Untangling the Web: Spiders and the Challenges of Invertebrate Conservation

Selection of informative focal groups is an important avenue to increasing the applications of invertebrates in conservation assessments and inventory studies, and to help overcome the challenges of high diversity, all-taxon surveys and taxonomic inadequacy. Spiders are a possible focal group of wide relevance in terrestrial ecosystems, but considerable further work is needed to clarify their broad values as indicators, the relevance of higher taxon surrogacy, the taxonomy of selected families and genera, and to develop standard sampling protocols. The current values of spiders in conservation asessment, and the prospects of enhancing their value as a focal group are enumerated and discussed.     

Temporal and Taxonomic Variability in Response of Fauna to Riparian Restoration

Evaluating the success of restoration projects requires well‐designed studies. Among the decisions that need to be made are what taxonomic groups to study and when to conduct the monitoring. To explore how these decisions can influence assessments of restoration success, we examined species richness and composition data collected over several years on different terrestrial fauna (landbirds, rodents, bees, and beetles) at Sacramento River restoration and remnant riparian sites. Our selection of study organisms enabled us to ask whether variability in species richness among restoration sites is less for vagile taxa than for sedentary taxa, and if invertebrates display greater variability among sites than vertebrates. Our results demonstrate that responses to restoration can vary depending upon the season when it is assessed, and the taxa that are studied. For all taxa except bees, there was considerable variability in the relative performance of taxa at restoration sites from one sampling date to the next, such that the relative ranking of the sites often changed dramatically. Comparisons of β ‐diversity (variability in species richness across sites) revealed that certain taxonomic groups were more spatially variable in their response to restoration than others. Among vertebrates, sedentary taxa (rodents) had significantly higher variability in species richness across sites than highly vagile taxa (birds); however, no such pattern was observed for invertebrates. Overall, vertebrates had lower variability than invertebrates, suggesting that evaluations of restoration success based on a few better‐known taxonomic groups (e.g., birds, rodents) may be inadequate to represent the biodiversity response of other groups (e.g., insects).     

Beyond Surrogacy—A Multi‐taxon Approach to Conservation Biogeography

Biogeographic analysis of species turnover (β diversity) of plants and animals among regions often yields conflicting results, with regions of high β diversity identified for some taxonomic groups but not others. Such discordance calls into question the use of surrogate taxa to forge conservation plans. This discordance begs for a means of comparing multiple taxa across phyla in a manner that is cost‐effective, considers limitations in computer resources in certain global regions, and is understood by policy makers and land managers. As a test case for a method taking into account these considerations, we used species lists for ten organismal groups (spanning plants, vertebrates, and invertebrates) to identify regions of high β diversity using Monmonier's algorithm, a spatially explicit technique that is readily implemented and interpreted. Data were for montane (>1000 m elevation) species across the Eastern Arc of Tanzania and Kenya and surrounding areas. Our results indicate that surrogacy does not make for the most effective expenditure of conservation efforts. We also show that to use a multi‐taxon approach one need not rely on intensive surveys of areas in order to make conservation decisions, including reserve selection. Our approach also eliminates the need for complex modeling and comparisons common to many GIS‐based complementarity techniques. Additionally, a wide variety of socioeconomic, political, demographic, geological, climatological, and evolutionary factors can be incorporated into the technique to help shape conservation biogeography from a local and regional perspective. This technique can bridge the gap between conservation biogeography theory and application in tropical regions and beyond.     

