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To test the hypothesis that the inclusion of the forage herbs chicory (Cichorium intybus L.) or plantain (Plantago lanceolata L.) or both in a grass–legume pasture mix improves persistence of sown vegetation and reduces weed and invertebrate pest ingress, a study was undertaken in 31–44 intensively managed pastures in each of five regions in New Zealand (Northland, Waikato, Bay of Plenty, Taranaki and Canterbury). The regions were stratified according to farm type (dairy or sheep and beef), pasture type (sown with grasses, legumes and herbs or grasses and legumes) and pasture age (young: 1–2 years old; medium: 3–4 years old; and old: 5+ years since sowing, n = 171). Sown species ground cover and emergence from the soil seed bank was greater, and unsown species ground cover and emergence lower, in sheep and beef pastures sown with herbs than without herbs (78% vs 68% ground cover, respectively, averaged over all pasture ages), but there was no difference between pasture types on dairy farms. Invert...

Experiments with artificial diets demonstrated that black field cricket (Teleogryllus commodus (Walker)) and Lepidogryllus sp. were highly responsive to presence of lolines in their diet-quantities of diet consumed declined exponentially with increasing loline concentration. Amount consumed by black field cricket and Lepidogryllus sp. on diet containing 5,600 µg/g lolines was only 8 and 2% relative to those on loline-free diet, respectively. Additional experiments with Festulolium seeds demonstrated that both cricket species predated heavily on endophyte-free seed but largely avoided Epichloë uncinata-infected seed. By 12 h, black field cricket had destroyed 98.8% of endophyte-free but only 24.8% of E. uncinata-infected, loline-containing seed. By 36 h, Lepidogryllus sp. crickets had destroyed 40% of endophyte-free but had not fed on E. uncinata-infected, loline-containing seed. Glasshouse experiments demonstrated this aversion to lolines greatly reduces the damage potential of black field cricket in E. uncinata-infected Festulolium. When microswards were sown with E. uncinata-infected Festulolium, seedling numbers were reduced 25-26%, and yields 29-40%, by black field crickets relative to microswards sown without insect infestation. This contrasts with 70-78% reduction in seedling numbers and 67-80% reduction in yields in microswards sown to either endophyte-free Festulolium, endophyte-free perennial ryegrass, or Epichloë festucae var. lolii-infected Festulolium. Yields of mature E. uncinata-infected Festulolium plants were not adversely affected by black field crickets, irrespective of the presence of the endophyte-free standard Festulolium sown as a companion. In contrast, yields of endophyte-free Festulolium, endophyte-free perennial ryegrass, and E. festucae var. lolii-infected Festulolium plants were reduced by 56-61% by crickets.

The South American weevil Listronotus bonariensis (Kuschel) is an important pest of pastures in New Zealand. As a component of management strategies for this pest, the South American parasitoid Microctonus hyperodae Loan (Hymenoptera: Braconidae) was released in northern New Zealand during 1991 as a biological control agent. Over the subsequent 5-6 yr, the reproductive biology of M. hyperodae and its relationship to, and effects on, the reproductive phenology and fitness of L. bonariensis were studied at three sites. M. hyperodae was first recovered in the field in December 1991. Subsequently, the incidence of parasitism in L. bonariensis increased to seasonal maxima of 75-90%. There was variable synchrony between parasitoid generations and the two generations of its host, leading to marked seasonal variation in rates of parasitism and parasitoid abundance. Despite marked inter-year variation, abundance of host adult and egg populations declined in the presence of parasitoids. Parasitized host females had lower ovarian maturity scores, had lower egg loads, and exhibited less investment in wing muscle development than females that had escaped parasitism. There was almost complete elimination of egg maturation in parasitized females and these hosts contributed little to population recruitment. Rate of buildup and seasonal maxima in parasitism, frequency of superparasitism, adult abundances, and wing muscle development in adult L. bonariensis varied among the three sites in a manner that was only partially related to climate differences across the 1.83° gradient of latitude. Site effects were weak to absent in measures of reproductive condition in L. bonariensis females.

A new Alaninema species associated with leaf-veined slugs (Athoracophoridae) in New Zealand is described and illustrated. Alaninema ngata n. sp. is characterised by a very long, cylindrical body, rounded head bearing two circles of cephalic papillae and amphids, shallow stoma with three bifid teeth, muscular, cylindroid pharynx lacking valves, enlarged, rounded cardia, nerve ring crossing the anterior intestine, excretory pore situated just posterior to nerve ring, monodelphic females with post-median vulva, thick-shelled eggs covered with prominent tubercles, monorchic males with short testis reflection, reduced number of male genital papillae, strong equal spicules and gubernaculum and tail tip bearing about 20 digitate processes. It is distinguished from A. venmansi and A. njoroensis by having a larger body size (mean 8.3 mm and 13.8 mm in males and females, respectively vs 8.2 mm females in A. venmansi and 4.8 mm and 5.5 mm in A. njoroensis males and females, respectively) and more anterior vulval position (V = 51.6 vs 70 and 66.4 in A. venmansi and A. njoroensis, respectively). Analyses of a partial 18S rDNA sequence indicate that A. ngata n. sp. is affiliated to the Panagrolaimorpha. The systematic position, host range and distribution of A. ngata n. sp. are discussed.

Habitat loss is one of the greatest threats to species persistence. Gauging the scale of this problem requires quantitative methods that can predict the number of extinctions resulting from habitat loss. For the past three decades, the species – area relationship, an empirical relationship between the number of species present in an area and the size of that area, has been this tool. However, it fails to incorporate threats to species aside from habitat loss and the heterogeneous distribution of these threats across habitats. Recent studies have improved species – area predictions by incorporating not only direct eff ects of area on richness, but also indirect eff ects of area (through area-mediated predator abundance), on prey species richness. We extend this work to test the hypotheses that the indirect eff ects of the multiple threats of grazing and trampling in addition to fragmentation will amplify the eff ect of area on species richness and that this eff ect will be greatest in zones closest to the fragment edge. Further, we test for species and population level eff ects of fragmentation and grazing, including the non-random pattern of species loss and the decline in population sizes. We test our hypotheses with a fi eld study of land snail richness in fragments with and without the additional threats of grazing and trampling. Our study supports the hypotheses that fragments with multiple threats in addition to habitat loss harbour fewer species than fragments without these threats, and that this eff ect is non-uniform across fragments, populations and species.

