Paulo A. V. Borges

Detection and Characterization of Wolbachia Infections in Natural Populations of Aphids: Is the Hidden Diversity Fully Unraveled?

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.

Ranking protected areas in the Azores using standardised sampling of soil epigean arthropods

Nineteen areas in seven of the nine Azorean islands were evaluated for species diversity and rarity based on soil epigean arthropods. Fifteen out of the 19 study areas are managed as Natural Forest Reserves and the remaining four were included due to their importance as indigenous forest cover. Four of the 19 areas are not included in the European Conservation network, NATURA 2000. Two sampling replicates were run per study area, and a total of 191 species were collected; 43 of those species (23%) are endemic to the archipelago and 12 have yet to be described. To produce an unbiased multiple-criteria index (importance value for conservation, IV-C) incorporating diversity and rarity based indices, an iterative partial multiple regression analysis was performed. In addition, an irreplaceability index and the complementarity method (using both optimisation and heuristic methods) were used for priority-reserves analyses. It was concluded that at least one well-managed reserve per island is absolutely necessary to have a good fraction of the endemic arthropods preserved. We found that for presence/absence data the suboptimal complementarity algorithm provides solutions as good as the optimal algorithm. For abundance data, optimal solutions indicate that most reserves are needed if we want that at least 50% of endemic arthropod populations are represented in a minimum set of reserves. Consistently, two of the four areas not included in the NATURA 2000 framework were considered of high priority, indicating that vascular plants and bird species used to determine NATURA 2000 sites are not good surrogates of arthropod diversity in the Azores. The most irreplaceable reserves are those located in older islands, which indicates that geological history plays an important role in explaining faunal diversity of arthropods in the Azores. Based both on the uniqueness of species composition and high species richness, conservation efforts should be focused on the unmanaged Pico Alto region in the archipelago’s oldest island, Santa Maria.

Harmonia axyridis: What will stop the invader?

In recent years Harmonia axyridis (Pallas, 1773) (Coleoptera: Coccinellidae) has become a very popular insect among biological control practitioners and scientists, not only for its potential to be an efficient biological control agent but also because it is considered invasive. Individuals of this species were deliberately introduced into several countries for biological control of different arthropods pests. However the predator itself became an invasive species, affecting the dynamics and composition of several guilds through direct or indirect interactions with established species, including intraguild predation. In this paper we discuss the reasons why the species has a high invasiveness and what are the limits to invasion by this species. It is not clear if the invasiveness of the beetle is linked to its biological, ecological and behavioural abilities, or to other factors such as invasibility and interactions between the invaders, the noninvaders, and the habitat, which may in part explain the reasons of its success and help us to answer the question “what will stop the invader?” We also discuss the reason for the absence of the predator in the Azores islands. Despite the intentional introduction of H. axyridis in the Azores and the high number of individuals released, there are no records of this species in the wild, despite recent extensive sampling effort. In this paper we discuss the reasons for the apparent failure or the delay in establishment of the predator. One factor which may hamper the establishment of H. axyridis in some of the Azores islands is the absence of winter environmental conditions, mainly the temperature which is seldom lower than 12°C, essential for the induction of diapause. The lack of success in the establishment could be also related to functional diversity saturation, that is species saturation and competitive exclusion of H. axyridis by other previously established species may be operating.

A spatial scale assessment of habitat effects on arthropod communities of an oceanic island

Most habitats in the Azores have undergone substantial land-use changes and anthropogenic disturbance during the last six centuries. In this study we assessed how the richness, abundance and composition of arthropod communities change with: (1) habitat type and (2) the surrounding land-use at different spatial scales. The research was conducted in Terceira Island, Azores. In eighty-one sites of four different habitat types (natural and exotic forests, semi-natural and intensively managed pastures), epigaeic arthropods were captured with pitfall traps and classified as endemic, native or introduced. The land-use surrounding each site was characterized within a radius ranging from 100 to 5000 m. Non-parametric tests were used to identify differences in species richness, abundance and composition between habitat types at different spatial scales. Endemic and native species were more abundant in natural forests, while introduced species were more abundant in intensively managed pastures. Natural forests and intensively managed pastures influenced arthropod species richness and composition at all spatial scales. Exotic forests and semi-natural pastures, however, influenced the composition of arthropod communities at larger scales, promoting the connectivity of endemic and native species populations. Local species richness, abundance and composition of arthropod communities are mostly determined by the presence of nearby natural forests and/or intensively managed pastures. However, semi-natural pastures and exotic forests seem to play an important role as corridors between natural forests for both endemic and native species. Furthermore, exotic forests may serve as a refuge for some native species.

