Use of insect rarity for biotope prioritisation: the tenebrionid beetles of the Central Apennines (Italy)

Insect conservation has been traditionally based mainly on the identification of priority biotopes. One of the most commonly used criteria for biotope prioritisation is the occurrence of priority species, hence the need for measures of species vulnerability. In this paper a two-step protocol for biotope prioritisation is proposed. Firstly, insect species vulnerability is estimated from rarity measures that can be easily derived from basic data. Then, these values of vulnerability are used to rank biotopes. The method was applied here to the tenebrionid beetles of the Central Apennines, a montane area of high conservation concern for South Europe. Their use in this paper is an example of the use of data hidden in museum collections for analyses dealing with traditionally overlooked insect groups. Most of conservation decisions for Mediterranean mountains are biased towards certain vertebrates. Although current management practices in the preserves of the study area are generally consistent with the conservation of tenebrionids (especially woodland protection, which is essential for arboreal species), results provided in this study stress the importance of preserving also open biotopes, which are crucial for many taxa, including several endemic forms. Moreover, there is indication that preservation of such kind of biotopes would be important also for vertebrates. The approach for biotope prioritisation based on species rarity was here applied to a single animal taxon, but it can be easily extended to other insect groups, in order to obtain a more general view of the relative importance of different biotope types for Apennine conservation.     

The origins of the arthropod nervous system: insights from the Onychophora

A revision of evolutionary relationships of the Arthropoda has provided fresh impetus to tracing the origins of the nervous system of this group of animals: other members of the Ecdysozoa possess a markedly different type of nervous system from both the arthropods and the annelid worms, with which they were previously grouped. Given their status as favoured sister taxon of the arthropods, Onychophora (velvet worms) are a key group for understanding the evolutionary changes that have taken place in the panarthropod (Arthropoda + Onychophora + Tardigrada) lineage. This article reviews our current knowledge of the structure and development of the onychophoran nervous system. The picture that emerges from these studies is that the nervous system of the panarthropod ancestor was substantially different from that of modern arthropods: this animal probably possessed a bipartite, rather than a tripartite brain; its nerve cord displayed only a limited degree of segmentation; and neurons were more numerous but more uniform in morphology than in living arthropods. These observations suggest an evolutionary scenario, by which the arthropod nervous system evolved from a system of orthogonally crossing nerve tracts present in both a presumed protostome ancestor and many extant worm-like invertebrates, including the onychophorans.     

Biotope prioritisation in the Central Apennines (Italy): species rarity and cross-taxon congruence

The conservation status of invertebrates is usually lesser known than that of vertebrates, and strategies to identify biotopes to preserve invertebrate diversity are typically based on a single surrogate taxon, or even on the use of vertebrates as surrogates. Aim of this research is to illustrate a method for biotope prioritisation that can be easily adapted to different animal groups and geographical contexts. A two-step protocol for biotope prioritisation is proposed on the basis of a multidimensional characterisation of species vulnerability. Firstly, species vulnerability is estimated from rarity measures which include geographical range, abundance and biotope specialisation. Then, these values of vulnerability are used to rank biotopes. The method was applied here to the tenebrionid beetles, the butterflies, the birds and the mammals of the Central Apennines, a montane area of high conservation concern for South Europe. This study provides evidence for the importance of including insects in conservation decisions, because vertebrates are poor surrogates for insects. Conservation efforts in the reserves included in the study area are mostly focused on vertebrates, for which woodlands are considered particularly important. However high altitude open biotopes are crucial for both tenebrionids and butterflies, and preservation of such kind of biotopes would be beneficial also for vertebrates. The approach applied here demonstrates that (1) vertebrates are poor surrogates for insects, and (2) measures of species rarity, typically used in vertebrate conservation, can be obtained also for insects, for which a veritable amount of data are hidden in specialised literature and museum collections.     