Hydrocarbon spills on Antarctic soils occur mainly near settlements where fuel is stored and aircraft and vehicles are refuelled. To investigate those factors that may preclude hexadecane mineralization activity in long-term hydrocarbon-contaminated soils from the Ross Sea Region, samples were collected from Scott Base, the site of former bases (Cape Evans, Marble Point, Vanda Station), and two oil spill sites in the Wright Valley (Bull Pass and Loop Moraine). The soils had low levels of nitrogen ( 24) reflecting hydrocarbon contamination. Following soil water adjustment to 10% (v/w), the influence of nutrient addition (250 mg/kg N added as monoammonium phosphate) and inoculation (spiking with Antarctic soil containing high numbers of hydrocarbon degraders) as required on hexadecane mineralization activity was determined. Hexadecane mineralization activity occurred in contaminated soils from Marble Point, Cape Evans and one sample from Vanda Station without nutrient addition. In contrast soils from Scott Base, Cape Evans, another sample from Vanda Station and Loop Moraine required nutrients, whereas Bull Pass soil required inoculation and nutrients before hexadecane mineralization proceeded. Hydrocarbon degrader numbers were highest in coastal soils from Scott Base and Marble Point (10 7 per gram) and less prevalent in inland soils from Wright Valley ( 5 per gram). The bacterial community structure of the soils differed between sites, but soils from the same sites tended to cluster together more closely, except for those from Vanda Station. Addition of nutrients did not cause large shifts in the soil bacterial communities. Results from this study indicate that hydrocarbon degradation may occur at some sites in summer when water is available. Long-term hydrocarbon-contaminated Antarctic soils may provide a valuable resource of hydrocarbon-degrading bacteria that can serve as inocula for more recent oil spills on land.

Four pedons on each of four drift sheets in the Lake Wellman area of the Darwin Mountains were sampled for chemical and microbial analyses. The four drifts, Hatherton, Britannia, Danum, and Isca, ranged from early Holocene (10 ka) to mid-Quaternary (c. 900 ka). The soil properties of weathering stage, salt stage, and depths of staining, visible salts, ghosts, and coherence increase with drift age. The landforms contain primarily high-centred polygons with windblown snow in the troughs. The soils are dominantly complexes of Typic Haplorthels and Typic Haploturbels. The soils were dry and alkaline with low levels of organic carbon, nitrogen and phosphorus. Electrical conductivity was high accompanied by high levels of water soluble anions and cations (especially calcium and sulphate in older soils). Soil microbial biomass, measured as phospholipid fatty acids, and numbers of culturable heterotrophic microbes, were low, with highest levels detected in less developed soils from the Hatherton drift. The microbial community structure of the Hatherton soil also differed from that of the Britannia, Danum and Isca soils. Ordination revealed the soil microbial community structure was influenced by soil development and organic carbon.

For many taxonomic groups, sparse information on the spatial distribution of biodiversity limits our capacity toanswer a variety of theoretical and applied ecological questions. Modelling community-level attributes (a- andb-diversity) over space can help overcome this shortfall in our knowledge, yet individually, predictions of a-or b-diversity have their limitations. In this study, we present a novel approach to combining models of a- andb-diversity, with sparse survey data, to predict the community composition for all sites in a region. We appliedour new approach to predict land snail community composition across New Zealand. As we demonstrate, thesepredictions of metacommunity composition have diverse potential applications, including predicting c-diversityfor any set of sites, identifying target areas for conservation reserves, locating priority areas for future ecologicalsurveys, generating realistic compositional data for metacommunity models and simultaneously predicting thedistribution of all species in a taxon consistent with known community diversity patterns.KeywordsAlpha, beta, community, dissimilarity, diversity, gamma, metacommunity, macroecology, richness, snail.Ecology Letters (2011) 14: 1043–1051

The Last Glacial Maximum (LGM) severely restricted forest ecosystems on NewZealand’s South Island, but the extent of LGM distribution for forest species is stillpoorly understood. We used mitochondrial DNA phylogeography (COI) and ecologicalniche modelling (ENM) to identify LGM refugia for the mycophagous beetle Agyrtodeslabralis (Leiodidae), a forest edge species widely distributed in the South Island. Boththe phylogenetic analyses and the ENM indicate that A. labralis refuged in Kaikoura,Nelson, and along much of the South Island’s west coast. Phylogeography of this speciesindicates that recolonization of the largely deforested east and southeast South Islandoccurred in a west–east direction, with populations moving through the Southern Alps,and that the northern refugia participated little in interglacial population expansion. Thiscontradicts published studies of other New Zealand species, in which recolonizationoccurs in a north–south fashion from many of the same refugia.Keywords: ecological niche model, fungus beetle, glacial refugia, Leiodidae, Maxent, PleistoceneReceived 26 July 2009; revision received 1 October 2009; accepted 3 October 2009

Aim Using New Zealand land snails as a case study, we evaluated recent spatial modelling approaches for the analysis of diversity in species-rich invertebrate groups. Applications and prospects for improved conservation assessment were investigated. Location New Zealand. Methods The study used a spatially extensive and taxonomically comprehensive, plot-based dataset on community structure in New Zealand land snails. Generalized regression analysis and spatial prediction (GRASP) was used to model and predict species richness as a function of environmental variables (including aspects of climate, soils and vegetation). Generalized dissimilarity modelling (GDM) was used to model turnover in species composition in relation to environmental and geographical distances, and to assess community similarity and the representativeness of the reserve network. Results Observed land snail richness in 20 × 20 m plots ranged from 1 to 74 (mean 17.5). The GRASP model explained a modest 27% of the variation in richness. The GDM model explained 57% of the variation in species turnover and indicated approximately equal amounts related to environmental (Cody’s beta diversity) and geographical distance (Cody’s gamma diversity). Temperature and moisture were the most important environmental variables. Results indicate that snail distributions are not only sorted by environment but are also strongly influenced by historical effects consistent with those expected of poorly dispersing taxa that have persisted in refugia during past climatic change. The GDM model enabled spatial classifications of snail communities, highlighting diverse communities in heterogeneous regions, such as the South Island mountains, and also enabled continuous depictions of community similarity and adequacy of New Zealand’s protected natural areas network. Main conclusions The GRASP and GDM analyses allowed us to model and depict spatial patterns of diversity in land snail communities involving 845 species, and produce community classifications and estimates of community similarity. These tools advance conservation assessment in species-rich groups, but require further conceptual and methodological development.

