The Non-Dereliction in Evolution: Trophic Specialisation Drives Convergence in the Radiation of Red Devil Spiders (Araneae: Dysderidae) in the Canary Islands.

Natural selection plays a key role in deterministic evolution, as clearly illustrated by the multiple cases of repeated evolution of ecomorphological characters observed in adaptive radiations. Unlike most spiders, Dysdera species display a high variability of cheliceral morphologies, which has been suggested to reflect different levels of specialization to feed on isopods. In this study, we integrate geometric morphometrics and experimental trials with a fully resolved phylogeny of the highly diverse endemic species from the Canary Islands to 1) quantitatively delimit the different cheliceral morphotypes present in the archipelago, 2) test their association with trophic specialization, as reported for continental species, 3) reconstruct the evolution of these ecomorphs throughout the diversification of the group, 4) test the hypothesis of convergent evolution of the different morphotypes, and 5) examine whether specialization constitutes a case of evolutionary irreversibility in this group. We show the existence of 9 cheliceral morphotypes and uncovered their significance for trophic ecology. Further, we demonstrate that similar ecomorphs evolved multiple times in the archipelago, providing a novel study system to explain how convergent evolution and irreversibility due to specialization may be combined to shape phenotypic diversification in adaptive radiations.

Dispersal ability and niche breadth influence interspecific variation in spider abundance and occupancy.

The relationship between species local abundance and their regional distribution (occupancy) is one of the most extensively recognized and investigated patterns in ecology. While exceptions exist, the generally held model is that locally abundant species also tend to be more widespread geographically. However, there is only a limited understanding of both the mechanisms driving this relationship, and their scale dependency. Here we use occupancy and abundance data for 123 species of spider from across the Canary Islands to understand how both dispersal ability and niche breadth might mediate variation among species for local abundance and occupancy. We test the predictions that (i) dispersal ability explains variation among species for both abundance and occupancy, and (ii) species with a higher degree of habitat specialization, reflecting more limited niche breadth, will have both higher occupancy and abundance. We find no evidence within habitat patches for an effect of dispersal ability on either local abundance or site occupancy, while across all patches species with higher dispersal ability tend to occupy more sites. Species largely restricted to laurel forests have higher abundance than species with broader niche breadth, but similar occupancy. The study revealed that dispersal ability and niche breadth were significant predictors of the abundance-occupancy relationship, highlighting the importance of both factors for understanding patterns of abundance and occupancy among spider species.

How much biodiversity is concealed in the word 'biodiversity'?

Amidst a global biodiversity crisis1, the word 'biodiversity' has become indispensable for conservation and management2. Yet, biodiversity is often used as a buzzword in scientific literature. Resonant titles of papers claiming to have studied 'global biodiversity' may be used to promote research focused on a few taxonomic groups, habitats, or facets of biodiversity - taxonomic, (phylo)genetic, or functional. This usage may lead to extrapolating results outside the target systems of these studies with direct consequences for our understanding of life on Earth and its practical conservation. Here, we used a random sample of papers with the word 'biodiversity' in their title to take a long view of the use of this term. Despite improvements in analytical tools, monitoring technologies, and data availability3,4, we found that the taxonomic scope of research articles has not increased in recent years. We also show that studies with a wider taxonomic scope attract more citations and online attention. Our results have broad ramifications for understanding how extrapolating from studies with narrow taxonomic scope affects our view of global biodiversity and conservation.

Global response of conservationists across mass media likely constrained bat persecution due to COVID-19.

Most people lack direct experience with wildlife and form their risk perception primarily on information provided by the media. The way the media frames news may substantially shape public risk perception, promoting or discouraging public tolerance towards wildlife. At the onset of the COVID-19 pandemic, bats were suggested as the most plausible reservoir of the virus, and this became a recurrent topic in media reports, potentially strengthening a negative view of this ecologically important group. We investigated how media framed bats and bat-associated diseases before and during the COVID-19 pandemic by assessing the content of 2651 online reports published across 26 countries, to understand how and how quickly worldwide media may have affected the perception of bats. We show that the overabundance of poorly contextualized reports on bat-associated diseases likely increased the persecution towards bats immediately after the COVID-19 outbreak. However, the subsequent interventions of different conservation communication initiatives allowed pro-conservation messages to resonate across the global media, likely stemming an increase in bat persecution. Our results highlight the modus operandi of the global media regarding topical biodiversity issues, which has broad implications for species conservation. Knowing how the media acts is pivotal for anticipating the propagation of (mis)information and negative feelings towards wildlife. Working together with journalists by engaging in dialogue and exchanging experiences should be central in future conservation management.

