A behavioural analysis of phase change in the desert locust

A programme of research into phase change in the desert locust, Schistocerca gregaria, is described. The ability to change phase between solitarious and gregarious forms in response to population density is a key feature of locusts and is central to their occasional yet catastrophic impact on humans. Phase polymorphism is an extreme form of phenotypic plasticity. The most labile phase characteristic is behaviour. It is argued that a fully integrated study of behavioural phase change provides a powerful tool for understanding both the mechanisms of phase change and locust population dynamics, both of which offer possibilities for improved management and control of desert locust plagues. An assay for measuring behavioural phase-state in individual locusts was derived, based on logistic regression analysis. Experiments are described that used the assay to quantify the time-course of behavioural change, both within the life of individual locusts and across generations. The locust-related stimuli that provoke behavioural gregarization were investigated. Complex interactions were found between tactile, visual and olfactory stimuli, with the former exerting the strongest effect. Behavioural analysis also directed a study of the mechanisms whereby adult females exert an epigenetic influence over the phase-state of their developing offspring. Female locusts use their experience of the extent and recency of being crowded to predict the probability that their offspring will emerge into a high-density population, and alter the development of their embryos accordingly through a gregarizing agent added to the foam that surrounds the eggs at laying. There is also a less pronounced paternal influence on hatchling phase-state. An understanding of the time-course of behavioural phase change led to a study of the effect of the fine-scale distribution of resources in the environment on interactions between individual locusts, and hence on phase change. This, in turn, stimulated an exploration of the implications of individual behavioural phase change for population dynamics. Cellular automata models were derived that explore the relationships between population density, density of food resources and the distribution of resources in the environment. The results of the simulation showed how the extent of gregarization within a population increases with rising population size relative to food abundance and increasing concentration of food resources. Of particular interest was the emergence of critical zones across particular combinations of resource abundance, resource distribution and population size, where a solitarious population would rapidly gregarize. The model provided the basis for further laboratory and field experiments, which are described.     

Responses of tropical cladocerans to a gradient of resource quality

1. The response of three tropical cladocerans to a gradient of resource quality was compared in a series of growth bioassays using seston collected from five lakes of different depth and trophic structure in Michigan, U.S.A. To assess the food quality in terms of digestibility, assimilation experiments were performed with ³²P‐labelled seston from the same lakes. Animals were also analysed for P‐content in their tissues at the end of these assays. 2. In general, assimilation efficiency was higher when animals fed on seston from shallow compared to deep lakes, and was significantly correlated with growth rates, suggesting that shallow lakes have the best food resources in terms of digestibility and P availability. 3. Results also showed that all cladoceran species responded similarly to the resource gradient, with lower growth rates in deep lakes and higher growth rates in shallow lakes, although the strength of response (sensitivity) was different among the species tested. 4. The cladoceran Moina micrura was the most sensitive species, and also displayed the highest P‐content and maximal growth rate, a pattern consistent with the growth rate hypothesis. 5. However, seston C : P ratio and growth rates in the different resources did not correlate with the animals’ P‐contents, showing an uncoupling between RNA‐phosphorus demands for growth and seston food quality. 6. In conclusion, our results support the idea that digestion resistance in algae is a major constraint to cladocerans in natural plankton communities.     

Reconsideration of the Phylogenetic Positions of Five Peritrich Genera,Vorticella, Pseudovorticella,Zoothamnopsis, Zoothamnium,and Epicarchesium (Ciliophora,Peritrichia, Sessilida), Based on Small Subunit rRNA Gene Sequences