In many agricultural landscapes, significant biodiversity gains can be made by improving the ecological condition of degraded remnants of semi‐natural habitat. Recent emphasis has been on the level of management intervention required to initiate vegetation recovery in small forest remnants, but no comparable emphasis has been placed on benefits for invertebrate communities. In the Waikato region, New Zealand, we tested the effects of livestock exclusion, mammalian pest control, and their interaction, on leaf‐litter invertebrate communities in 30 forest remnants, using a space‐for‐time substitution approach. A total of 87 376 invertebrates were extracted from 964 leaf‐litter samples. Invertebrate density was an order of magnitude lower in remnants than in nearby large forest reserves. For key taxa, such as Diplopoda, Isopoda, Coleoptera and Mollusca, 10‐ to 100‐fold lower densities were recorded in remnants with no pest control, particularly where livestock were not excluded. By contrast, other ta...

A recently discovered and threatened large land snail, Powelliphanta augusta n. sp. is described from Mt Augustus on the western scarp of the Stockton Plateau, North Westland. On shell characters it is readily distinguishable from all other Powelliphanta, with narrow red spiral lines underlying irregular and variable dark reddish‐brown axial bands, a small adult size and sculptured dorsal surface. Differences in the shell morphology of P. augusta, its closest phylogenetic relative P. lignaria, and its closest geographic neighbour P. patrickensis, were investigated using principal components analysis. P. lignaria is much larger than either P. augusta or P. patrickensis, and there are significant differences in shell shape between the three taxa. Earlier mitochondrial DNA sequencing data, which supported specific status of P. augusta, is discussed. The snail's only habitat on the Mt Augustus ridgeline is described, but most has now been removed by coal mining. Snails salvaged before the destruction...

Soils of the Ross Sea region of Antarctica are among the least diverse ecosystems on earth. Few plants and animals have colonised these soils, but bacteria are distributed throughout. We compared the bacterial communities of soil samples from 5 locations in the Ross Sea region, representing different soil taxonomic groups, using culture-independent methods. Four mineral soils and one ornithogenic soil were analysed. Organic C, Total N, and moisture of the mineral soils were low, total P and EC were variable. In contrast, the water content, organic C, and Total N and P, and EC of the ornithogenic soil were 10–100 fold higher than the mineral soils. Soil pH was alkaline. One thousand and fourteen clones were screened by restriction fragment length polymorphisms. While the majority of the ribotypes were rare, some were dominant and occurred ≥ 3 times in the respective clone libraries. The dominant ribotypes were sequenced and shown to be phylogenetically affiliated with the phyla Acidobacteria , Actinobacteria , Bacteroidetes , Proteobacteria , Deinococcus-Thermus , Firmicutes , Cyanobacteria and Candidate TM7. The closest relatives of the clones retrieved from the mineral soils were uncultured environmental clones or bacterial isolates not assigned to recognised genera. In contrast, the clones from the ornithogenic soil were most closely related to the endospore-formers; Oceanobacillus profundus , Clostridium acidiurici and Sporosarcina aquimarina. We found no convincing evidence that the Haplorthels from Scott Base, Marble Point, and Cape Hallett, have bacterial communities structured fundamentally different from those of Anhyorthels from Bull Pass and Vanda. Nonetheless the composition of the communities did vary substantially and were readily discriminated by multivariate statistics on the basis of soil properties.

Human‐assisted spread of species poses a major challenge to border security agencies. Ideally, limited resources need to be targeted at species posing the most risk. Climate matching is an important component of assessing risk but often little or no biological information is available to enable detailed modelling. To assess if distribution records alone provide useful establishment predictions we compare the climate in New Zealand and its outlying islands to that found in the current native and introduced ranges of 12 tramp ant species, three of which are already established in New Zealand, using the climate module of BIOSECURE, a risk assessment tool. Eleven species showed a similar general pattern, with mean annual temperature being the climatic variable with the least overlap between the estimated realised niche and the predicted New Zealand niche. The twelfth species, of temperate origin, is less restricted by temperature, but much of New Zealand may have too high a rainfall. The present and ...

Although there has been much recent interest in the effect of litter mixing on decomposition processes, much remains unknown about how litter mixing and diversity affects the abundance and diversity of decomposer organisms. We conducted a litter mixing experiment using litterbags in a New Zealand rainforest, in which treatments consisted of litter monocultures of each of 8 forest canopy and understory plant species, as well as mixtures of 2, 4 and 8 species. We found litter mixing to have little effect on net decomposition rates after either 279 or 658 days, and for each species decomposition rates in mixture treatments were the same as in monoculture. Litter species identity had important effects on litter microfauna, mesofauna and macrofauna, with different litter types promoting different subsets of the fauna. Litter mixing had few effects on densities of mesofauna and macrofauna, but did have some important effects on components of the microfauna, notably microbe-feeding and predatory nematodes. At day 279, litter mixing also consistently reduced the ratio of bacterial-feeding to microbe-feeding (bacterial-feeding+fungal-feeding) nematodes, pointing to mixing causing a significant switch from the bacterial-based to the fungal-based energy channel. Litter mixing sometimes influenced the community composition and diversity of nematodes and macrofauna, but effects of litter mixing on diversity were not necessarily positive, and were much weaker than effects of litter species identity on diversity. We conclude that litter mixing effects on the abundance and diversity of decomposer biota, when they occur, are likely to be of secondary and generally minor significance when compared to the effects of litter species identity and composition.