Flora and fauna assemblages vary with local topography in a subtropical eucalypt forest

Local‐scale spatial patterning (over hundreds of metres) in terrestrial assemblages was investigated by sampling a variety of organisms within a 400 ha eucalypt forest area in the lowlands of south‐east Queensland. Organisms were trees, shrubs, birds, insects extracted from the litter layer, and insects caught in pitfall traps. Each group was sampled using a standardized methodology, and the component taxa were counted and identified to a level commonly used in ecological studies of that organism – varying from species to order levels. Sites adjacent to drainage lines or ephemeral streams were biotically more similar to one another than they were to paired upslope sites 100–200 m distant and 15–35 m higher in altitude. This phenomenon occurred irrespective of the level of taxonomic resolution or type of organism. Within each taxonomic group, some components were mainly riparian, while others were more characteristic of upslope sites. Characteristically riparian taxa included trees in the genus Glochidion, the shrub genus Leptospermum, birds in the Pachycephalidae and Meliphagidae families, and litter invertebrates in the order Acarina. Upslope taxa included shrubs in the Rutaceae, birds in the Artamidae, and ants. Within the groups of trees, birds and litter invertebrates, more taxa were characteristic of riparian than upslope sites. Local scale biotic patterns were more strongly correlated with altitude than with measured soil characteristics; however, microtopographical differences would also be highly correlated with a large suite of covarying environmental features. The patterns of diversity and the implications for survey design and conservation are discussed.     

Three reasons to re-evaluate fungal diversity ‘on Earth and in the ocean’

Attempts to assess fungal global species richness are confounded by several problems: uncertainty about the number of described species, incomplete fungal inventories even at a high taxonomic level, high diversity of unknown, often small and elusive taxa, high levels of morphological conservation, and incomplete knowledge of their ecological and biogeographical distributions. The two main bases for estimating total fungal diversity are (1) the number of described species and their taxonomic structure, and (2) extrapolating species-area relationships. We argue that knowledge of fungal taxonomy and environmental sampling of fungi are both too incomplete for either approach to be reliable. However, it is likely that the true number of fungal species on the planet is a seven-digit number, and may even be an order of magnitude higher.     

Three reasons to re-evaluate fungal diversity ‘on Earth and in the ocean’

Attempts to assess fungal global species richness are confounded by several problems: uncertainty about the number of described species, incomplete fungal inventories even at a high taxonomic level, high diversity of unknown, often small and elusive taxa, high levels of morphological conservation, and incomplete knowledge of their ecological and biogeographical distributions. The two main bases for estimating total fungal diversity are (1) the number of described species and their taxonomic structure, and (2) extrapolating species-area relationships. We argue that knowledge of fungal taxonomy and environmental sampling of fungi are both too incomplete for either approach to be reliable. However, it is likely that the true number of fungal species on the planet is a seven-digit number, and may even be an order of magnitude higher.     

Indicators of species richness at the local scale in an alpine region: a comparative approach between plant and invertebrate taxa

Studies investigating congruent variations in species richness patterns in alpine habitats are scarce. We investigated the potential of using the indicator taxa approach for species richness in alpine habitats of the Scandes (Norway). In four areas, we investigated seven functional and taxonomic terrestrial groups of organisms and evaluated their contribution to the species diversity. The function of each group as a surrogate for the overall species diversity or for the diversity of another taxon was analysed. Three groups of invertebrates (spiders without Lycosids, Lycosids only, and ground beetles), three groups of plants (shrubs, graminoids, and herbs), and lichens were used for a cross-taxon analysis of species diversity. Congruence in species richness was restricted to several significant results, with vascular plants and spiders (Araneae) being best suited as surrogate taxa in alpine habitats of the Scandes. In the cross-taxon analyses they showed strongest significant positive correlations, covering the total species richness of the alpine habitats best. Species counts in one group account for up to 70% of the variation in total species richness. We found only limited evidence for an ideal, efficient biodiversity indicator taxon that could be applied without restrictions at different alpine habitats in low and middle alpine areas. Thus, our results suggest that it is very important to use more than one taxon as indicator for species richness in terrestrial alpine habitats. This should facilitate future conservation planning in alpine areas.     