Effect of island geological age on the arthropod species richness of Azorean pastures

Species richness of six pasture arthropod assemblages (total arthropod species, total herbivore species, sucking and chewing herbivores, total predatory species and spiders) were regressed against several geographical variables (area, distance from the nearest mainland, maximum elevation and geological age of the islands) of three Azorean islands (S. Maria, Terceira and Pico). The species were sampled by the fixed-quadrat size sampling method and the results obtained are consistent with the geological age hypothesis, i.e. the species richness of the six indigenous arthropod assemblages increases with the geological age of the islands, both at local and regional scales. Higher values of indigenous and endemic species richness were consistendy found on the older island (S. Maria), and the lowest values on the most recent island (Pico). Moreover, when considering the age of Faial (an older island probably once connected with Pico) as a estimate of the age of Pico, correlations between species richness and island age were improved, thereby strengthening the relationship. The older island (S. Maria) has more specialized herbivores and a greater proportion of herbivores in relation to predatory arthropods. Ecological and biogeographical studies in the Azores should take into account the effects of the time each island has been available for colonization and evolution.     

Time, area and isolation: factors driving the diversification of Azorean arthropods

Aim R. J. Whittaker et al. recently proposed a ‘general dynamic model of oceanic island biogeography’ (GDM), providing a general explanation of island biodiversity patterns by relating fundamental biogeographical processes – speciation, immigration, extinction – to area (A) and time (T; maximum island geological age). We adapt their model, which predicts a positive relationship with area combined with a humped relationship to time (designated the ATT² model), to study the factors promoting diversification on the Azores for several arthropod groups. Location The Azorean archipelago (North Atlantic; 37–40° N, 25–31° W). Methods We use the number of single‐island endemics (SIEs) as a measure of diversification, to evaluate four different predictions for the variation in SIEs between different islands, derived from the GDM theory and our knowledge of the fauna and history of the Azores. We calculated the number of SIEs for seven out of the nine Azorean islands and six groups of species (all arthropods, beetles, cavernicolous and non‐cavernicolous species, and taxa with high and low dispersal abilities). Several variables accounting for island characteristics (area, geological age, habitat diversity and isolation) and generalized linear models were used to evaluate the reliability of each prediction. Results A linear and positive relationship between SIEs and an AT (area + time) model was the most parsimonious explanation for overall arthropod diversification. However, cavernicolous species showed the opposite pattern (more SIEs inhabiting the youngest islands). Also, isolation was an important predictor of diversification for all groups except for the species with high dispersal ability; while the former were negatively related to the distance from the main source of colonizing lineages (Santa Maria island in most cases), the latter were related to area. Dispersal ability was also a key factor affecting the diversification of most groups of species. Main conclusions In general, the diversification of Azorean arthropods is affected by age, area and isolation. However, different groups are affected by these factors in different ways, showing radically different patterns. Although the ATT² model fails to predict the diversification pattern of several groups, it provides a framework for integrating these deviations into a general theory. Further improvements of the GDM theory need to take into account the particular traits of each group and the role of isolation in shaping island diversity.     

Trophic segregation of Falkland Islands seabirds: insights from stable isotope analysis

Seabird colonies provide rare opportunities to study trophic segregation in an entire bird community. We here present data on nitrogen and carbon isotope ratios of eight species of seabirds from New Island, Falkland Islands, and compare trophic levels (TL) and foraging distributions. We included adult feathers representing the interbreeding season, as well as chick feathers or down representing the breeding season. The stable isotope ratios indicated differences in feeding areas and TLs between species, consistent with the data of previous conventional diet analyses and observations at sea. We further reviewed conventional and stable isotope seabird community studies calculating the means and ranges of TLs observed across these studies. The mean TL (3.7) of the seabird community on New Island was at the lower end of the mean value range (3.5–4.5), but not significantly different, from the reviewed seabird communities. Seabirds on New Island had a range of 1.3 TLs, which is on the upper end of ranges within a community (0.4–1.5), indicating strong trophic structuring.     