Five experiments investigated the importance of herbivory by Sminthurus viridis in structuring botanical composition in developing grasslands, and how these effects may be modified by diversity in collembolan and plant species. Differential susceptibility to S. viridis feeding was demonstrated in 23 dicotyledonous and three monocotyledonous plants assayed as seedlings at the first true leaf stage. The composition of seedling communities developing from natural and artificially constructed soil seed banks varied with the level of S. viridis infestation, with plant species least susceptible to herbivory making the greatest contribution to plant biomass. The combined effect of herbivory by S. viridis and Bourletiella hortensis on Trifolium repens biomass was shown to be less than the effect of S. viridis alone, indicating competitive interference. The adverse effects of herbivory by S. viridis on T. repens biomass was reduced by increased diversity of plants growing in association with the legume, and the presence of four non-herbivorous arthropleonan Collembola. S. viridis was shown to reduce seedling numbers, species diversity and biomass in communities developing from the soil seed bank, but the presence of non-herbivorous arthropleonan species reduced the effect of S. viridis. The experiments demonstrate the potential for herbivory by S. viridis to significantly alter species composition in developing grassland communities. However, interactions with collembolan and plant species profoundly modified S. viridis herbivory impacts, either by reducing feeding intensity or enhancing plant growth. These results highlight the fact that data from simple, synthetic systems may be poor predictors of herbivory impacts under field conditions where more complex species interactions occur.

The parasitoid Microctonus hyperodae Loan (Hymenoptera: Braconidae) was introduced into New Zealand to control the weevil Listronotus bonariensis (Kuschel) (Coleoptera: Curculionidae), a major pest of graminaceous plants. Four experiments were conducted to investigate the effects of various pesticides that are commonly used in the pastoral environments of L. bonariensis and M. hyperodae . To pical applications of aqueous solutions prepared from commercial formulations of five herbicides were not toxic, but the surfactant Silwett L-77 increased M. hyperodae mortality relative to the watertreated controls. Laboratory assays showed that M. hyperodae adults were susceptible to chlorpyrifos residues on pasture foliage following application of the insecticide to field plots at ≥ 5 g a.i. ha − 1 . Maintenance of L. bonariensis on ryegrass in the laboratory showed that treatment of the food plants with chlorpyrifos at ≥ 96 g a.i. ha − 1 reduced L. bonariensis survivorship and ultimately reduced M. hyperodae prepupal emergence from those hosts. Initially, mortalities of non-parasitized L. bonariensis were significantly greater than for parasitized L. bonariensis . Maintenance of parasitized L. bonariensis on diflubenzuron-treated ryegrass plants arrested M. hyperodae larval development in the host and ultimately reduced prepupal emergence of M . hyperodae from those hosts. Despite the arrested development of M. hyperodae , the mortality of L. bonariensis hosts was increased. Adult M. hyperodae successfully reared from hosts maintained on diflubenzuron (12.5 g a.i. ha − 1 ) treated food plants had reduced reproductive potential. The consequences of pasture management strategies that employ pesticides are discussed in relation to biocontrol of L. bonariensis by M . hyperodae .

Summary 1There is increasing awareness that similar suites of plant traits may govern foliage palatability and litter decomposability, but whether there is an association between the response of vegetation to herbivory and litter decomposition rates across plant species remains unexplored. 2We collected 141 samples of litter from 59 understorey and 18 canopy tree species from a total of 28 sites under natural forest throughout New Zealand. We assessed whether variables related to decomposition and quality of litter of the understorey species showed a statistical relationship with the response of vegetation density (assessed using a pole-intercept method) of the same species at the same locations to browsing by deer and goats. Decomposition and nutrient-loss data from litter were obtained using standardized laboratory bioassays. 3There was a significant positive correlation between litter decomposition rate and the extent to which vegetation density was reduced by browsing mammals (r = 0·488, P

We established a greenhouse experiment based on replicated mini-ecosystems to evaluate the effects of defoliation intensity on soil food-web properties in grasslands. Plant communities, composed of white clover (Trifolium repens), perennial ryegrass (Lolium perenne) and plantain (Plantago lanceolata) with well-established root and shoot systems, were subjected to five defoliation intensity treatments: no trimming (defoliation intensity 0, or DI 0), and trimming of all plant material to 35 cm (Dl 1), 25 cm (DI 2), 15 cm (DI 3) and 10 cm (Dl 4) above soil surface every second week for 14 weeks. Intensification of defoliation reduced shoot production and standing shoot and root mass of plant communities but increased their root to shoot ratio. Soil microbial activity and biomass decreased with intensification of defoliation. Concentrations of NO 3 -N in soil steadily increased with intensifying defoliation, whereas NH 4 -N concentrations did not vary between treatments. Numbers of microbi-detritivorous enchytraeids, bacterial-feeding rotifers and bacterial-feeding nematodes steadily increased with intensifying defoliation, while the abundance of fungal-feeding nematodes was significantly enhanced only in Dl 3 and DI 4 relative to DI 0. The abundance of herbivorous nematodes per unit soil mass was lower in DI 3 and DI 4 than in DI 0, Dl I and DI 2. but when calculated per unit root mass, their abundance tended to increase with defoliation intensity, The abundance of omnivorous and predatory nematodes appeared to be highest in the most intensely defoliated systems The ratio of abundance of fungal-feeding nematodes to that of bacterial-feeding nematodes was not significantly affected by defoliation intensity. The results s infer that defoliation intensity may significantly alter the structure of soil food webs in grasslands, and that defoliation per se is able to induce patterns observed in grazing studies in the field. The results did not support hypotheses that defoliation per se would cause a shift between the bacterial-based and fungal-based energy channels in the decomposer food web, or that herbivore and detritivore densities in soil would be highest under intermediate defoliation. Furthermore, our data for microbes and microbial feeders implies that the effects of defoliation intensity on soil food-web structure may depend on the duration of defoliation and are therefore likely to be dynamic rather than constant in nature.