Variation in invertebrate–bryophyte community structure at different spatial scales along altitudinal gradients

Aim This study assessed changes in diversity and assemblage composition in bryophytes and their associated invertebrates along altitudinal gradients in Australia and New Zealand. The importance of altitude in shaping these communities and for the diversity of both invertebrates and bryophytes was examined at different spatial scales, including local, altitudinal, regional and biogeographical. Location Samples were taken from four Australasian mountain ranges between 42° and 43°S: Mt Field and Mt Rufus, Tasmania, Australia, and Otira Valley and Seaward Kaikoura Mountains, South Island, New Zealand. Methods On both Tasmanian mountains, five altitudes were assessed (250, 500, 750, 1000 and 1250 m). At each location (mountain/altitude combination) two sites were chosen and six samples were taken. Six altitudes were assessed on New Zealand mountains (Otira: 250, 500, 750, 1000, 1250 and 1500 m; Kaikoura: 1130, 1225, 1325, 1425, 1525 and 2000 m). Bryophyte substrate was collected, and all samples were stored in 70% ethanol. Invertebrates were extracted from bryophytes using kerosene‐phase separation and all invertebrates were identified to family. At each location in Tasmania, all bryophyte species within six 25‐cm² grids per site were collected and identified to species. Bryophytes from New Zealand were identified to species from the invertebrate sample substrate because of sampling constraints. Results Altitude did have a significant effect on diversity, however, no general trend was found along the altitudinal gradient on the four mountains. There were distinct differences in diversity between biogeographical regions, mountains, altitudes and sites. In Tasmania, Mt Field had the highest diversity in invertebrates and bryophytes at 750 m. In contrast, Mt Rufus had consistent low invertebrate and bryophyte diversity along the entire altitudinal gradient. There were also distinctive differences between locations in the composition of invertebrate and bryophyte communities in Tasmania. Along the two altitudinal gradients in New Zealand, Otira had highest diversity for both invertebrates and bryophytes at low altitudes, whereas Kaikoura had highest invertebrate and lowest bryophyte diversity at the highest altitude. Main conclusions There was an effect of altitude, however, there were no consistent changes in diversity or composition on the four different mountains. There was considerable local and regional variation, and, despite a strong sampling design, no underlying altitudinal trends were detectable. This study demonstrates the importance of examining a range of spatial scales if patterns in community structure along altitudinal gradients are to be studied. The implications of this study are discussed with reference to survey design, taxonomic resolution, climate change and conservation of habitat.     

Cryptic diversity in vertebrates: molecular data double estimates of species diversity in a radiation of Australian lizards (Diplodactylus,Gekkota)

A major problem for biodiversity conservation and management is that a significant portion of species diversity remains undocumented (the ‘taxonomic impediment’). This problem is widely acknowledged to be dire among invertebrates and in developing countries; here, we demonstrate that it can be acute even in conspicuous animals (reptiles) and in developed nations (Australia). A survey of mtDNA, allozyme and chromosomal variation in the Australian gecko, genus Diplodactylus, increases overall species diversity estimates from 13 to 29. Four nominal species each actually represent multi-species complexes; three of these species complexes are not even monophyletic. The high proportion of cryptic species discovered emphasizes the importance of continuing detailed assessments of species diversity, even in apparently well-known taxa from industrialized countries.     

Taxonomic surrogacy in biodiversity assessments,and the meaning of Linnaean ranks

The majority of biodiversity assessments use species as the base unit. Recently, a series of studies have suggested replacing numbers of species with higher ranked taxa (genera, families, etc.); a method known as taxonomic surrogacy that has an important potential to save time and resources in assesments of biological diversity. We examine the relationships between taxa and ranks, and suggest that species/higher taxon exchanges are founded on misconceptions about the properties of Linnaean classification. Rank allocations in current classifications constitute a heterogeneous mixture of various historical and contemporary views. Even if all taxa were monophyletic, those referred to the same rank would simply denote separate clades without further equivalence. We conclude that they are no more comparable than any other, non‐nested taxa, such as, for example, the genus Rattus and the phylum Arthropoda, and that taxonomic surrogacy lacks justification. These problems are also illustrated with data of polychaetous annelid worms from a broad‐scale study of benthic biodiversity and species distributions in the Irish Sea. A recent consensus phylogeny for polychaetes is used to provide three different family‐level classifications of polychaetes. We use families as a surrogate for species, and present Shannon‐Wiener diversity indices for the different sites and the three different classifications, showing how the diversity measures rely on subjective rank allocations.     