Body size, rarity, and phylogenetic community structure: insights from diving beetle assemblages of Alberta

Although it is implicit that interactions between species depend on their traits, studies on the probability of finding related species in a community are in their infancy. Community composition and species richness of predaceous diving beetles (Dytiscidae: Coleoptera) have been used as indicators of freshwater ecosystem function yet no incorporation of phylogenetic relationships of coexisting dytiscids has been attempted to date. Improved knowledge of phylogenetic relationships and phylogenetic community structure analysis methods may provide additional insight into the relationships between community composition and species richness, thus impacting our interpretation of aquatic indicator species metrics. Here, we use museum records of dytiscid beetles in 53 lakes of Alberta, Canada to: (1) compile a supertree of dytiscid beetles that live in the province, (2) examine whether coexisting dytiscids tend to be more or less related than expected by chance, and (3) examine whether phylogenetic structuring depends on species richness or mean size of coexisting species. We find that, although the majority of dytiscid assemblages exhibited phylogenetic clustering, the extent to which this occurred depended on the mean size of dytiscids. We discuss the potential mechanisms and implications of the observed patterns in phylogenetic clustering, along with data that would further improve our understanding of community dynamics in dytiscid beetles.     

Separating the wheat from the chaff: a prioritisation pipeline for the analysis of metabolomics datasets

Liquid Chromatography Mass Spectrometry (LC-MS) is a powerful and widely applied method for the study of biological systems, biomarker discovery and pharmacological interventions. LC-MS measurements are, however, significantly complicated by several technical challenges, including: (1) ionisation suppression/enhancement, disturbing the correct quantification of analytes, and (2) the detection of large amounts of separate derivative ions, increasing the complexity of the spectra, but not their information content. Here we introduce an experimental and analytical strategy that leads to robust metabolome profiles in the face of these challenges. Our method is based on rigorous filtering of the measured signals based on a series of sample dilutions. Such data sets have the additional characteristic that they allow a more robust assessment of detection signal quality for each metabolite. Using our method, almost 80% of the recorded signals can be discarded as uninformative, while important information is retained. As a consequence, we obtain a broader understanding of the information content of our analyses and a better assessment of the metabolites detected in the analyzed data sets. We illustrate the applicability of this method using standard mixtures, as well as cell extracts from bacterial samples. It is evident that this method can be applied in many types of LC-MS analyses and more specifically in untargeted metabolomics.

     

Separating the wheat from the chaff: a prioritisation pipeline for the analysis of metabolomics datasets

Liquid Chromatography Mass Spectrometry (LC-MS) is a powerful and widely applied method for the study of biological systems, biomarker discovery and pharmacological interventions. LC-MS measurements are, however, significantly complicated by several technical challenges, including: (1) ionisation suppression/enhancement, disturbing the correct quantification of analytes, and (2) the detection of large amounts of separate derivative ions, increasing the complexity of the spectra, but not their information content. Here we introduce an experimental and analytical strategy that leads to robust metabolome profiles in the face of these challenges. Our method is based on rigorous filtering of the measured signals based on a series of sample dilutions. Such data sets have the additional characteristic that they allow a more robust assessment of detection signal quality for each metabolite. Using our method, almost 80% of the recorded signals can be discarded as uninformative, while important information is retained. As a consequence, we obtain a broader understanding of the information content of our analyses and a better assessment of the metabolites detected in the analyzed data sets. We illustrate the applicability of this method using standard mixtures, as well as cell extracts from bacterial samples. It is evident that this method can be applied in many types of LC-MS analyses and more specifically in untargeted metabolomics.     

Stoichiometry of actual vs. potential predator–prey interactions: insights into nitrogen limitation for arthropod predators

Literature‐compiled data sets demonstrate wide interspecific variation in nitrogen content among terrestrial arthropods and raise the possibility of nitrogen (N) limitation for predatory species. It remains unclear, however, whether the same disparities between N supply and demand that appear in literature compilations also exist in particular ecological communities. To address this uncertainty, we compared arthropod predator–prey stoichiometries derived from a compiled database with those from a natural Spartina saltmarsh community. Separate assessments of potential N‐limitation were made for arthropod predators feeding on herbivores and for intraguild predators feeding on intraguild prey. Relative to the compiled database, saltmarsh consumer–resource interactions exhibited increased disparity between N‐content of herbivores and N‐demand by predators. The high N content of saltmarsh arachnids relative to predatory insects at large may contribute to the supply‐demand disparity. Whether N‐limitation of terrestrial arthropod predators is widespread in the marsh, and in nature in general, depends sensitively on the predatory species’ gross growth efficiencies for N and carbon. Obtaining hard empirical data for these efficiency parameters should be a research goal.     