This is the first published invertebrate survey focusing on a low‐altitude shrubland community in New Zealand. Invertebrates were collected from a remnant native shrubland (450 m) protected by the Brookdale Conservation Covenant, Rock and Pillar Range, Otago, New Zealand in late summer/autumn 1999. Sampling was carried out by beating 30 randomly chosen shrubs of each of two native species: Olearia bullata H. D. Wilson & Garnock‐Jones (Asteraceae) and Coprosma propinqua A. Cunn. (Rubiaceae). Fifty pitfall traps were also set under the same shrubs and on nearby open patches of exotic grassland. Three Phyla, six Classes, 25 Orders and approximately 280 species were recorded. An annotated list of taxa is presented, and plant/host associations plus other observations on the fauna are discussed. Approximately 90% of the identified species were endemic, emphasising the importance of such remnant habitats for the protection of New Zealand's biodiversity.

Although factors affecting plant growth and plant carbon/nutrient balance – e.g., light availability and defoliation by herbivores – may also propagate changes in below-ground food webs, few studies have aimed at linking the above-ground and below-ground effects. We established a 29-week laboratory experiment (∼one growing season) using autotrophic microcosms to study the effects of light and defoliation on plant growth, plant carbon/nutrient balance, soil inorganic N content, and microbial activity and biomass in soil. Each microcosm contained three substrate layers – mineral soil, humus and plant litter – and one Nothofagus solandri var. cliffortioides seedling. The experiment constituted of the presence or absence of two treatments in a full factorial design: shading (50% decrease in light) and artificial defoliation (approximately 50% decrease in leaf area in the beginning of the growing season). At the end of the experiment a range of above-ground and below-ground properties were measured. The shading treatment reduced root and shoot mass, root/shoot ratio and leaf production of the seedlings, while the defoliation treatment significantly decreased leaf mass only. Leaf C and N content were not affected by either treatment. Shading increased NO 3–N concentration and decreased microbial biomass in humus, while defoliation did not significantly affect inorganic N or microbes in humus. The results show that plant responses to above-ground treatments have effects which propagate below ground, and that rather straightforward mechanisms may link above-ground and below-ground effects. The shading treatment, which reduced overall seedling growth and thus below-ground N use and C allocation, also led to changes in humus N content and microbial biomass, whereas defoliation, which did not affect overall growth, did not influence these below-ground properties. The study also shows the carbon/nutrient balance of N. solandri var. cliffortioides seedlings to be highly invariant to both shading and defoliation.

While there has been a rapidly increasing research effort focused on understanding whether and how composition and richness of species and functional groups may determine ecosystem properties, much remains unknown about how these community attributes affect the dynamic properties of ecosystems. We conducted an experiment in 540 mini-ecosystems in glasshouse conditions, using an experimental design previously shown to be appropriate for testing for functional group richness and composition effects in ecosystems. Artificial communities representing 12 different above-ground community structures were assembled. These included treatments consisting of monoculture and two- and four-species mixtures from a pool of four plant species; each plant species represented a different functional group. Additional treatments included two herbivore species, either singly or in mixture, and with or without top predators. These experimental units were then either subjected to an experimentally imposed disturbance (drought) for 40 d or left undisturbed. Community composition and drought both had important effects on plant productivity and biomass, and on several below-ground chemical and biological properties, including those linked to the functioning of the decomposer subsystem. Many of these compositional effects were due to effects both of plant and of herbivore species. Plant functional group richness also exerted positive effects on plant biomass and productivity, but not on any of the below-ground properties. Above-ground composition also had important effects on the response of below-ground properties to drought and thus influenced ecosystem stability (resistance); effects of composition on drought resistance of above-ground plant response variables and soil chemical properties were weaker and less consistent. Despite the positive effects of plant functional group richness on some ecosystem properties, there was no effect of richness on the resistance of any of the ecosystem properties we considered. Although herbivores had detectable effects on the resistance of some ecosystem properties, there were no effects of the mixed herbivore species treatment on resistance relative to the single species herbivore treatments. Increasing above-ground food chain length from zero to three trophic levels did not have any consistent effect on the stability of ecosystem properties. There was no evidence of either above-ground composition or functional group richness affecting the recovery rate of ecosystem properties from drought and hence ecosystem resilience. Our data collectively point to the role of composition (identity of functional group), but not functional group richness, in determining the stability (resistance to disturbance) of ecosystem properties, and indicates that the nature of the above-ground community can be an important determinant of the consistency of delivery of ecosystem services.

Summary Aim The New Zealand terrestrial mollusc fauna is among the most speciose in the world, with often remarkably high richness at lowland forest sites. We sought to elucidate general explanations for patterns of richness in terrestrial mollusc communities by analysis of species coexistence and habitat relationships within a New Zealand district fauna. Location Pukeamaru Ecological District, eastern North Island, New Zealand. Methods We sampled molluscs using qualitative methods at twenty-three sites and quantitatively by frame sampling of scrubland-forest floor litter at sixteen of these sites and analysed patterns of species richness and turnover in relation to regional species pools and local habitat attributes. We then tested for nonrandom assemblage of taxa along diversity and habitat gradients. Results Ninety-four indigenous mollusc species were recorded from a district fauna estimated at 102 indigenous species: only two species were endemic. From the presumptive geological history of the district, the low endemism, and Brooks parsimony and indicator species analyses of faunal relationships, the communities were indicated to have resulted by accumulation of colonists from other New Zealand districts since the Miocene. Richness ranged from two or three indigenous species in dune habitats to fifty-nine species in a floristically rich forest. Beta diversity was high and site occupancy per species was low, indicating communities structured by successive replacement of ecological equivalents. Sites differing in vegetation had characteristic species assemblages, indicating a degree of habitat specialization. Canonical correspondence analysis indicated that canopy tree species, canopy height, floristic diversity, altitude, litter mass, and litter pH were important determinants of species assemblage in scrubland and forest. Richness was strongly associated with site floristic diversity and, for litter-dwelling species, the pH of litter substrate. High richness occurred at those sites supporting molluscs in high abundance. Shell-shape distributions were essentially Cainian unimodal, with communities dominated by snail species with subglobose to discoidal shells. Mean and variance of shell size increased with mollusc species richness and floristic diversity at sites, indicating dominance of communities by small-shelled species at early successional or floristically poor sites, and increased richness resulting from addition of larger snails into vacant niches. Shifts in shell form were associated with sympatry in several congeneric taxa. Main conclusions The underdispersion of shell shape, relative to faunas elsewhere in the world, indicates that community structure in New Zealand land snail faunas has been constrained by limited phylogenetic diversity and/or by convergence upon successful adaptations. The remarkably high richness that characterizes these communities indicates special conditions allow coexistence of numerous species. The relationship between floristic diversity at sites and the richness, diversity, and shell-size distributions of the molluscs suggests assemblages structured around niche partitioning among competing species. While there is an element of congruence between vegetation and mollusc pattern, this study indicates that assembly rules will be defined, and spatial pattern predicted, only through a better understanding of the linkage between regional species pool, organism traits, environment, and local community assemblage.