Invertebrate survey of coastal habitats and podocarp forest on Ulva Island,Rakiura National Park,New Zealand

Abstract

Inventory of the invertebrate fauna is important to establish taxonomic diversity, abundance and distribution, and hence the conservation of indigenous biodiversity. Invertebrate assemblages have been documented in some broadleaf‐podocarp forests and grassland habitats in New Zealand, but not in dense stands of coastal forest or in mature podocarp forest. This survey aimed to provide a taxonomic inventory of terrestrial invertebrates and their habitat associations on Ulva Island (Rakiura National Park, Stewart Island), an off‐shore sanctuary of significant conservation value in New Zealand. We systematically documented the invertebrate assemblages collected in ground litter and on tree trunks on the island. The invertebrate specimens identified represented 4 phyla, 6 classes, 25 orders and 62 species. The invertebrate fauna reported in this survey was distinct from those of lowland shrubland and broadleaved‐Nothofagus forests on the mainland, but shared species with that reported from another similar off‐shore island, Codfish Island (Whenua Hou).     

Climatic vulnerabilities and ecological preferences of soil invertebrates across biomes

Unlike plants and vertebrates, the ecological preferences, and potential vulnerabilities of soil invertebrates to environmental change, remain poorly understood in terrestrial ecosystems globally. We conducted a cross‐biome survey including 83 locations across six continents to advance our understanding of the ecological preferences and vulnerabilities of the diversity of dominant and functionally important soil invertebrate taxa, including nematodes, arachnids and rotifers. The diversity of invertebrates was analyzed through amplicon sequencing. Vegetation and climate drove the diversity and dominant taxa of soil invertebrates. Our results suggest that declines in forest cover and plant diversity, and reductions in plant production associated with increases in aridity, can result in reductions of the diversity of soil invertebrates in a drier and more managed world. We further developed global atlases of the diversity of these important soil invertebrates, which were cross‐validated using an independent database. Our study advances the current knowledge of the ecological preferences and vulnerabilities of the diversity and presence of functionally important soil invertebrates in soils from across the globe. This information is fundamental for improving and prioritizing conservation efforts of soil genetic resources and management policies.     

Rapid biodiversity assessment of arthropods for monitoring average local species richness and related ecosystem services

Rapid biodiversity assessment (RBA) is proposed as an affordable indicator for monitoring local species richness of arthropods and sustainability of related ecosystem services. The indicator is based on strictly standardised sampling procedures and the identification of parataxonomic units (morphospecies) instead of species identification. The collection of arthropods was optimized with regard to trap types, time and length of collecting period, selection of four out of seven weekly samples, and choice of counted taxa and trophic guilds. By measuring arthropod activity, RBA is an indicator for functional diversity. Over a period of 8 years, average yearly numbers of morphospecies were assessed in Switzerland in 15 agricultural habitats, 15 managed forests, and in 12 unmanaged habitats ranging from protected lowland wetlands to Alpine meadows. The yearly RBA-trend in unmanaged habitats is used for assessing the influence of climate and weather on biodiversity, and as a reference for measuring the relative influences of recent management changes in agriculture and forestry. The average number of morphospecies per sampling station per year depends on temperature, and was only marginally significantly increasing over time in agriculture, but not in forestry or unmanaged areas. Three RBA indices considered to be relevant for maintaining ecosystem services were calculated from the average number of morphospecies per location per year: (1) indicator for ecological resilience and sustainability (all morphospecies); (2) indicator for pollinator diversity (taxa with a majority of pollinators) and (3) indicator for biocontrol diversity (ratio between carnivore and herbivore guilds).     