ABSTRACT. In order to re‐evaluate the systematics of sessilid peritrich ciliates, small subunit (SSU) rRNA gene sequences were determined for 12 species belonging to five genera: Vorticella, Pseudovorticella, Epicarchesium, Zoothamnium, and Zoothamnopsis. Phylogenetic trees were deduced using Bayesian inference, maximum parsimony, and maximum likelihood methods. The phylogenetic analyses suggest that (1) sessilids which have stalks with continuous myonemes that contract in a zig‐zag fashion form a separate clade from those which have stalks that contract independently and in a spiral fashion, supporting the separation of the family Zoothamniidae from the family Vorticellidae and (2) Epicarchesium and Pseudovorticella, both of which have reticulate silverline systems, are more closely related to each other than to other vorticellids, suggesting that differences in the silverline system (i.e. transverse vs. reticulate) may be the result of genuine evolutionary divergence among sessilid peritrichs. However, the newly sequenced Zoothamnopsis sinica, which has a reticulate silverline pattern, nests within the unresolved Zoothamnium species that have transverse silverline patterns. Thus, there were at least two evolutions of the reticulate silverline pattern character state from a plesiomorphic transverse state in the peritrichid ciliates. The molecular work demonstrates the genus Zoothamnium to be paraphyletic in relation to morphological studies, and suggests that Astylozoon, Opisthonecta, and Vorticella microstoma possibly share a SSU rRNA secondary structure in the helix E10‐1 region.     

Impacts of grazing and burning on spider assemblages in dry eucalypt forests of north‐eastern New South Wales,Australia

Abstract In the dry eucalypt forests of north‐eastern New South Wales, Australia, cattle grazing occurs at low intensities and is accompanied by frequent low‐intensity burning. This study investigated the combined effects of this management practice on the ground‐dwelling and arboreal (low vegetation) spider assemblages. Spiders were sampled at 49 sites representing a range of grazing intensities, using pitfall trapping, litter extraction and sweep sampling. A total of 237 spider morphospecies from 37 families were collected using this composite sampling strategy. The abundance, richness, composition and structure of spider assemblages in grazed and ungrazed forest sites were compared and related to a range of environmental variables. Spider assemblages responded to a range of environmental factors at the landscape, habitat and microhabitat scales. Forest type, spatial relationships and habitat variability at the site scale were more important in determining spider assemblages than localized low‐intensity grazing and burning. However, it is possible that a threshold intensity of grazing may exist, above which spiders respond to grazing and burning. Although low‐intensity grazing and burning may not affect spider assemblages below a threshold stocking rate, that stocking rate has yet to be established.     

Habitat selection by the swamp wallaby (Wallabia bicolor) in relation to diel period,food and shelter

Patterns of resource selection by animals may be influenced by sex, and often change over a 24‐h period. We used a dry sclerophyll landscape managed for commercial timber production to investigate the effects of sex and diel period on habitat selection by the swamp wallaby (Wallabia bicolor). We predicted that selection would be (i) affected by both sex and diel period; and (ii) positively related to lateral cover during the day, but to food resources at night. Non‐metric multidimentional scaling indicated that some of the available habitats differed markedly with respect to visibility (an indicator of lateral cover), fern cover, forb cover, wallaby density and a forage quality index, providing the basis for non‐random habitat selection. At the landscape scale, wallabies showed strong selection for 5‐year‐old regenerating sites, selected against 10‐year‐old regenerating sites and unharvested forest, and avoided recently harvested (3–10 months post‐harvest) sites completely. At the scale of individual home ranges, a pooled male and female sample demonstrated selection for unharvested forest over recently harvested sites during both diurnal and nocturnal periods. A separate analysis showed that both sex and diel period influenced the selection of 5‐ and 10‐year‐old sites and the surrounding unharvested forest. Using a novel approach, we demonstrated that diurnal habitat selection by both sexes was negatively correlated with visibility, representing stronger selection for areas with more lateral cover. Nocturnal selection by females was positively correlated with values of a forage quality index, but this was not the case for males. We hypothesise that the observed patterns of selection were driven by the need to find food and avoid predators, but were also affected by the different reproductive strategies of males and females. Our results demonstrate the importance of incorporating factors such as sex and diel period into analyses of habitat selection.     

Three new species of deep‐sea Gromia (Protista,Rhizaria) from the bathyal and abyssal Weddell Sea,Antarctica

We describe three new species of the genus Gromia from bathyal and abyssal depths in the Weddell Sea. The new species are characterized by a combination of morphological and molecular criteria. All three species possess a distinct oral capsule and a layer of ‘honeycomb membranes’, which form the inner part of the organic test wall. Both these features are typical of gromiids. Their identification as gromiids is confirmed by analyses of partial small subunit ribosomal DNA (SSU rDNA) gene sequences. Gromia marmorea sp. nov. is a rounded species with a prominent oral capsule and a characteristically mottled appearance. In Gromia melinus sp. nov. , the test surface exhibits a polygonal pattern of ridges, with a layer of clay particles coating the surface between the ridges. Gromia winnetoui sp. nov. represents an elongate morphotype in which the organic test is enclosed within an agglutinated case, a feature previously unknown in gromiids. Phylogenetic analysis using the maximum‐likelihood method revealed that all three species form distinct clades, reflecting the morphological differences among Weddell Sea species, as well as between deep‐water Southern Ocean Gromia and previously described gromiids. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 157 , 451–469.     