Integrating landscape disturbance and indicator species in conservation studies.

Successful conservation plans are conditioned by our ability to detect anthropogenic change in space and time and various statistical analyses have been developed to handle this critical issue. The main objective of this paper is to illustrate a new approach for spatial analysis in conservation biology. Here, we propose a two-step protocol. First, we introduce a new disturbance metric which provides a continuous measure of disturbance for any focal communities on the basis of the surrounding landscape matrix. Second, we use this new gradient to estimate species and community disturbance thresholds by implementing a recently developed method called Threshold Indicator Taxa ANalysis (TITAN). TITAN detects changes in species distributions along environmental gradients using indicators species analysis and assesses synchrony among species change points as evidence for community thresholds. We demonstrate our method with soil arthropod assemblages along a disturbance gradient in Terceira Island (Azores, Portugal). We show that our new disturbance metric realistically reflects disturbance patterns, especially in buffer zones (ecotones) between land use categories. By estimating species disturbance thresholds with TITAN along the disturbance gradient in Terceira, we show that species significantly associated with low disturbance differ from those associated with high disturbance in their biogeographical origin (endemics, non-endemic natives and exotics) and taxonomy (order). Finally, we suggest that mapping the disturbance community thresholds may reveal areas of primary interest for conservation, since these may host indigenous species sensitive to high disturbance levels. This new framework may be useful when: (1) both local and regional processes are to be reflected on single disturbance measures; (2) these are better quantified in a continuous gradient; (3) mapping disturbance of large regions using fine scales is necessary; (4) indicator species for disturbance are searched for and; (5) community thresholds are useful to understand the global dynamics of habitats.

Species Distribution Models Do Not Account for Abundance: The Case of Arthropods on Terceira Island

The objective of this study is to investigate whether presence/absence models can be used as surrogates of arthropod abundance, and eventually under which circumstances such surrogacy is guaranteed. Presence/absence data for 48 arthropod species from Terceira Island were modelled using artificial neural networks. Probabilities of occurrence were correlated with abundance data from a standardized arthropod survey programme. Although a tendency was found for vagile species to show relationships, only nine species showed significant positive correlations between probability of presence and abundance. Five of these were exotic spider species with high abundances and wide distributions in several human-modified habitats. The patchy distribution of pristine habitats, the capacity to reach them and the probable low dependence on limiting resources, other than food, enhance the relationship. A lack of significant correlations for the majority of the species may be due to historical events, inappropriate scale, demographic controls of density, or the incapacity of presence/absence models to account for environmental suitability. The difficulty to identify a priori the species for which the relationship will hold advises against the use of species distribution models as surrogates of arthropod abundance.

Functional biogeography of oceanic islands and the scaling of functional diversity in the Azores

Significance Biogeographic theory builds upon a long history of analyzing species-diversity patterns of remote islands, but no previous studies have attempted to investigate corresponding patterns in functional traits on islands. Our analyses of functional diversity (FD) for spiders and beetles in the Azorean archipelago reveal that FD increases with species richness, which, in turn scales with island area regardless of the taxa and distributional group considered (endemics, natives, and exotics). Our results also support the hypothesis that each group contributes to FD in proportion to their species richness and that, being dominant, exotic species have significantly extended the realized trait space of the Azorean islands. Further analyses in other archipelagos are needed to establish whether our findings are representative of oceanic islands. Analyses of species-diversity patterns of remote islands have been crucial to the development of biogeographic theory, yet little is known about corresponding patterns in functional traits on islands and how, for example, they may be affected by the introduction of exotic species. We collated trait data for spiders and beetles and used a functional diversity index (FRic) to test for nonrandomness in the contribution of endemic, other native (also combined as indigenous), and exotic species to functional-trait space across the nine islands of the Azores. In general, for both taxa and for each distributional category, functional diversity increases with species richness, which, in turn scales with island area. Null simulations support the hypothesis that each distributional group contributes to functional diversity in proportion to their species richness. Exotic spiders have added novel trait space to a greater degree than have exotic beetles, likely indicating greater impact of the reduction of immigration filters and/or differential historical losses of indigenous species. Analyses of species occurring in native-forest remnants provide limited indications of the operation of habitat filtering of exotics for three islands, but only for beetles. Although the general linear (not saturating) pattern of trait-space increase with richness of exotics suggests an ongoing process of functional enrichment and accommodation, further work is urgently needed to determine how estimates of extinction debt of indigenous species should be adjusted in the light of these findings.

