The BALA project: A pioneering monitoring of Azorean forest invertebrates over two decades (1999-2022).

Globally, there is a concerning decline in many insect populations, and this trend likely extends to all arthropods, potentially impacting unique island biota. Native non-endemic and endemic species on islands are under threat due to habitat destruction, with the introduction of exotic, and potentially invasive, species, further contributing to this decline. While long-term studies of plants and vertebrate fauna are available, long-term arthropod datasets are limited, hindering comparisons with better-studied taxa. The Biodiversity of Arthropods of the Laurisilva of the Azores (BALA) project has allowed gathering comprehensive data since 1997 in the Azorean Islands (Portugal), using standardised sampling methods across islands. The dataset includes arthropod counts from epigean (pitfall traps) and canopy-dwelling (beating samples) communities, enriched with species information, biogeographic origins, and IUCN categories. Metadata associated with the sample protocol and events, like sample identifier, archive number, sampled tree species, and trap type are also recorded. The database is available in multiple formats, including Darwin Core, which facilitates the ecological analysis of pressing environmental concerns, such as arthropod population declines and biological invasions.

Assessing nature exposure: a study on the reliability and validity of a Portuguese version of the nature exposure scale.

Introduction: Engaging with nature has been widely acknowledged for its positive impact on well-being. Traditionally, assessments of nature exposure focus on estimating green space exposure and outdoor time. However, the Nature Exposure Scale (NES) offers a unique approach by evaluating the quality of nature experiences, encompassing both deliberate and spontaneous encounters. Methods: This study aimed to assess the validity and reliability of the NES in the context of Portugal. Exploratory Factor Analysis (PCA) and Confirmatory Factor Analysis (CFA) were employed to examine the underlying structure of the scale. Additionally, reliability assessments, along with tests of convergent and divergent validity, were conducted. Data were collected from a sample of 558 adults and 241 adolescents between 2016 and 2018. Results: The findings revealed strong internal consistency of the NES, supported by acceptable correlation values and robust factor loadings within a unidimensional model. The scale effectively predicted variations in nature exposure across diverse professional activities. Discussion: In summary, the 4-item NES emerged as a reliable tool for assessing nature exposure in various settings, effectively bridging a gap in the Portuguese context. The scale demonstrated potential for cross-cultural research and was particularly adept at predicting nature exposure in different professional contexts. The results suggest that the NES can enhance our understanding of the impact of nature on well-being in diverse cultural settings. Conclusion: The study underscores the reliability of the NES in assessing nature exposure in Portugal, paving the way for further exploration in Portuguese-speaking regions. The scale holds promise for advancing research on the relationship between nature and well-being, contributing valuable insights across different cultural contexts.

SLAM Project - Long Term Ecological Study of the Impacts of Climate Change in the natural forest of Azores: IV - The spiders of Terceira and Pico Islands (2019-2021) and general diversity patterns after ten years of sampling.

Background: Long-term studies are key to understand the drivers of biodiversity erosion, such as land-use change and habitat degradation, climate change, invasive species or pollution. The long-term project SLAM (Long Term Ecological Study of the Impacts of Climate Change in the natural forest of Azores) started in 2012 and focuses on arthropod monitoring, using SLAM (Sea, Land and Air Malaise) traps, aiming to understand the impact of the drivers of biodiversity erosion on Azorean native forests (Azores, Portugal). This is the fourth contribution including SLAM project data and the second focused on the spider fauna (Arachnida, Araneae) of native forests on two islands (Pico and Terceira). In this contribution, we describe data collected between 2019 and 2021 and we analyse them together with a previously published database that covered the 2012-2019 period, in order to describe changes in species abundance patterns over the last ten years. New information: We present abundance data of Azorean spider species for the 2019-2021 period in two Azorean Islands (Terceira and Pico). We also present analyses of species distribution and abundance of the whole sampling period. In the period of 2019-2021, we collected a total of 5110 spider specimens, of which 2449 (48%) were adults. Most juveniles, with the exception of some exotic Erigoninae, were also included in the data presented in this paper, since the low diversity of spiders in the Azores allows a relatively precise species-level identification of this life-stage. We recorded a total of 45 species, belonging to 39 genera and 16 families. The ten most abundant species were composed mostly of endemic or native non-endemic species and only two exotic species (Tenuiphantestenuis (Blackwall, 1852) and Dysderacrocata C. L. Koch, 1838). They included 4308 individuals (84%) of all sampled specimens and were the dominant species in Azorean native forests. The family Linyphiidae was the richest and most abundant taxon, with 15 (33%) species and 2630 (51%) specimens. We report Cheiracanthiummildei L. Koch, 1864, a non-native species, from Pico Island for the first time. We found no new species records on Terceira Island. This publication contributes to increasing the baseline information for future long-term comparisons of the spiders on the studied sites and the knowledge of the arachnofauna of the native forests of Terceira and Pico, in terms of species abundance, distribution and diversity across seasons for a 10 years period.