The World Spider Trait database: a centralized global open repository for curated data on spider traits.

Spiders are a highly diversified group of arthropods and play an important role in terrestrial ecosystems as ubiquitous predators, which makes them a suitable group to test a variety of eco-evolutionary hypotheses. For this purpose, knowledge of a diverse range of species traits is required. Until now, data on spider traits have been scattered across thousands of publications produced for over two centuries and written in diverse languages. To facilitate access to such data, we developed an online database for archiving and accessing spider traits at a global scale. The database has been designed to accommodate a great variety of traits (e.g. ecological, behavioural and morphological) measured at individual, species or higher taxonomic levels. Records are accompanied by extensive metadata (e.g. location and method). The database is curated by an expert team, regularly updated and open to any user. A future goal of the growing database is to include all published and unpublished data on spider traits provided by experts worldwide and to facilitate broad cross-taxon assays in functional ecology and comparative biology. Database URL:https://spidertraits.sci.muni.cz/.

Prey acceptance and metabolic specialisations in some Canarian Dysdera spiders.

Spiders of the genus Dysdera are peculiar for their preying on terrestrial isopods though the preference for this prey type varies between species. We tested prey acceptance of two isopod and two non-isopod prey in 8 species endemic to the Canary Islands, and analyzed growth and metabolic parameters (growth efficiency; dry mass, lipid and N extraction efficiency; lipid:protein consumption ratio) of 6 of these species when fed either house flies (Musca domestica) or isopods (Porcellio scaber) in the laboratory. The species represented four morphological types (unmodified chelicerae, slightly elongated chelicerae, concave chelicerae, flattened fang), supposedly reflecting different specializations to isopod prey. The results showed reduced relative acceptance of non-isopod and increased acceptance of isopod prey in species groups with specialised morphologies compared to the unmodified species group. All species had similar or lower growth and growth efficiency when feeding on isopods than on flies. Extraction efficiency of dry mass and lipid were higher for flies than for isopods, while extraction efficiency of protein was higher for isopods than for flies. All species also utilized isopod protein equally well, but protein utilization of flies was lower in the presumed specialist compared to generalist species, indicating a possible metabolic trade-off from isopod specialisation. Thus, morphological adaptations were associated with increased behavioural preferences for isopods and reduced metabolic ability to handle 'generalist' prey.

Standardised spider (Arachnida, Araneae) inventory of Kilpisjärvi, Finland.

BACKGROUND: A spider taxonomy and ecology field course was organised in Kilpisjärvi Biological Station, northern Finland, in July 2019. During the course, four 50 × 50 m plots in mountain birch forest habitat were sampled following a standardised protocol. In addition to teaching and learning about spider identification, behaviour, ecology and sampling, the main aim of the course was to collect comparable data from the Kilpisjärvi area as part of a global project, with the purpose of uncovering global spider diversity patterns. NEW INFORMATION: A total of 2613 spiders were collected, of which 892 (34%) were adults. Due to uncertainty of juvenile identification, only adults are included in the data presented in this paper. The observed adult spiders belong to 51 species, 40 genera and 11 families, of which the Linyphiidae were the most rich and abundant with 28 (55%) species and 461 (52%) individuals. Lycosidae had six species and 286 individuals, Gnaphosidae five species and 19 individuals, Thomisidae four species and 24 individuals, Theridiidae two species and 23 individuals. All other six families had one species and less than 40 individuals. The most abundant species were the linyphiid Agnyphantes expunctus (204) and the lycosids Pardosa eiseni (164) and Pardosa hyperborea (107).

A database of functional traits for spiders from native forests of the Iberian Peninsula and Macaronesia.