New frameworks for species surrogacy in monitoring highly variable coastal ecosystems: Applying the BestAgg approach to Mediterranean coastal lagoons

Benthic invertebrates are good indicators of aquatic ecosystem health. Yet, environmental monitoring and assessment of community changes in relation to both natural and human sources of disturbance involves considerable efforts for sample processing and time-consuming identifications of organisms, which make challenging large-scale and continuous monitoring programs required under the current regulatory frameworks at European scale. The use of higher taxa (e.g. families) as surrogates for species is a mainstream approach to reduce cost and time associated to fine taxonomic resolution in environmental studies, especially concerning macro-invertebrate communities. However, this approach of ‘taxonomic sufficiency’ simply relies on the static grouping of organisms in taxa belonging to a single higher taxonomic level irrespective of their ecological relevance or difficulties in their taxonomic identification, leading to unnecessary losses of taxonomic detail or ecological information. A new approach, namely the Best Practicable Aggregation of Species (BestAgg), has been recently developed as an alternative procedure for species surrogacy. BestAgg is based on aggregating species in the minimum number of surrogates sufficient to depict species-level community patterns consistently, while capitalizing on ecological information. Although the approach has been successfully applied to marine and freshwater invertebrate assemblages, its effectiveness in transitional water systems, where the complex and highly variable environmental conditions may affect the performance of surrogacy methods, still remain untested. Here, we applied the BestAgg approach to quantifying spatio-temporal patterns of variability of macro-invertebrate assemblages from Mediterranean coastal lagoons (Northern Adriatic Sea). Surrogates were defined using species-level data from a representative lagoon system, which served as pilot study. Then, they were used to analyze macro-invertebrate assemblages in two independent lagoons in the same geographic area. Results showed that BestAgg surrogates were effective in depicting multivariate patterns of macro-invertebrate assemblages as at species level over and beyond potential variations among the investigated lagoons. The use of families, following a more classic approach based on taxonomic sufficiency, also provided results comparable to those obtained using species. However, with respect to families, BestAgg surrogates allowed an estimated timesaving of 40% higher while still retaining an equivalent amount of information on species-level patterns. More importantly, BestAgg allowed exploiting different criteria of species aggregation, leading to a set of surrogates more aligned with ecological/functional characteristics of organisms, suggesting that BestAgg approach may provide a fresh perspective for optimizing trade-offs between pragmatism and the need for relevant ecological information in environmental assessment and monitoring.     

Resource seasonality and fish diets in an Illinois stream

Synopsis The purpose of this study was to evaluate the intensity of competition for food among 9 species of stream fishes that primarily eat aquatic invertebrates. The taxonomic and size composition, and numerical abundance of aquatic invertebrates were monitored for one year using drift and benthic samples. Diet data were obtained from stomachs of fishes captured at the same time and place that invertebrate sampling was done. Diet characteristics examined included taxonomic and size composition, number of prey per fish, and diet breadth. Drifting invertebrates were more abundant early in the year (March–June) than later (July–January). The summer-early fall scarcity of invertebrates was especially notable among those>3.6 mm long, which comprised the bulk of prey found in fish stomachs. Average prey size eaten by a fish species was positively correlated with fish mouth size, but interspecific overlap in prey size was extensive. Cyprinids as a group (5 species) ate proportionally fewer small (< 3.6 mm long) prey from July to January than did the centrarchids and stonecat. Taxonomic compositions of available invertebrates and fish diets varied markedly among sampling dates, but the use of prey taxa by fishes was not correlated with the availability of those taxa. Use of aquatic prey taxa was generally similar among fish species, but cyprinids as a group ate proportionally more terrestrial prey from July to January than did the centrarchids and stonecat. Diet breadths for all species increased as food levels declined, indicating that these fishes experienced resource depression. Food scarcity was evidently more severe for cyprinids since their stomachs contained few prey through the summer and fall relative to the centrarchids and stonecat. Though the fish species studied probably compete for food in the summer and fall, this competition did not account for the community structure observed.     

