► We identify seven impediments to invertebrate conservation. ► Three dilemmas: public, political and scientific. ► Four shortfalls: Linnean, Wallacean, Prestonian and Hutchinsonian. ► We present ...possible solutions for each impediment.
Despite their high diversity and importance for humankind, invertebrates are often neglected in biodiversity conservation policies. We identify seven impediments to their effective protection: (1) invertebrates and their ecological services are mostly unknown to the general public (the public dilemma); (2) policymakers and stakeholders are mostly unaware of invertebrate conservation problems (the political dilemma); (3) basic science on invertebrates is scarce and underfunded (the scientific dilemma); (4) most species are undescribed (the Linnean shortfall); (5) the distribution of described species is mostly unknown (the Wallacean shortfall); (6) the abundance of species and their changes in space and time are unknown (the Prestonian shortfall); (7) species ways of life and sensitivities to habitat change are largely unknown (the Hutchinsonian shortfall).
Numerous recent developments in taxonomy, inventorying, monitoring, data compilation, statistical analysis and science communication facilitate overcoming these impediments in both policy and practice. We suggest as possible solutions for the public dilemma: better public information and marketing. For the political dilemma: red-listing, legal priority listing and inclusion in environmental impact assessment studies. For the scientific dilemma: parataxonomy, citizen science programs and biodiversity informatics. For the Linnean shortfall: biodiversity surrogacy, increased support for taxonomy and advances in taxonomic publications. For the Wallacean shortfall: funding of inventories, compilation of data in public repositories and species distribution modeling. For the Prestonian shortfall: standardized protocols for inventorying and monitoring, widespread use of analogous protocols and increased support for natural history collections. For the Hutchinsonian shortfall: identifying good indicator taxa and studying extinction rates by indirect evidence.
1. Fifteen species richness estimators (three asymptotic based on species accumulation curves, 11 nonparametric, and one based in the species-area relationship) were compared by examining their ...performance in estimating the total species richness of epigean arthropods in the Azorean Laurisilva forests. Data obtained with standardized sampling of 78 transects in natural forest remnants of five islands were aggregated in seven different grains (i.e. ways of defining a single sample): islands, natural areas, transects, pairs of traps, traps, database records and individuals to assess the effect of using different sampling units on species richness estimations. 2. Estimated species richness scores depended both on the estimator considered and on the grain size used to aggregate data. However, several estimators (ACE, Chao1, Jackknife1 and 2 and Bootstrap) were precise in spite of grain variations. Weibull and several recent estimators proposed by Rosenzweig et al., and Ugland et al. performed poorly. 3. Estimations developed using the smaller grain sizes (pair of traps, traps, records and individuals) presented similar scores in a number of estimators (the above-mentioned plus ICE, Chao2, Michaelis-Menten, Negative Exponential and Clench). The estimations from those four sample sizes were also highly correlated. 4. Contrary to other studies, we conclude that most species richness estimators may be useful in biodiversity studies. Owing to their inherent formulas, several nonparametric and asymptotic estimators present insensitivity to differences in the way the samples are aggregated. Thus, they could be used to compare species richness scores obtained from different sampling strategies. Our results also point out that species richness estimations coming from small grain sizes can be directly compared and other estimators could give more precse results in those cases. We propose a decision framework based on our results and on the literature to assess which estimator should be used to compare species richness scores of different sites, depending on the grain size of the original data, and of the kind of data available (species occurrence or abundance data).
Aphids are a serious threat to agriculture, despite being a rather small group of insects. The about 4,000 species worldwide engage in highly interesting and complex relationships with their ...microbial fauna. One of the key symbionts in arthropods is Wolbachia, an α-Proteobacterium implicated in many important biological processes and believed to be a potential tool for biological control. Aphids were thought not to harbour Wolbachia; however, current data suggest that its presence in aphids has been missed, probably due to the low titre of the infection and/or to the high divergence of the Wolbachia strains of aphids. The goal of the present study is to map the Wolbachia infection status of natural aphids populations, along with the characterization of the detected Wolbachia strains. Out of 425 samples from Spain, Portugal, Greece, Israel and Iran, 37 were found to be infected. Our results, based mainly on 16S rRNA gene sequencing, indicate the presence of two new Wolbachia supergroups prevailing in aphids, along with some strains belonging either to supergroup B or to supergroup A.