Drivers of extinction: the case of Azorean beetles

Oceanic islands host a disproportionately high fraction of endangered or recently extinct endemic species. We report on species extinctions among endemic Azorean beetles following 97% habitat loss since AD 1440. We infer extinctions from historical and contemporary records and examine the influence of three predictors: geographical range, habitat specialization and body size. Of 55 endemic beetle species investigated (out of 63), seven can be considered extinct. Single-island endemics (SIEs) were more prone to extinction than multi-island endemics. Within SIEs restricted to native habitat, larger species were more extinction-prone. We thus show a hierarchical path to extinction in Azorean beetles: species with small geographical range face extinction first, with the larger bodied ones being the most threatened. Our study provides a clear warning of the impact of habitat loss on island endemic biotas.     

Public preferences for health care: prioritisation in the United Kingdom

The Government in the UK is encouraging consumerism within health care and is requiring Health Authorities to consult with the public on prioritisation of resources. Public consultation within the National Health Service (NHS) has had limited success in the past. Many of the techniques used are flawed. Despite the limited scope of the public surveys conducted so far, a number of themes have emerged: a willingness to pay for experimental, 'high-tech' life-saving treatments rather than more cost-effective treatments which will improve quality of life, which are more likely to maximise utility from the scarce resources available; preference for treating the young rather than the old; preference for treating patients with dependents (e.g. spouse, children) rather than those who have none; a willingness to discriminate against those patients who were partially responsible for their illness due to choice of 'unhealthy' lifestyle (e.g. smoking cigarettes, drinking excess alcohol). These public preferences raise ethical problems. For example, is it just to spend more on heroic treatments which are likely to fail? Is there a right to health care irrespective of whether you have had 'a fair innings' or whether a patient is in part responsible for their illness due to an unhealthy lifestyle? If there are ethical concerns about these preferences, should health authorities consult with the public at all? Is human life and suffering incommensurable, and hence is it impossible to prioritise anyway? Some of the ethical consequences of using empirical data on public preferences are discussed.     

Footprint on a coprolite: A rarity from the Eocene of Vietnam

A rare ichnological example of a crocodilian footprint on an Eocene crocodilian coprolite has been discovered. The enigmatic specimen, measuring 4 cm in length, was found among 100 coprolites at the Na Duong coal mine in Lang Son province, northern Vietnam. Almost all the discovered coprolites were ascribed to a potential crocodilian producer, suggesting the existence of an ancient lacustrine environment. Furthermore, neoichnological analyses of extant crocodilian footprints also corroborate the inference of a crocodilian as the producer of the footprint. We concluded that the 4-cm-long Na Duong fingerprints most likely belonged to a 2-m-long crocodilian whose IV or V finger of the right manus might have made the fingerprints. Since no other records of foot traces or trackways were identified during the fieldwork in the fall of 2018, the Na Duong fossil site should be investigated for further ichnofossil analysis.     

Using range filling rather than prevalence of invasive plant species for management prioritisation: the case of Spathodea campanulata in the Society Islands (South Pacific)

Biological invasion science lacks standardised measures of invasion success that would provide effective prioritisation of invasive species and invaded areas management. Prevalence (area of occupancy) of invasive species is often used as proxy of their success but this metric ignores the extent to which a species fills its potential distribution. This study aims to estimate the performance of invasive tree species by computing the ratio between the compressed canopy area (CCA), assessed through remote sensing, and their potential distribution, estimated using invasive species distribution modelling. This index of ‘range filling’ (RF) has applicability to a broad set of invasive plant species in any biome. A case study is provided using the invasive African tulip tree Spathodea campanulata (Bignoniaceae) on three small tropical oceanic islands (South Pacific) exhibiting different invasion levels to test for differences between CCA and RF. The results show that the RF of Spathodea campanulata varied within islands depending on elevation but not proportionally to the CCA of the species. Another key result was that the RF of the species and its CCA provided different between-island perspectives on the invasions and lead to distinct ranking among islands to prioritise for management. Therefore, managers should disregard species’ prevalence as a measure of success and rather weight it with potential distribution to quantify how an invader is performing in a given environment.     