Modelling the distribution of Mustela nivalis and M. putorius in the Azores archipelago based on native and introduced ranges.

The aims of this study were to predict the potential distribution of two introduced Mustelidae, Mustela nivalis and M. putorius in the Azores archipelago (Portugal), and evaluate the relative contribution of environmental factors from native and introduced ranges to predict species distribution ranges in oceanic islands. We developed two sets of Species Distribution Models using MaxEnt and distribution data from the native and introduced ranges of the species to project their potential distribution in the archipelago. We found differences in the predicted distributions for the models based on introduced and on native occurrences for both species, with different most important variables being selected. Climatic variables were most important for the introduced range models, while other groups of variables (i.e., human-disturbance) were included in the native-based models. Most of the islands of the Azorean archipelago were predicted to have suitable habitat for both species, even when not yet occupied. Our results showed that predicting the invaded range based on introduced range environmental conditions predicted a narrower range. These results highlight the difficulty to transfer models from native to introduced ranges across taxonomically related species, making it difficult to predict future invasions and range expansion.

Fundamental research questions in subterranean biology.

Five decades ago, a landmark paper in Science titled The Cave Environment heralded caves as ideal natural experimental laboratories in which to develop and address general questions in geology, ecology, biogeography, and evolutionary biology. Although the 'caves as laboratory' paradigm has since been advocated by subterranean biologists, there are few examples of studies that successfully translated their results into general principles. The contemporary era of big data, modelling tools, and revolutionary advances in genetics and (meta)genomics provides an opportunity to revisit unresolved questions and challenges, as well as examine promising new avenues of research in subterranean biology. Accordingly, we have developed a roadmap to guide future research endeavours in subterranean biology by adapting a well-established methodology of 'horizon scanning' to identify the highest priority research questions across six subject areas. Based on the expert opinion of 30 scientists from around the globe with complementary expertise and of different academic ages, we assembled an initial list of 258 fundamental questions concentrating on macroecology and microbial ecology, adaptation, evolution, and conservation. Subsequently, through online surveys, 130 subterranean biologists with various backgrounds assisted us in reducing our list to 50 top-priority questions. These research questions are broad in scope and ready to be addressed in the next decade. We believe this exercise will stimulate research towards a deeper understanding of subterranean biology and foster hypothesis-driven studies likely to resonate broadly from the traditional boundaries of this field.

Wildlife inventory from camera-trapping surveys in the Azores (Pico and Terceira islands).