Climate change is causing shifts in species distributions worldwide. Understanding how species distributions will change with future climate change is thus critical for conservation planning. Impacts ...on oceanic islands are potentially major given the disproportionate number of endemic species and the consequent risk that local extinctions might become global ones. In this study, we use species climate envelope models to evaluate the current and future potential distributions of Azorean endemic species of bryophytes, vascular plants, and arthropods on the Islands of Terceira and São Miguel in the Azores archipelago (Macaronesia). We examined projections of climate change effects on the future distributions of species with particular focus on the current protected areas. We then used spatial planning optimization software (PRION) to evaluate the effectiveness of protected areas at preserving species both in the present and future. We found that contractions of species distributions in protected areas are more likely in the largest and most populated island of São Miguel, moving from the coastal areas towards inland where the current protected areas are insufficient and inadequate to tackle species distribution shifts. There will be the need for a revision of the current protected areas in São Miguel to allow the sustainable conservation of most species, while in Terceira Island the current protected areas appear to be sufficient. Our study demonstrates the importance of these tools for informing long-term climate change adaptation planning for small islands.
► IUCN criteria have shortcomings for invertebrates. ► Risk categories may be wrongly applied. ► Assessments should rely on Area of Occupancy and Extent of Occurrence and use ecological modelling. ► ...Co-extinction should be considered.
The IUCN Red List is the most useful list of species that are at risk for extinction worldwide, as it relies on a number of objective criteria. Nevertheless, there is a taxonomic bias that excludes species with small body sizes, narrow distribution ranges and low dispersal abilities, which constitute the vast majority of the planet’s biota, particularly local endemics.
By evaluating each IUCN criterion separately, we (i) identify the shortcomings for invertebrate applications, (ii) explain how risk categories may be wrongly applied due to inapplicable and/or misleading thresholds, (iii) suggest alternative ways of applying the existing criteria in a more realistic way and (iv) suggest possible new criteria that were not considered in the current evaluation framework but that could allow a more comprehensive and effective assessment of invertebrates.
By adapting the criteria to rely more explicitly on the Area of Occupancy and the Extent of Occurrence, their respective trends and by using ecological modelling methods, the criteria’s applicability would be increased. The change in some thresholds or, eventually, the creation of sub-categories would further increase their adequacy. Additionally, co-extinction could be introduced as an explicit part of the classification process.
As a case study, we evaluated 48 species of Azorean arthropods and Iberian spiders according to the current criteria. More than one-quarter (27%) of all evaluated species were classified as Critically Endangered, 19% as Endangered, 6% as Vulnerable and 8% as Least Concern. The remaining 40% did not have enough data to reach a classification.
Oceanic islands have been providing important insights on the structuring of ecological communities and, under the context of the present biodiversity crisis, they are paramount to assess the effects ...of biological invasions on community assembly. In this study we compare the taxonomic and functional diversity of insect herbivore assemblages associated with the dominant tree species of Azorean native forests and investigate the ecological processes that may have originated current patterns of plant-herbivore associations. Five dominant trees-Erica azorica, Ilex perado subsp. azorica, Juniperus brevifolia, Laurus azorica and Vaccinium cylindraceum-were sampled in the remnants of the native forest of Terceira Island (Azores) using a standardised methodology. The taxonomic and functional diversity of insect herbivore assemblages was assessed using complementary metrics and beta diversity partitioning analysis (species replacement and richness differences) aiming to evaluate the variation in insect herbivore assemblages within and between the study plant species. Sixty two insect species, mostly bugs (Hemiptera) and caterpillars (Lepidoptera), were found in the five study plants with indigenous (endemic and native non-endemic) insects occurring with higher species richness and abundance than introduced ones. Species replacement was the most important component of insect herbivore taxonomic beta diversity while differences in trait richness played a major role on functional beta diversity. The endemic E. azorica stands out from the other study plants by having associated a very distinct insect herbivore assemblage with a particular set of functional attributes, mainly composed by large bodied and long shaped species that feed by chewing. Despite the progressive biotic homogenization witnessed in the Azores during the last few decades, several strong associations between the endemic trees and their indigenous insect herbivores remain.
