minimales en protéines pour la maturation des ovocytes, c'est à dire que quelle que soit leur taille (fécondité potentielle) les femelles doivent obtenir les protéines suffisantes à la formation d'au moins 110 ovocytes pour devenir gravides. Les femelles dont l'absorption de protéines dépasse ce minimum, mais est insuffisant pour permettre la maturation de la totalité de leurs ovocytes, en résorbent quelques uns et conduisent à maturité le reste. Puisque la fécondité potentielle augmente avec la taille des mouches, les grosses femelles résorbent plus d'ovocytes que les petites avec un régime suboptimal en protéines, c'est à dire que la proportion d'ovocytes résorbés augmente avec la taille des mouches. Dans les conditions de la nature, la totalité des ovocytes parvient rarement à maturité, ce qui indique une limitation générale en aliments riches en protéines. La réduction moyenne de la fécondité chez les femelles de la nature a été au cours de cette étude de 53.4 oeufs par cycle (23,2%).

     

Embryopathies due to spermatozoal impairment in Xenopus laevis.

An in vitro test system, involving gametes of Xenopus laevis has been used to study the effect of antifertility agents upon spermatozoa as manifest in developing embryos. Exposure to either the isomeric forms of dimethylmyleran, methyl methanesulphonate or gamma-radiation led to development of a range of embryopathies, whereas treatment with steroidal drugs or the rodent epididymal chemosterilants alpha-chlorohydrin and trimethylphosphate was compatible with production of apparently normal offspring.     

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.     

Factors affecting abundance and oviposition rates of a field population of the Old World screw-worm fly, Chrysomya bezziana (Diptera: Calliphoridae)

Sentinel cattle and a grid of swormlure-baited sticky traps were used to monitor a Malaysian population of the Old World screw-worm fly, Chrysomya bezziana Villeneuve. Observations were carried out on an isolated cattle station at monthly intervals during the period August 1996 to June 2000. The number of flies caught was unaffected by weather conditions at the time of trapping, but was positively correlated with the total rainfall and the average daily air temperature prevailing 15-28 days earlier, when trapped flies were still juveniles. Trap catches were biased in favour of females, but daily catch rates of both sexes increased significantly the longer traps were open, suggesting that efficacy was related to the differential volatility of the chemicals comprising swormlure. Oviposition on sentinel cattle occurred mostly in late afternoon or early evening but increased significantly as the wound aged. Oviposition rates were positively correlated with female catch rates, but the relationship was curvilinear, suggesting that fly populations may be subject to some form of density-dependent constraint. Consistent differences in oviposition rates on sentinel cattle at different localities on the cattle station suggested the existence of highly clumped, quasi-stationary populations. Differences in trap catches between traps located in pastoral areas and those sited in nearby oil palm or rubber plantations supported this interpretation of the data. These findings are discussed in relation to the use of the sterile insect technique for the control of screw-worm fly infestations.     

Canopy invertebrate community composition on rainforest trees: Different microhabitats support very different invertebrate communities

Tropical rainforest canopies are renowned for their high invertebrate diversity and abundance. The canopy comprises a range of microhabitats representing very different food resources (including photosynthetic, reproductive, and structural tissues). As these resources vary considerably in temporal and spatial availability, nutritional quality, chemical protection and other attributes, we hypothesized that microhabitats support structurally different invertebrate communities. To test this we used the Australian Canopy Crane to sample invertebrates from mature leaves, flush leaves, flowers, fruit and suspended dead wood from 23 plant species. Invertebrate faunas on different microhabitats varied in taxonomic composition and feeding guild structure in support of the microhabitat differentiation hypothesis. Herbivores were found predominantly on new leaves (Hemiptera, Lepidoptera) and especially flowers (Coleoptera, Thysanoptera), but were relatively uncommon on mature leaves. Instead, the mature foliage community was dominated by predators, especially spiders and ants, and supported high abundances of saprophages. Ripe fruit and dead wood were scarce canopy resources that were utilized by a relatively small number of invertebrates, mostly saprophages and fungivores. Flowers supported a more heterogeneous fauna than the leaves in terms of proportional abundances of taxonomic groups and feeding guilds, both within tree species (evenness) and between tree species (non‐uniformity). These results demonstrate microhabitat differentiation in a rainforest canopy and are the first to quantify differences in taxonomic composition, guild structure and abundance patterns between such diverse invertebrate assemblages within host trees. We conclude that studies based only on sampling one microhabitat, and leaves in particular, may provide a distorted picture of invertebrate community structure.     

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.     

Vertical stratification in the spatial distribution of the beech scale insect (Ultracoelostoma assimile) in Nothofagus tree canopies in New Zealand

Abstract. 1. The degree of infestation by New Zealand sooty beech scale insects (Ultracoelostoma assimile, Homoptera: Margarodidae) varies dramatically among adjacent southern beech trees (Nothofagus spp., Fagaceae), but has previously been assumed to be uniformly or randomly distributed within individual host trees. In this study, a full‐census survey was conducted from ground level to canopy level on 14 naturally occurring, canopy‐dominant red beech (Nothofagus fusca) trees (size range 38.7–107.6 cm diameter at breast height) to determine the degree of within‐tree heterogeneity in herbivore density. 2. The within‐tree distribution of the sooty beech scale was vertically stratified and highly heterogeneous, with the greatest densities occurring on bark surfaces in the canopy rather than on the trunk, and on the lower rather than upper sides of the branches. The spatial distribution was strongly negatively correlated with trunk and branch diameter, and increasing bark thickness (as a function of diameter) provides a plausible explanation for differences in the establishment and population density of sooty beech scale insects with trunk and branch size. Furthermore, there was a significant change in the spatial distribution of scale insect populations on trunks and branches of trees of increasing diameter at breast height. This indicates a strong temporal component to the spatial dynamics of the sooty beech scale insect driven by changing host phenology. Future studies on phytophagous insects infesting large host trees need to consider more explicitly changes in population dynamics through space and time. 3. Because of the high degree of within‐tree heterogeneity in population density, the total population size of scale insects on an individual tree could not be predicted from any measure of population density low on the trunk. However, the dry weight biomass of sooty mould fungi growing on the ground beneath infested trees was a remarkably accurate predictor of the total population size of scale insects. The use of sooty mould fungi as a relative measure of population size could be incorporated into studies of other honeydew‐producing hemipterans, since the growth of sooty mould is a distinctive feature synonymous with high concentrations of honeydew production worldwide.