BACKGROUND: The present publication provides a dataset from five camera-trapping sampling campaigns on two islands of the Azorean archipelago (Pico and Terceira islands), between 2013-2018. This dataset was obtained as a by-product of campaigns designed for different purposes. The sampling campaigns were designed to: (i) study the ecology of introduced mammals; (ii) assess the impact of introduced mammals on native birds (Azores woodpigeon - Columba palumbus azorica and Cory's shearwater - Calonectris diomeda borealis), through nest predation; and (iii) obtain information about the impact of vertebrates on agricultural systems, particularly on Azorean traditional vineyards. A total of 258 sites and 47 nests were sampled using camera traps. These sampling campaigns provided a large data series that allowed the creation of a vertebrate wildlife inventory. NEW INFORMATION: We obtained a total of 102,095 camera-trap records, which allowed us to to identify 30 species of vertebrates: one amphibian, one reptile, 17 birds and ten mammal species. This represented 100% of the amphibians and terrestrial mammals, 58% of the breeding birds and 50% of the reptile species known for Pico and/or Terceira islands. Concerning the colonisation status of the species, we recorded 15 indigenous (native non-endemic or endemic) and three introduced bird species; all known terrestrial amphibians, reptiles and mammals in the Azores are introduced species. The data collected contribute to increasing knowledge on the distribution of vertebrate species on Pico and Terceira islands, where most existing records of some species were only available to Island level (e.g. mustelids and hedgehogs). None of the identified species was previously unknown to the study area.

Species conservation profiles of cave-dwelling arthropods from Azores, Portugal.

BACKGROUND: Azorean volcanic cave biodiversity is under considerable pressure due to ongoing threats of pollution, land use change, touristic activities or climate change. In this contribution, we present the IUCN Red List profiles of 15 cave-adapted arthropod species, endemic to the Azorean archipelago, including species belonging to the speciose genus Trechus (Carabidae), which is represented in Azores by seven species. The objective of this paper is to assess all endemic Azorean cave-adapted species and advise on possible future research and conservation actions critical for the long-term survival of the most endangered species. NEW INFORMATION: Most species have a restricted distribution (i.e. occur in one or two caves), very small extent of occurrence (EOO) and a small area of occupancy (AOO). A continuing decline in the number of mature individuals is inferred from the ongoing cave habitat degradation. The two troglobitic species of the homopteran genus Cixius are in great danger of extinction due to major land-use changes in epigean habitats above their known localities. We suggest, as future measures of conservation, the regular monitoring of the species (every five years), the creation of additional protected caves, the limitation of several aggressive activities around the caves (e.g. decreasing pasture intensification) and in some cases the creation of fences in the entrance of the most important caves.

Standardised inventories of spiders (Arachnida, Araneae) of Macaronesia I: The native forests of the Azores (Pico and Terceira islands).

BACKGROUND: The data presented here come from samples collected as part of two recent research projects (NETBIOME - ISLANDBIODIV and FCT - MACDIV) which aimed at understanding the drivers of community assembly in Macaronesian islands. We applied the sampling protocol COBRA (Conservation Oriented Biodiversity Rapid Assessment, Cardoso 2009) in sixteen 50 m x 50 m native forest plots in the Azorean Islands of Pico (6 plots) and Terceira (10 plots) to assess spider diversity. Through this publication, we contribute to the knowledge of the arachnofauna of the Azores and, more specifically, to that of the islands of Pico and Terceira. NEW INFORMATION: The collected samples yielded 8,789 specimens, of which 45% were adults (3,970) belonging to 13 families, 36 species and three morphospecies that have yet to be described. Species of the family Linyphiidae dominated the samples, with 17 species and two morphospecies that have yet to be described (48% of the taxa). Out of the identified (morpho)species, 16 were introduced, 13 Azorean endemic (three of which were undescribed) and seven native (five of them Macaronesian endemics). We report the first record of the introduced species Haplodrassus signifer and Agyneta decora in Pico Island.

Species conservation profile of moths (Insecta, Lepidoptera) from Azores, Portugal.