Beta diversity and nestedness are central concepts of ecology and biogeography and evaluation of their relationships is in the focus of contemporary ecological and conservation research. Beta ...diversity patterns are originated from two distinct processes: the replacement (or turnover) of species and the loss (or gain) of species leading to richness differences. Nested distributional patterns are generally thought to have a component deriving from beta diversity which is independent of replacement processes. Quantification of these phenomena is often made by calculating a measure of beta diversity, and the resulting value being subsequently partitioned into a contribution by species replacement plus a fraction shared by beta diversity and nestedness. Three methods have been recently proposed for such partitioning, all of them based on pairwise comparisons of sites. In this paper, the performance of these methods was evaluated on theoretical grounds and tested by a simulation study in which different gradients of dissimilarity, with known degrees of species replacement and species loss, were created. Performance was also tested using empirical data addressing land-use induced changes in endemic arthropod communities of the Terceira Island in the Azores. We found that the partitioning of β cc (dissimilarity in terms of the Jaccard index) into two additive fractions, β₋₃ (dissimilarity due to species replacement) plus β rich (dissimilarity due to richness differences) reflects the species replacement and species loss processes across the simulated gradients in an ecologically and mathematically meaningful way, whilst the other two methods lack mathematical consistency and prove conceptually self-contradictory. Moreover, the first method identified a selective local extinction process for endemic arthropods, triggered by land-use changes, while the latter two methods overweighted the replacement component and led to false conclusions. Their basic flaw derives from the fact that the proposed replacement and nestedness components (deemed to account for species loss) are not scaled in the same way as the measure that accounts for the total dissimilarity (Sørensen and Jaccard indices). We therefore recommend the use of β cc = β₋₃ + β rich , since its components are scaled in the same units and their responses are proportional to the replacement and the gain/loss of species.
Researchers measuring beta diversity have rarely concerned themselves with the problems of how complete the species lists of studied communities are, and of how the varying degrees of completeness ...can actually change estimates of beta diversity. No comprehensive assessment has been made regarding the behaviour of most beta diversity indices when applied to incomplete samples, a situation which is more common than usually recognized. Our objective was to assess the behaviour and robustness of a number of beta diversity measures for incidence data from undersampled communities. Mainland Portugal and the Azorean archipelago (North Atlantic). Data from intensive sampling of spiders in mainland Portugal and arthropods in Azores were collected. We examined the properties of 15 beta diversity measures developed for incidence data. We simulated varying degrees of completeness, whereas computing beta diversity for selected pairs of samples. The robustness of these beta diversity accumulation curves was assessed for the purpose of finding the best measures for undersampled communities. The Harrison et al.β₋₂ and the Williams β₋₃ are particularly robust to undersampling. These measures are also insensitive to differences of alpha diversity (species richness) between communities, and therefore to nestedness. Colwell & Coddington βcc and the related Jaccard βj and Gaston et al.βg performed best of the measures sensitive to alpha diversity. They performed poorly, however, when compared communities exhibited very low values of beta diversity. In such cases, the Routledge βr performed the best. No index was found to perform without bias in all circumstances. Overall, β₋₂, β₋₃ and βcc (or related measures βj and βg) are recommended as they seem to be the most robust to undersampling.