Experimental herbivory of native Australian macrophytes by the introduced Mozambique tilapia Oreochromis mossambicus

This study describes experimental herbivory and detritivory of three common native aquatic macrophyte species by the introduced Mozambique tilapia Oreochromis mossambicus (Peters) (Pisces: Cichlidae), and its physiological response to their consumption. There was a highly significant effect of fish herbivory on plant weight for each of the macrophyte species, but this effect was not influenced by any preference for periphyton. Despite the herbivory, there was a highly significant loss of fish body weight across all plant species and weight could only be maintained by supplementary feeding of a high protein fish flake. These results suggest that despite eating these plants, an alternative food resource may be needed for survival and may trigger trophic plasticity in O. mossambicus.     

The importance of passerine birds as tick hosts and in the transmission of Borrelia burgdorferi,the agent of Lyme disease: a case study from Scotland

Borrelia burgdorferi sensu lato (s.l.) is the causative agent of Lyme borreliosis, the most common tick‐borne zoonosis of humans in Europe and North America. Here, we assessed the relative importance of different passerine bird species as tick hosts and their contribution to the B. burgdorferi s.l. transmission cycle in a rural residential area in Scotland. We caught 1229 birds of 22 species during the tick‐questing season. On average, 29% carried larval ticks (0.8 larvae per individual) and 5% carried nymph ticks (0.06 nymphs per individual). All attached ticks tested were Ixodes ricinus. Using a nested‐PCR, we found that 20% of nymphs tested positive to B. burgdorferi s.l. and all these were of the genospecies Borrelia garinii. We identified two new bird species carrying infected nymphs: Eurasian Siskin Carduelis spinus and European Greenfinch Carduelis chloris. Ground‐foraging species were more important than arboreal species in hosting I. ricinus nymphs and B. burgdorferi s.l. Common Blackbirds Turdus merula were the most common hosts, with Song Thrushes Turdus philomelos, Dunnocks Prunella modularis, European Greenfinches and Chaffinches Fringilla coelebs also hosting high rates of infection.     

Determining uptake of 'non-labile' soil cadmium by Thlaspi caerulescens using isotopic dilution techniques

We assessed the ability of several populations of the metal-hyperaccumulator species, Thlaspi caerulescens , to mobilize non-labile cadmium in soils historically contaminated by Pb/Zn mine spoil or sewage sludge. Radio- labile Cd was determined chemically as an ' E -value', [Cd _E ], and biologically as an ' L -value', [Cd _L ]. For comparison, chloride-extractable Cd, [Cd_chlor], was also determined using 1 M CaCl₂ as a single-step soil extractant. Values of [Cd _L ] were measured for six populations of T. caerulescens that varied substantially in their ability to assimilate soil Cd, and a non-accumulator species with a similar growth habit, Lepidium heterophyllum . Seeds were sown in soil spiked with ¹⁰⁹Cd and grown for 9–12 wk in a controlled environment room. Values of [Cd _L ] were determined from the specific activity of ¹⁰⁹Cd and concentration of Cd in the plant leaves. For the six soils studied, [Cd _E ] ranged from 4.9 to 49% of total soil Cd [Cd_T]. Values of [Cd _L ] were, in general, in close agreement with both [Cd _E ] and [Cd_chlor] and substantially less than [Cd_T]. However, [Cd _L ] showed no correlation with the concentration of Cd in plant tissue, [Cd_shoot]. This suggests that, in the soils studied, T. caerulescens did not mobilize non-labile soil Cd by producing root exudates or altering rhizosphere pH. The results imply that there may be significant restrictions to metal bioavailability, even to hyperaccumulator species, in heavily contaminated soils in which a large proportion of the metal may be present in 'non-labile' forms.