BACKGROUND: The few remnants of Azorean native forests harbour a unique set of endemic moths (Insecta, Lepidoptera), some of them under severe long term threats due to small sized habitats or climatic changes. In this contribution, we present the IUCN Red List profiles of 34 endemic moths to the Azorean archipelago, including species belonging to two diverse families: Noctuidae (11 species) and Crambidae (eight species). The objective of this paper is to assess all endemic Azorean moth species and advise on possible future research and conservation actions critical for the long-trem survival of the most endangered species. NEW INFORMATION: Most species have a large distribution (i.e. 58% occur in at least four islands), very large extent of occurrence (EOO) and a relatively large area of occupancy (AOO). Only nine species are single-island endemics, three of them from Flores, three from São Miguel and one from Pico, São Jorge and Faial. Most of the species also experience continuing decline in habitat quality, number of locations and subpopulations caused by the ongoing threat from pasture intensification, forestry, invasive plant species and future climatic changes. The lack of new records may indicate that one of the species previously named is extinct (Eupithecia ogilviata). Therefore, we suggest as future conservation actions: (1) a long-term species monitoring plan and (2) control of invasive species.

Conservation status of the forest beetles (Insecta, Coleoptera) from Azores, Portugal.

BACKGROUND: Island biodiversity is under considerable pressure due to the ongoing threats of invasive alien species, land use change or climate change. The few remnants of Azorean native forests harbour a unique set of endemic beetles, some of them possibly already extinct or under severe long term threat due to the small areas of the remaining habitats or climatic changes. In this contribution we present the IUCN Red List profiles of 54 forest adapted beetle species endemic to the Azorean archipelago, including species belonging to four speciose families: Zopheridae (12 species), Carabidae (11 species), Curculionidae (11 species) and Staphylinidae (10 species). NEW INFORMATION: Most species have a restricted distribution (i.e. 66% occur in only one island) and a very small extent of occurrence (EOO) and area of occupancy (AOO). Also common to most of the species is the severe fragmentation of their populations, and a continuing decline in EOO, AOO, habitat quality, number of locations and subpopulations caused by the ongoing threat from pasture intensification, forestry, invasive species and future climatic changes. Therefore, we suggest as future measures of conservation: (1) a long-term monitoring plan for the species; (2) control of invasive species; (3) species-specific conservation action for the most highly threatened species.

Dispersal ability determines the scaling properties of species abundance distributions: a case study using arthropods from the Azores.

Species abundance distributions (SAD) are central to the description of diversity and have played a major role in the development of theories of biodiversity and biogeography. However, most work on species abundance distributions has focused on one single spatial scale. Here we used data on arthropods to test predictions obtained with computer simulations on whether dispersal ability influences the rate of change of SADs as a function of sample size. To characterize the change of the shape of the SADs we use the moments of the distributions: the skewness and the raw moments. In agreement with computer simulations, low dispersal ability species generate a hump for intermediate abundance classes earlier than the distributions of high dispersal ability species. Importantly, when plotted as function of sample size, the raw moments of the SADs of arthropods have a power law pattern similar to that observed for the SAD of tropical tree species, thus we conjecture that this might be a general pattern in ecology. The existence of this pattern allows us to extrapolate the moments and thus reconstruct the SAD for larger sample sizes using a procedure borrowed from the field of image analysis based on scaled discrete Tchebichef moments and polynomials.

New insights into the genetic composition and phylogenetic relationship of wolves and dogs in the Iberian Peninsula.

This study investigates the gene pool of Portuguese autochthonous dog breeds and their wild counterpart, the Iberian wolf subspecies (Canis lupus signatus), using standard molecular markers. A combination of paternal and maternal molecular markers was used to investigate the genetic composition, genetic differentiation and genetic relationship of native Portuguese dogs and the Iberian wolf. A total of 196 unrelated dogs, including breed and village dogs from Portugal, and other dogs from Spain and North Africa, and 56 Iberian wolves (wild and captive) were analyzed for nuclear markers, namely Y chromosome SNPs, Y chromosome STR loci, autosomal STR loci, and a mitochondrial fragment of the control region I. Our data reveal new variants for the molecular markers and confirm significant genetic differentiation between Iberian wolf and native domestic dogs from Portugal. Based on our sampling, no signs of recent introgression between the two subspecies were detected. Y chromosome data do not reveal genetic differentiation among the analyzed dog breeds, suggesting they share the same patrilineal origin. Moreover, the genetic distinctiveness of the Iberian wolf from other wolf populations is further confirmed with the description of new mtDNA variants for this endemism. Our research also discloses new molecular markers for wolf and dog subspecies assignment, which might become particularly relevant in the case of forensic or noninvasive genetic studies. The Iberian wolf represents a relic of the once widespread wolf population in Europe and our study reveals that it is a reservoir of unique genetic diversity of the grey wolf, Canis lupus. These results stress the need for conservation plans that will guarantee the sustainability of this threatened top predator in Iberia.