1 The development of general principles regarding biotic interactions involving plants, or plant species effects in ecosystems, is best achieved through simultaneous evaluation of several species. We utilized a comparative approach involving 20 dicotyledonous herbaceous species, to explore possible relationships between several plant ecophysiological traits and plant litter decomposition, interactions involving competition and herbivory, and plant species effects on soil properties. 2 Decomposition rates of plant stem and leaf litter were negatively related to plant mass, time until flowering and vegetative growth rate, and positively related to stem nitrogen content. Root decomposition was also related to several traits. Multiple regression relationships showed that 74% and 84% of the variation across species for stem and root litter decomposition, respectively, could be predicted by plant traits; this suggests that plant traits may be powerful predictors of decomposition and have potential as alternative predictors to the litter quality characteristics that previous studies have concentrated on. 3 Palatability of both seedlings and leaf discs by the invertebrate herbivores Deroceras reticulatum and Listronotus bonariensis were frequently related to plant traits. Those traits that showed the strongest relationships with the palatability data included various vegetative growth characteristics and (for the leaf disc data) nitrogen concentrations of flowering plant stems. 4 Competitive effects of the dicotyledonous species against a phytometer species, the grass Lolium perenne, were negatively related to leaf nitrogen concentration, and multiple regression relationships involving this trait in combination with others explained over 50% of the variation across species. The competitive response of both plant mass and total seed production to L. perenne was poorly related to plant traits. 5 The effects of plant species on soil properties including microbial biomass and activity, pH, nitrate concentration and total nitrogen were often closely related to various plant traits. Multiple regression relationships revealed that combinations of several traits were often important in determining these effects; the strongest relationships found were for effects of senescent plants on soil respiration and for the effects of flowering plants on soil nitrate. Plant traits were therefore clearly important in determining plant species effects on soils. 6 Our study emphasizes the importance of plant traits in understanding (and predicting) species interactions and effects in communities and ecosystems, and shows that properties considered at the whole plant level have the potential to manifest themselves over much larger scales. We therefore conclude that there are clear linkages between plant ecophysiological traits, biotic interactions involving plants, and ecosystem level properties and processes.

The parasitoids Microctonus aethiopoides Loan and Microctonus hyperodae Loan (Hymenoptera: Braconidae) have been introduced into New Zealand to control the adult stage of the forage pests Sitona discoideus and Listronotus bonariensis (Coleoptera: Curculionidae), respectively. These parasitoids have been chosen to examine the environmental impact of introduced biological control agents on native and non‐target species. A survey of the Curculionoidea fauna of pasture, lucerne, and modified native grassland in parts of the southern South Island, Canterbury, and the northern North Island of New Zealand, where these parasitoids are present, was carried out in order to identify Curculionoidea with taxonomic and ecological affinities with the target hosts, and hence, potential non‐target hosts. At a total of 155 sites, 85 species of Curculionoidea were identified, of which 64 (75%) were native species. Three tribes in the subfamily Brachycerinae (broad‐nosed weevils) accounted for 61 species, and 7 trib...

Laboratory host specificity of 2 biological control agents, already introduced in New Zealand, was compared with actual field parasitism. The parasitoids were Microctonus aethiopoides Loan and Microctonus hyperodae Loan, braconids imported to control the curculionid forage pests Sitona discoideus Gyllenhal and Listronotus bonariensis (Kuschel), respectively. The nontarget weevil species tested included native, introduced, and beneficial species. M. aethiopoides oviposited in 11 of the 12 species to which it was exposed and successfully parasitized 9 species. M. hyperodae oviposited in 5 of the 11 species to which it was exposed and developed successfully in 4 species. Higher percentage parasitism was recorded with M. aethiopoides than with M. hyperodae . Field collections of weevils from Otago, Canterbury, and Waikato indicated that 10 New Zealand native species and 3 other nontarget species, including the weed biological control agent Rhinocyllus conicus (Froehlich), were parasitized by M. aethiopoides . M. hyperodae has been found parasitizing 1 native species, as well as Sitona lepidus Gyllenhal, which was accidentally introduced to New Zealand recently. In nontarget species, parasitism levels in the field of >70% have been recorded for M. aethiopoides and

Laboratory experiments were done to examine the effect of ryegrass infection by the endophytic fungus Acremonium lolii Latch, Christensen & Samuels (Ascomycetes: Clavicipitaceae) on Microctonus hyperodae Loan, a parasitoid of Listronotus bonariensis (Kuschel). Infection of ryegrass did not influence the rate of parasitism in weevils confined with naive parasitoids to individual plants and to ryegrass swards in small plots. In contrast, M. hyperodae with prior experience of hosts feeding on endophyte infected grass subsequently were more efficient in parasitizing hosts on this diet. Feeding on individual plants and small plot swards by weevil for 3–4 d increased subsequent parasitoid attack rate.

A new genus and a new species are described for nematodes removed from the genital tract of Athoracophorus bitentaculatus in New Zealand. Hugotdiplogaster neozelandia n. gen., n. sp. is different from other genera in the Diplogasteridae by the shape of the stoma and by possessing a very robust and large gubernaculum. It is the first time that a diplogasterid has been reported associated with a mollusc. The localisation of the nematode in the genital tract of slugs raises questions about its life cycle and effect on the host.