Different surrogacy approaches for stream macroinvertebrates in discriminating human disturbances in Central China

Due to the difficulty of identifying many taxa of freshwater invertebrates to species, many researchers have assessed the utility of surrogates for species-level identifications (e.g. higher taxa) in bioassessment programs. Here, we examined the efficiency of two different approaches to species surrogacy, one using coarser taxonomic resolution and a second approach based on random aggregation (“Best practicable aggregation of species”, BestAgg), in portraying patterns of stream macroinvertebrates in Central China. The main objectives were: (1) to compare the discriminatory power of biodiversity indices and assemblage structure for different levels of human disturbances based on different taxonomic resolution and on BestAgg; (2) to identify the congruence of assemblage-environment and biodiversity-indices-environment relationships for datasets at the species level versus those at surrogate levels. We found that genus-level and BestAgg datasets accurately reproduced the pattern of species-level communities, whereas family- and order-level datasets did not. Specifically, both genus-level and BestAgg approaches performed almost as well as species-level data in distinguishing sites subjected to different disturbance levels. Most of the environmental variables that were important for species-level assemblages, also emerged as significant when analyzing genera and BestAgg surrogates, as shown by both analyses of indices and assemblage composition according to distance-based ordination models. Our results suggest that genus-level taxonomy, which resulted in the least loss of ecological information relative to species-level identification, is sufficient in studies of community ecology and bioassessment of stream macroinvertebrates in Central China. In addition, the BestAgg approach, which required identification of fewer taxa than genus-level analysis, has a similar ability to depict multivariate patterns of macroinvertebrate assemblages and differentiate different disturbance levels. Applying our results could enhance speed and cost-effectiveness of freshwater biomonitoring and bioassessment programs; however, independent determination of best taxonomic level and BestAgg will be required whenever a new geographic area or habitat type is assessed.     

Land‐use effects on the functional distinctness of arthropod communities

Land‐use change is a major driver of the global loss of biodiversity, but it is unclear to what extent this also results in a loss of ecological traits. Therefore, a better understanding of how land‐use change affects ecological traits is crucial for efforts to sustain functional diversity. To this end we tested whether higher species richness or taxonomic distinctness generally leads to increased functional distinctness and whether intensive land use leads to functionally more narrow arthropod communities. We compiled species composition and trait data for 350 species of terrestrial arthropods (Araneae, Carabidae and Heteroptera) in different land‐use types (forests, grasslands and arable fields) of low and high land‐use intensity. We calculated the average functional and taxonomic distinctness and the rarified trait richness for each community. These measures reflect the range of traits, taxonomic relatedness and number of traits that are observed in local communities. Average functional distinctness only increased significantly with species richness in Carabidae communities. Functional distinctness increased significantly with taxonomic distinctness in communities of all analyzed taxa suggesting a high functional redundancy of taxonomically closely related species. Araneae and Heteroptera communities had the expected lower functional distinctness at sites with higher land‐use intensity. More frequently disturbed land‐use types such as managed grasslands or arable fields were characterized by species with smaller body sizes and higher dispersal abilities and communities with lower functional distinctness or trait richness. Simple recommendations about the conservation of functional distinctness of arthropod communities in the face of future land‐use intensification and species loss are not possible. Our study shows that these relationships depend on the studied taxa and land‐use type. However, for some arthropod groups functional distinctness is threatened by intensification and conversion from less to more frequently disturbed land‐uses.     

Bivalves in a bottleneck: taxonomy,phylogeography and conservation of freshwater mussels (Bivalvia: Unionoida) in Australasia

The conservation biology of Australasian freshwater mussels is hindered by lack of a taxonomic framework that employs molecular data as a complement to shell characters, larval forms and internal anatomy. The fauna includes more than 32 known species (30+ Hyriidae, 2 Unionidae), but has not been revised for 55 years, despite minor amendments. The hyriids are relics of Gondwana, represented in Australia and New Guinea by the ancestral Velesunioninae and in Australia and New Zealand by the Hyriinae (Tribe Hyridellini). Many taxonomic and phylogeographic issues await resolution, including the relationships between Australasian and South American species, and between Australian and New Zealand species, and the status of species in New Guinea (including uncertain reports of Unionidae) and the Solomon Islands. Once these are clarified, it will be easier to identify threatened species and evaluate the conservation status of the fauna. At present, only seven taxa are named in the IUCN Red List or under national/state legislation, and these are not representative. Threatening processes include altered flow regimes, catchment disturbances, salinisation, pollution and invasive species. While the need for a taxonomic revision is paramount, progress in conservation may depend also upon involving the wider community.     