Cryptic diversity in the Azorean beetle genus Tarphius Erichson, 1845 (Coleoptera: Zopheridae): An integrative taxonomic approach with description of four new species.

Recent findings based on molecular data support the occurrence in the Azores of several independently evolving lineages of the beetle genus Tarphius Erichson, 1845 (Coleoptera: Zopheridae Solier, 1834) and higher species richness masked by cryptic diversity, needing formal taxonomic description. All Tarphius from the Azores are revised using an integrative taxonomic approach, using evidence from morphology, morphometrics and molecular data to delimit species. Our results reveal that Azorean Tarphius comprise at least five phyletic lineages, two of which share a similar morphology, despite being divergent at the molecular level. A total of four new species are described grouped into two complexes: i) two new species in the "complex tornvalli" with the new taxa Tarphius relictus sp. nov. (Terceira) and Tarphius furtadoi sp. nov. (São Jorge, Faial and Pico) and; ii) two new species in the "complex azoricus-wollastoni-depressus" with the new taxa Tarphius gabrielae sp. nov. (Pico) and Tarphius floresensis sp. nov. (Flores). Descriptions, photographs of holotypes and morphological details, and remarks on diagnostic features comparing similar species are presented. Additional information on the distribution and conservation status of the 12 described species in the archipelago is also provided.

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 GDM's 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.

New records and detailed distribution and abundance of selected arthropod species collected between 1999 and 2011 in Azorean native forests.

BACKGROUND: In this contribution we present detailed distribution and abundance data for arthropod species identified during the BALA - Biodiversity of Arthropods from the Laurisilva of the Azores (1999-2004) and BALA2 projects (2010-2011) from 18 native forest fragments in seven of the nine Azorean islands (all excluding Graciosa and Corvo islands, which have no native forest left). NEW INFORMATION: Of the total 286 species identified, 81% were captured between 1999 and 2000, a period during which only 39% of all the samples were collected. On average, arthropod richness for each island increased by 10% during the time frame of these projects. The classes Arachnida, Chilopoda and Diplopoda represent the most remarkable cases of new island records, with more than 30% of the records being novelties. This study stresses the need to expand the approaches applied in these projects to other habitats in the Azores, and more importantly to other less surveyed taxonomic groups (e.g. Diptera and Hymenoptera). These steps are fundamental for getting a more accurate assessment of biodiversity in the archipelago.

Cave microbial community composition in oceanic islands: disentangling the effect of different colored mats in diversity patterns of Azorean lava caves.

Processes determining diversity and composition of bacterial communities in island volcanic caves are still poorly understood. Here, we characterized colored microbial mats in 14 volcanic caves from two oceanic islands of the Azores using 16S rRNA gene sequences. Factors determining community diversity (α) and composition (ß) were explored, namely colored mats, caves and islands, as well as environmental and chemical characteristics of caves. Additive partitioning of diversity using OTU occurrence showed a greater influence of ß-diversity between islands and caves that may relate to differences in rare OTUs (singletons and doubletons) across scales. In contrast, Shannon diversity partitioning revealed the importance of the lowest hierarchical level (α diversity, colored mat), suggesting a dominance of cosmopolitan OTUs (>1%) in most samples. Cosmopolitan OTUs included members involved in nitrogen cycling, supporting the importance of this process in Azorean caves. Environmental and chemical conditions in caves did not show any significant relationship to OTU diversity and composition. The absence of clear differences between mat colors and across scales may be explained by (1) the geological youth of the cave system (cave communities have not had enough time to diverge) or/and (2) community convergence, as the result of selection pressure in extreme environments.