Long-term monitoring of invertebrate communities is needed to understand the impact of key biodiversity erosion drivers (e.g. habitat fragmentation and degradation, invasive species, pollution, ...climatic changes) on the biodiversity of these high diverse organisms.The data we present are part of the long-term project SLAM (Long Term Ecological Study of the Impacts of Climate Change in the natural forest of Azores) that started in 2012, aiming to understand the impact of biodiversity erosion drivers on Azorean native forests (Azores, Macaronesia, Portugal). In this contribution, the design of the project, its objectives and the first available data for the spider fauna of two Islands (Pico and Terceira) are described.Passive flight interception SLAM traps (Sea, Land and Air Malaise traps) were used to sample native forest plots in several Azorean islands, with one trap being set up at each plot and samples taken every three months following the seasons.The key objectives of the SLAM project are: 1) collect long-term ecological data to evaluate species distributions and abundance at multiple spatial and temporal scales, responding to the Wallacean and Prestonian shortfalls, 2) identify biodiversity erosion drivers impacting oceanic indigenous assemblages under global change for conservation management purpose, 3) use species distribution and abundance data in model-based studies of environmental change in different islands, 4) contribute to clarifying the potential occurrence of an "insect decline" in Azores and identifying the spatial and temporal invasion patterns of exotic arthropod species, 5) contribute with temporal data to re-assess the Red-list status of Azorean endemic arthropods and 6) perform studies about the relationship between diversity (taxonomic, functional and phylogenetic) and ecosystem function.
The project SLAM (Long Term Ecological Study of the Impacts of Climate Change in the natural forest of Azores) is described in detail.Seasonal distribution and abundance data of Azorean spiders, based on a long-term study undertaken between 2012 and 2019 in two Azorean Islands (Terceira and Pico), is presented. A total of 14979 specimens were collected, of which 6430 (43%) were adults. Despite the uncertainty of juvenile identification, juveniles are also included in the data presented in this paper, since the low diversity allows a relatively precise identification of this life-stage in Azores.A total of 57 species, belonging to 50 genera and 17 families, were recorded from the area, which constitutes baseline information of spiders from the studied sites for future long-term comparisons. Linyphiidae were the richest and most abundant family, with 19 (33%) species and 5973 (40%) specimens. The ten most abundant species are composed mostly of endemic or native non-endemic species and only one exotic species (
(Blackwall, 1852)). Those ten most abundant species include 84% of all sampled specimens and are clearly the dominant species in the Azorean native forests.
L. Koch, 1872 was firstly reported from Terceira and Pico Islands,
Clerck, 1757 was firstly reported from Terceira Island,
(Sundevall, 1830) and
(Blackwall, 1867) were firstly reported from Pico Island.This publication contributes not only to a better knowledge of the arachnofauna present in native forests of Terceira and Pico, but also to understand the patterns of abundance and diversity of spider species, both seasonally and between years.
During the last few centuries oceanic island biodiversity has been drastically modified by human-mediated activities. These changes have led to the increased homogenization of island biota and to a ...high number of extinctions lending support to the recognition of oceanic islands as major threatspots worldwide. Here, we investigate the impact of habitat changes on the spider and ground beetle assemblages of the native forests of Madeira (Madeira archipelago) and Terceira (Azores archipelago) and evaluate its effects on the relative contribution of rare endemics and introduced species to island biodiversity patterns. We found that the native laurel forest of Madeira supported higher species richness of spiders and ground beetles compared with Terceira, including a much larger proportion of indigenous species, particularly endemics. In Terceira, introduced species are well-represented in both terrestrial arthropod taxa and seem to thrive in native forests as shown by the analysis of species abundance distributions (SAD) and occupancy frequency distributions (OFD). Low abundance range-restricted species in Terceira are mostly introduced species dispersing from neighbouring man-made habitats while in Madeira a large number of true rare endemic species can still be found in the native laurel forest. Further, our comparative analysis shows striking differences in species richness and composition that are due to the geographical and geological particularities of the two islands, but also seem to reflect the differences in the severity of human-mediated impacts between them. The high proportion of introduced species, the virtual absence of rare native species and the finding that the SADs and OFDs of introduced species match the pattern of native species in Terceira suggest the role of man as an important driver of species diversity in oceanic islands and add evidence for an extensive and severe human-induced species loss in the native forests of Terceira.
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