The subterranean fauna of a biodiversity hotspot region - Portugal: an overview and its conservation

INTRODUCTION Extensive biological studies have been made in the main karst areas around the world, namely in the eastern United States of America and in the region from the Pyrenees to Slovenia and the Dinaric karst (Culver et al., 2000; Culver & Pipan, 2009). Information about subterranean fauna in mainland Portugal is sparse and scattered along many publications mostly from the middle of the 20th century. The main thrust in the subterranean biology was given by the survey of caves made by Barros Machado during the 1940s, and by the prospection of well-dwelling crustaceans in the north of the country by researchers of the former Instituto de Zoologia “Dr. Augusto Nobre” from Porto University (Gama & Afonso, 1994). In addition, endogean habitats have been subject of considerable coleopterological

Using taxonomically unbiased criteria to prioritize resource allocation for oceanic island species conservation

Oceanic islands have been the grand stage of documented extinctions. In view of limited resources, efficient prioritization is crucial to avoid the extinction of taxa. This work lists the top 100 management priority species for the European archipelagos of the Macaronesian region (Azores, Madeira and the Canary Islands), taking into account both their protection priority and their management feasibility. Bryophytes, vascular plants, molluscs, arthropods and vertebrates were scored by species experts following two sets of criteria: (i) protection priority, including ecological value, singularity, public institutions’ management responsibilities and social value; (ii) management feasibility, including threats knowledge and control feasibility, external socio-economical support for management and biological recovery potential. Environmental managers weighted the same criteria according to their management importance. Final species scores were determined by the combination of both species valuation and criteria weighting. Vascular plants dominate the Top 100 list, followed by arthropods and vertebrates. The majority of listed taxa are endemic to one archipelago or even to a single island. The management feasibility criteria did not dictate that all taxa must be eminently endangered, as for most of the species it should be relatively easy to control threats. The main advantages of this process are the independent participation of scientists and conservation managers, the inclusion of criteria on both protection priority and management feasibility and the taxonomically unbiased nature of the process. This study provides a potentially useful biodiversity conservation tool for the Macaronesian archipelagos that could be readily implemented by the respective regional governments in future legislation.

Oceanic island biogeography through the lens of the general dynamic model: Assessment and prospect

The general dynamic model of oceanic island biogeography (GDM) has added a new dimension to theoretical island biogeography in recognizing that geological processes are key drivers of the evolutionary processes of diversification and extinction within remote islands. It provides a dynamic and essentially non‐equilibrium framework generating novel predictions for emergent diversity properties of oceanic islands and archipelagos. Its publication in 2008 coincided with, and spurred on, renewed attention to the dynamics of remote islands. We review progress, both in testing the GDMs predictions and in developing and enhancing ecological–evolutionary understanding of oceanic island systems through the lens of the GDM. In particular, we focus on four main themes: (i) macroecological tests using a space‐for‐time rationale; (ii) extensions of theory to islands following different patterns of ontogeny; (iii) the implications of GDM dynamics for lineage diversification and trait evolution; and (iv) the potential for downscaling GDM dynamics to local‐scale ecological patterns and processes within islands. We also consider the implications of the GDM for understanding patterns of non‐native species diversity. We demonstrate the vitality of the field of island biogeography by identifying a range of potentially productive lines for future research.