Direct Measurement Versus Surrogate Indicator Species for Evaluating Environmental Change and Biodiversity Loss

The enormity and complexity of problems like environmental degradation and biodiversity loss have led to the development of indicator species and other surrogate approaches to track changes in environments and/or in biodiversity. Under these approaches particular species or groups of species are used as proxies for other biota, particular environmental conditions, or for environmental change. The indicator species approach contrasts with a direct measurement approach in which the focus is on a single entity or a highly targeted subset of entities in a given ecosystem but no surrogacy relationships with unmeasured entities are assumed. Here, we present a broad philosophical discussion of the indicator species and direct measurement approaches because their relative advantages and disadvantages are not well understood by many researchers, resource managers and policy makers. A goal of the direct measurement approach is to demonstrate a causal relationship between key attributes of the target ecosystem system (for example, particular environmental conditions) and the entities selected for measurement. The key steps in the approach are based on the fundamental scientific principles of hypothesis testing and associated direct measurement that drive research activities, management activities and monitoring programs. The direct measurement approach is based on four critical assumptions:(1) the ‘right’ entities to measure have been selected, (2) these entities are well known, (3) there is sufficient understanding about key ecological processes and (4) the entities selected can be accurately measured. The direct measurement approach is reductionist and many elements of the biota, many biotic processes and environmental factors must be ignored because of practical considerations. The steps in applying the indicator species approach are broadly similar to the direct measurement approach, except surrogacy relationships also must be quantified between a supposed indicator species or indicator group and the factors for which it is purported to be a proxy. Such quantification needs to occur via: (1) determining the taxonomic, spatial and temporal bounds for which a surrogacy relationship does and does not hold. That is, the extent of transferability of a given surrogate such as an indicator species to other biotic groups, to landscapes, ecosystems, environmental circumstances or over time in the same location can be determined; and (2) determining the ecological mechanisms underpinning a surrogacy relationship (for example, through fundamental studies of community structure). Very few studies have rigorously addressed these two tasks, despite the extremely widespread use of the indicator species approach and similar kinds of surrogate schemes in virtually all fields of environmental, resource and conservation management. We argue that this has the potential to create significant problems; thus, the use of an indicator species approach needs to be better justified. Attempts to quantify surrogacy relationships may reveal that, in some circumstances, the alternative of direct measurement of particular entities of environmental or conservation interest will be the best option.     

Diversification of New Zealand weta (Orthoptera: Ensifera: Anostostomatidae) and their relationships in Australasia

New Zealand taxa from the Orthopteran family Anostostomatidae have been shown to consist of three broad groups, Hemiandrus (ground weta), Anisoura/Motuweta (tusked weta) and Hemideina-Deinacrida (tree-giant weta). The family is also present in Australia and New Caledonia, the nearest large land masses to New Zealand. All genera are endemic to their respective countries except Hemiandrus that occurs in New Zealand and Australia. We used nuclear and mitochondrial DNA sequence data to study within genera and among species-level genetic diversity within New Zealand and to examine phylogenetic relationships of taxa in Australasia. We found the Anostostomatidae to be monophyletic within Ensifera, and justifiably distinguished from the Stenopelmatidae among which they were formerly placed. However, the New Zealand Anostostomatidae are not monophyletic with respect to Australian and New Caledonian species in our analyses. Two of the New Zealand groups have closer allies in Australia and one in New Caledonia. We carried out maximum-likelihood and Bayesian analyses to reveal several well supported subgroupings. Our analysis included the most extensive sampling to date of Hemiandrus species and indicate that Australian and New Zealand Hemiandrus are not monophyletic. We used molecular dating approaches to test the plausibility of alternative biogeographic hypotheses for the origin of the New Zealand anostostomatid fauna and found support for divergence of the main clades at, or shortly after, Gondwanan break-up, and dispersal across the Tasman much more recently.