For many taxonomic groups, sparse information on the spatial distribution of biodiversity limits our capacity to answer a variety of theoretical and applied ecological questions. Modelling community-level attributes (α- and β-diversity) over space can help overcome this shortfall in our knowledge, yet individually, predictions of α- or β-diversity have their limitations. In this study, we present a novel approach to combining models of α- and β-diversity, with sparse survey data, to predict the community composition for all sites in a region. We applied our new approach to predict land snail community composition across New Zealand. As we demonstrate, these predictions of metacommunity composition have diverse potential applications, including predicting γ-diversity for any set of sites, identifying target areas for conservation reserves, locating priority areas for future ecological surveys, generating realistic compositional data for metacommunity models and simultaneously predicting the distribution of all species in a taxon consistent with known community diversity patterns.

The Last Glacial Maximum (LGM) severely restricted forest ecosystems on New Zealand's South Island, but the extent of LGM distribution for forest species is still poorly understood. We used mitochondrial DNA phylogeography (COI) and ecological niche modelling (ENM) to identify LGM refugia for the mycophagous beetle Agyrtodes labralis (Leiodidae), a forest edge species widely distributed in the South Island. Both the phylogenetic analyses and the ENM indicate that A. labralis refuged in Kaikoura, Nelson, and along much of the South Island's west coast. Phylogeography of this species indicates that recolonization of the largely deforested east and southeast South Island occurred in a west-east direction, with populations moving through the Southern Alps, and that the northern refugia participated little in interglacial population expansion. This contradicts published studies of other New Zealand species, in which recolonization occurs in a north-south fashion from many of the same refugia.

Life cycles of the slugs Deroceras reticulatum and Arion intermedius were studied over a 2.5-year period in a sheep-grazed pasture in the Kaimai Range, New Zealand. D. reticulatum approximated a bivoltine phenology, with intervals between consecutive generations ranging from 4 to 7 months and maximum life span from 8 to 12 months. Egg laying occurred in autumn and spring-early summer. Periods of egg hatching were followed by a phase of juvenile growth characterised by linear increase in the logarithm of population mean body weights. This was followed by a phase of reproductive activity. A. intermedius had an annual life cycle. Eggs laid during late summer and autumn hatched during autumn-winter. Immature slugs were characterised by a period of low growth rate during winter followed by a period of rapid growth culminating in reproductive maturity. In both species, the phase of rapid growth rate was associated with enlargement of the hermaphrodite gland. The size of the hermaphrodite gland relative to body weight reached a maximum during the spermatozoon stage, generally coincident with maximum body weight. During the reproductive period of D. reticulatum and A. intermedius the body weight remained relatively stable but the hermaphrodite gland became progressively smaller as the slugs approached the post-reproductive stage. The albumen gland attained maximum weight at the oocyte stage of the hermaphrodite gland, at the onset of reproductive activity. The pasture, strongly dominated by grasses, was heterogenous in slope, vegetation and treading/grazing influences from sheep due to stratification of the habitat into contour tracks, associated with sheep movement around the slope contours, and intertrack areas. High population densities of D. reticulatum were associated with the track edges (kerbs) and the upper intertrack slopes, while densities of A. intermedius were highest for the intertrack slopes. Low densities of both species occurred on the tracks and at sheep campsites. Analysis of cohort life tables indicated that these dispersion patterns were important in the dynamics of the populations. The stage mortality contributing most to the variance in reproductive populations of both species was that between hatching and onset of reproduction; most of the mortality occurred shortly after hatching on south-facing slopes, but tended to occur later in the life cycle on north slopes. For D. reticulatum this mortality was inversely related to initial density and associated, at least in part, with predation by carabids and birds and with treading by sheep. Density relationships, both at whole plot and plot stratum level, indicated that variations in natality had a stabilizing influence on D. reticulatum populations.

We review the diversity and conservation status of Placostylinae, land snails endemic to the western Pacific. Their narrow-range endemism, large size and associated vulnerability, consumptive exploitation by people, and habitat loss and degradation (inclusive of invasive predators) threaten their survival. There has been considerable attention from conservation biologists in New Caledonia, Lord Howe Island and New Zealand aimed at species recovery. Nonetheless, only on uninhabited, pest-free islands do these native snails persist in high numbers, and these remaining ‘sanctuaries’ are dependent on biosecurity vigilance. For other populations in New Caledonia, Lord Howe Island and New Zealand, the benefits of control of invasive mesopredators have been demonstrated, but it remains unclear if long-term persistence of Placostylinae can be achieved in degraded landscapes that continue to be subject to anthropogenic pressures. For species in Fiji, Vanuatu, and the Solomon Islands – the centre of Placostylinae diversity – their conservation status is not known with any certainty due to lack of basic data on range and population trends. Recent IUCN Red List assessments indicate a high level of extinction risk among Fijian species due to narrow geographic range coupled with decline in habitat extent and quality. Further inventory and ecological work is urgently needed in the Solomon Islands and Vanuatu to enable assessment of extinction risk and identify threatening processes. We identify four priority areas for advancing the conservation of Placostylinae, especially in Melanesia, and discuss the most pressing methodological issues. Molecular phylogenetic analyses are needed to provide an evolutionary framework for taxonomic revision and to underpin development of both conservation policy and species recovery plans.