The terrestrial arthropod fauna of the Byers Peninsula,Livingston Island,South Shetland Islands

We report the first detailed study of the terrestrial invertebrate fauna of the Byers Peninsula SSSI, Livingston Island, South Shetland Islands. Fourteen micro-arthropod taxa (10 Acari, four Collembola) and two Diptera are recorded, including the first record of the mite Edwardzetes dentifer from the maritime Antarctic. The first record of the midge Belgica antarctica from neighbouring Snow Island is also given. Population composition and density were described in samples from a wide range of terrestrial and freshwater habitats. There was no strong relationship between habitat and microarthropod species occurrence, although comparison of completely vegetated and more stony sites revealed greater population densities at the vegetated sites, and different species proportions at each. Some individual samples contained a wide range of species with none achieving numerical dominance, whilst others from superficially similar sites were dominated by one species. Dipterans were limited to a small number of lakes, streams and seepage areas, where they were sometimes abundant. Population density data and species occurrence are compared with previously published studies from the maritime Antarctic and elsewhere.     

The rarity concept and the commonness of rarity in freshwater zooplankton

1. Regional distribution, frequency of occurrence and relative abundance were scored in 2467 Norwegian lakes for all the recorded 130 species of crustacean zooplankton. The majority of species were rare in the sense that 65% of species were recorded in fewer than 10% of localities. Only six species were recorded in more than 50% of localities, and the median number of species in a given locality was 14 (i.e. 10% of the total species pool). 2. Abundances of all species were scored according to the fraction of lakes in which they were recorded, their geographical range of distribution, and their numerical abundance. Typically the most rare species were rare by all three criteria, and vice versa for the common species, pointing to rarity as an inherent property of some species. For some species this rarity reflects being on the edge of their distributional range, while for others rarity seems to be a consequence of their life cycle strategies. 3. Some of the truly rare species have high dispersal rates and high colonization abilities, but are rapidly replaced by other species. Others are confined to specific habitats, often highly eutrophic, pointing to highly specialized niche adaptations. 4. A major cause for the few truly common species seems to be the limited number of species that are able to coexist within a given locality, reflecting ‘the ghost of competition past’ and predation pressure. 5. While species composition and species richness may reflect colonization abilities and stochastic events, the presence or absence of species is not only a random lottery but also a consequence of species‐specific attributes.     

Factors influencing the distribution of Azorean mountain vegetation: implications for nature conservation

In order to understand the main ecological factors that influence the distribution of mountain vegetation in Azores and to use it as a model for nature conservation, a study on Santa Bárbara Mountain on Terceira Island was performed. A multivariate analysis was done on vegetation data and the abiotic factors governing the vegetation pattern are discussed. The major factors contributing to the observed vegetation patterns are wind exposure, soil water saturation and historical human impacts. The vegetation analysis showed that Azorean Mountain habitats have a high variation of vegetation structure as a result of the habitat heterogeneity and the extreme ecological conditions. Using the co-dominant plants as indicator species, eco-zones are distinguished, characterized by the combination of different ecological–human factors which contribute to the distribution of specific types of natural vegetation communities. The studies of vegetation distribution give important information about the relation between vegetation and landscape that is essential for conservation, restoration proposals and landscape planning. Azores terrestrial Natura 2000 sites are largely dominated by mountain vegetation, therefore this work provides essential background data for the management plans of these sites.     

A new cytotoxic tambjamine alkaloid from the Azorean nudibranch Tambja ceutae

The chemical investigation of Azorean nudibranch mollusk Tambja ceutae led us to isolate a new member of the tambjamine family, tambjamine K (1). The bryozoan Bugula dentata, prey of the nudibranch, was also analyzed and found to contain compound 1 in very small amounts together with known blue pigment 2 and tambjamines A (3) and B (4). The structure of tambjamine 1 was elucidated by the interpretation of the spectroscopic data as well as by the comparison with related compounds. Compounds 1 and 2 possess antiproliferative activity, in particular, tambjamine K (1) displayed high cytotoxicity against both tumor and non-tumor mammalian cells.