Comparative phylogeography of endemic Azorean arthropods.

BACKGROUND: For a remote oceanic archipelago of up to 8 Myr age, the Azores have a comparatively low level of endemism. We present an analysis of phylogeographic patterns of endemic Azorean island arthropods aimed at testing patterns of diversification in relation to the ontogeny of the archipelago, in order to distinguish between alternative models of evolutionary dynamics on islands. We collected individuals of six species (representing Araneae, Hemiptera and Coleoptera) from 16 forest fragments from 7 islands. Using three mtDNA markers, we analysed the distribution of genetic diversity within and between islands, inferred the differentiation time-frames and investigated the inter-island migration routes and colonization patterns. RESULTS: Each species exhibited very low levels of mtDNA divergence, both within and between islands. The two oldest islands were not strongly involved in the diffusion of genetic diversity within the archipelago. The most haplotype-rich islands varied according to species but the younger, central islands contributed the most to haplotype diversity. Colonization events both in concordance with and in contradiction to an inter-island progression rule were inferred, while a non-intuitive pattern of colonization from western to eastern islands was also inferred. CONCLUSIONS: The geological development of the Azores has followed a less tidy progression compared to classic hotspot archipelagos, and this is reflected in our findings. The study species appear to have been differentiating within the Azores for <2 Myr, a fraction of the apparent life span of the archipelago, which may indicate that extinction events linked to active volcanism have played an important role. Assuming that after each extinction event, colonization was initiated from a nearby island hosting derived haplotypes, the apparent age of species diversification in the archipelago would be moved closer to the present after each extinction-recolonization cycle. Exploiting these ideas, we propose a general model for future testing.

Spatial distribution of Madeira Island Laurisilva endemic spiders (Arachnida: Araneae).

Madeira island presents a unique spider diversity with a high number of endemic species, many of which are still poorly known. A recent biodiversity survey on the terrestrial arthropods of the native forest, Laurisilva, provided a large set of standardized samples from various patches throughout the island. Out of the fifty two species recorded, approximately 33.3% are Madeiran endemics, many of which had not been collected since their original description. Two new species to science are reported - Ceratinopsis n. sp. and Theridion n. sp. - and the first records of Poecilonetavariegata (Blackwall, 1841) and Tetragnathaintermedia Kulczynski, 1891 are reported for the first time for Madeira island. Considerations on species richness and abundance from different Laurisilva locations are presented, together with distribution maps for endemic species. These results contribute to a better understanding of spider diversity patterns and endemic species distribution in the native forest of Madeira island.

Comparison of Bacterial Diversity in Azorean and Hawai'ian Lava Cave Microbial Mats.

Worldwide, lava caves host colorful microbial mats. However, little is known about the diversity of these microorganisms, or what role they may play in the subsurface ecosystem. White and yellow microbial mats were collected from four lava caves each on the Azorean island of Terceira and the Big Island of Hawai'i, to compare the bacterial diversity found in lava caves from two widely separated archipelagos in two different oceans at different latitudes. Scanning electron microscopy of mat samples showed striking similarities between Terceira and Hawai'ian microbial morphologies. 16S rRNA gene clone libraries were constructed to determine the diversity within these lava caves. Fifteen bacterial phyla were found across the samples, with more Actinobacteria clones in Hawai'ian communities and greater numbers of Acidobacteria clones in Terceira communities. Bacterial diversity in the subsurface was correlated with a set of factors. Geographical location was the major contributor to differences in community composition (at the OTU level), together with differences in the amounts of organic carbon, nitrogen and copper available in the lava rock that forms the cave. These results reveal, for the first time, the similarity among the extensive bacterial diversity found in lava caves in two geographically separate locations and contribute to the current debate on the nature of microbial biogeography.