Where conservation status of island non-marine molluscs is known, snails tend to be one of the most threatened faunal groups. However, published information regarding island gastropod conservation status, diversity and endemism is frequently unavailable despite the importance of this information for the formulation of biodiversity action plans and conservation strategy. Fiji, for example, has a diverse native land snail fauna (>240 species) with an endemism level of ~80%, but only within the last few years has any information about any of these species been available to the national biodiversity reporting repository. For one lineage in particular, members of the tree snail family Partulidae, with four endemic Fiji Island species, the conservation status of the group has never been assessed. However, based on the alarming extinction rates documented in partulid snail species on other Pacific Islands, information about the occurrence and status of these taxa is urgently needed for Fiji’s biodiversity action plan. To redress this information void, we formulated the Fijian Partulid Tree Snail Project, consisting of five components: (1) raising awareness; (2) locating populations and monitoring population trends; (3) elucidating patterns of genetic diversity; (4) creating action partnerships; and (5) conducting disturbance gradient analyses. The overall goal was to characterise mechanisms leading to persistence of partulids in the face of increasing anthropogenic disturbance. In the initial stages of this project, existing information on Fiji’s partulids was collated and two small, remote islands in the Fiji archipelago were surveyed to investigate whether tree snails persisted there. Living populations of Partula lanceolata and empty shells of Partula leefei were found on Cicia Island in Lau, and on Rotuma Island in the Rotuma Group, respectively. DNA analyses confirm a sister relationship between the two Partula species in north-eastern Lau, P. lirata and P. lanceolata, with both sharing a sister relationship with a member of the same genus in Vanuatu – P. auraniana Hartman, 1888. Prioritisation and further sampling of additional islands, and residual native habitat on less accessible islands and islets, is needed to fully assess the conservation status of all four Fijian species via the IUCN Red List process. Moreover, the basic descriptive information and associated studies reported here will serve to raise awareness of Fiji’s endemic tree snails particularly in communities that had no prior knowledge of their special conservation status; and also at a wider national, regional and global level. Community awareness is particularly vital as the willing support of land owners in the relevant small island communities is critical to implementing any future conservation action plans.

The two main biotic factors affecting grassland plant species are herbivory and competition. We investigated the significance of both these factors in establishing grassland communities through manipulation experiments conducted in both winter and summer in glasshouse conditions. Manipulations consisted of addition of above-ground and below-ground herbivores, reduction of herbivory through use of an insecticide, and varying of competitive pressure by removing either dicotyledonous or monocotyledonous seedlings as they appeared. In the winter experiment, the total biomass of dicotyledonous plants was reduced by both above-ground and below-ground herbivory, while monocotyledonous plant biomass was unaffected. However, in the summer experiment, only those treatments in which competition was manipulated had any detectable effect. This suggests that the relative importance of competition and herbivory can vary considerably. The above-ground herbivory treatment sometimes stimulated and sometimes reduced plant species diversity, and sometimes also induced significantly higher dicotyledonous species diversity than the root herbivory treatment. Those treatments in which competitive pressure was manipulated had no detectable effects on diversity. There were many strong effects detected when plant species were considered individually, with different species responding to different treatments, and with all treatments affecting at least some species. Different species therefore respond quite differently to competition and herbivory. None of the treatments affected soil microbial activity or active soil microbial biomass in the winter experiment, although active biomass was significantly positively correlated with dicotyledonous plant biomass. In the summer experiment, soil activity was positively affected by monocotyledonous plants, while active biomass was the same in all treatments except for the treatment with all plants removed, where biomass was less. None of the soil biotic variables showed any relationship with plant species richness. Our study shows that competition and herbivory can both exert important influences at both the community and ecosystem levels of resolution.

The flight behavior of Listronotus bonariensis (Kuschel) in Waikato (New Zealand) pastures was studied by recording catches on aerial sticky traps and sampling ground populations; this was supported by dissection studies and laboratory experimentation. All flight activity in the field occurred between 25 August and 5 May. Flight phenology and intensity varied greatly between localities and years. Intensity was generally low, although occasional mass flights were observed. Sampled ground populations indicated that only a proportion of the weevils exhibited development of flight musculature and hence flight capability: the proportion of these “flight capable” weevils varied greatly from generation to generation. Field sampling and laboratory experimentation indicated partitioning of reproductive and dispersive effort among individuals in the population. Following emergence, some weevils became reproductive and others developed flight musculature before ovarian development and dispersal. The proportion of weevils in the latter category was highest in the field following peak emergence and was shown by laboratory experimentation to be increased by weevil crowding, particularly on unfavorable host plants. The laboratory experiments also indicated that alleviation of crowding resulted in degeneration of flight musculature and resumption of reproductive activity. Field sampling indicated overwintering in the pastures (summer breeding grounds) and no migration to adjacent potential hibernation sites. These results are discussed in relation to the climate of the Waikato and the dynamics of the populations.

Argentine stem weevil (Listronotus bonariensis (Kuschel)) damage to pasture grasses was studied in relation to life cycle events in the Waikato. For a grazed pasture, numbers of adults at the onset of spring oviposition were low. First generation egg and larval numbers were correspondingly low, and the larvae caused negligible pasture damage. The second generation was considerably larger, and the larvae caused extensive damage. Applications of systemic insecticides increased the yield of grazed Nui ryegrass pasture by 17% over the spring-autumn period. Similarly in autumn-sown plots, Manawa, Ruanui, Nui, and Ellett ryegrasses, browntop, cocksfoot, paspalum, phalaris, and tall fescue, without clover, gave 15–91% higher yields under insecticide protection during the first spring-autumn period. Prairie grass production did not respond to insecticide application. Results show that responses in yield were caused by insecticidal control of stem weevil at a stage when the establishing grasses were parti...

Plots in a grazed trial in which 4 ryegrass cultivars received regular insecticidal applications of oxamyl were sampled at about 2-month intervals from September 1981 to May 1983 and generic nematode populations assessed. Significant treatment, sampling time, block, and cultivar effects were found. Oxamyl reduced most nematode populations, including Heterodera, Meloidogyne, Heticotylenchus, Paratylenchus, and Pungentus, possibly contributing to increased herbage production and influencing sward composition. The greatest cultivar effect was depression qf Paratylenchus beneath the low-yielding ‘Grasslands Manawa’. There was no evidence of nematodes developing tolerance to oxamyl, and population reduction occurred to 90 cm depth.

Survival of Argentine stem weevil from eggs to fourth instar larvae was compared in endophyte-infected and endophyte-free ryegrass in the screenhouse. Survival was significantly poorer in endophyte-infected plants. The results are discussed in relation to larval survival and damage in the field.