BACKGROUND: There is an increasing demand for databases including species trait information for biodiversity and community ecology studies. The existence of trait databases is useful for comparative studies within taxa or geographical regions, but there is low availability of databases for certain organisms. Here we present an open access functional trait database for spiders from Macaronesia and the Iberian Peninsula, recording several morphological and ecological traits related to the species life histories, microhabitat and trophic preferences. NEW INFORMATION: We present a database that includes 12 biological traits for 506 spider species present in natural forests of the Iberian Peninsula (Spain) and three Macaronesian archipelagoes (Azores, Madeira and Canary Islands). The functional trait database consists of two sections:individual-level data for six morphological traits (total body size, prosoma length, prosoma width, prosoma height, tibia I length and fang length), based on direct measurements of 2844 specimens of all spider species; andspecies-level aggregate data for 12 traits (same 6 morphological traits as in the previous section plus dispersal ability, vertical stratification, circadian activity, foraging strategy, trophic specialization and colonization status), based on either the average of the direct measurements or bibliographic searches.This functional trait database will serve as a data standard for currently ongoing analyses that require trait and functional diversity statistics.

Standardised inventories of spiders (Arachnida, Araneae) of Macaronesia II: The native forests and dry habitats of Madeira archipelago (Madeira and Porto Santo islands).

BACKGROUND: Here we present the data obtained from the samples collected as part of a large research project (MACDIV) which aims at understanding the drivers of spider (Araneae) community assembly in Macaronesian islands. To obtain the data, we applied the sampling protocol COBRA (Conservation Oriented Biodiversity Rapid Assessment), in twelve 50 m x 50 m native forest plots and five dry habitat plots on the island of Madeiraand in 5 dry habitat plots on the island of Porto Santo. Through this publication, we contribute to the knowledge of the arachnofauna of the Madeiran archipelago. NEW INFORMATION: From the samples that we collected, we obtained a total of 14,902 specimens, of which 49% were adults (7,263). We identified these specimens to 87 species and 18 morphospecies (undescribed), belonging to 26 families. Species of the family Linyphiidae dominated the samples, with 24 (morpho)species. Out of the 105 recorded (morpho)species, 34 were endemic, 26 native non-endemic, 22 introduced and 23 species of unknown origin. We report seven new records of possibly recently introduced species in the Madeiran archipelago. We also present 21 new records for Madeira island and 32 for Porto Santo (33 for the whole archipelago).

Chance and predictability in evolution: The genomic basis of convergent dietary specializations in an adaptive radiation.

The coexistence of multiple eco-phenotypes in independently assembled communities makes island adaptive radiations the ideal framework to test convergence and parallelism in evolution. In the radiation of the spider genus Dysdera in the Canary Islands, species diversification occurs concomitant with repeated events of trophic specialization. These dietary shifts, to feed primarily on woodlice, are accompanied by modifications in morphology (mostly in the mouthparts), behaviour and nutritional physiology. To gain insight into the molecular basis of this adaptive radiation, we performed a comprehensive comparative transcriptome analysis of five Canary Island Dysdera endemics representing two evolutionary and geographically independent events of dietary specialization. After controlling for the potential confounding effects of hemiplasy, our differential gene expression and selective constraint analyses identified a number of genetic changes that could be associated with the repeated adaptations to specialized diet of woodlice, including some related to heavy metal detoxification and homeostasis, the metabolism of some important nutrients and venom toxins. Our results shed light on the genomic basis of an extraordinary case of dietary shift convergence associated with species diversification. We uncovered putative molecular substrates of convergent evolutionary changes at different hierarchical levels, including specific genes, genes with equivalent functions and even particular amino acid positions. This study improves our knowledge of rapid adaptive radiations and provides new insights into the predictability of evolution.

A DNA barcode-assisted annotated checklist of the spider (Arachnida, Araneae) communities associated to white oak woodlands in Spanish National Parks.

BACKGROUND: A large scale semi-quantitative biodiversity assessment was conducted in white oak woodlands in areas included in the Spanish Network of National Parks, as part of a project aimed at revealing biogeographic patterns and identify biodiversity drivers. The semi-quantitative COBRA sampling protocol was conducted in sixteen 1-ha plots across six national parks using a nested design. All adult specimens were identified to species level based on morphology. Uncertain delimitations and identifications due to either limited information of diagnostic characters or conflicting taxonomy were further investigated using DNA barcode information. NEW INFORMATION: We identified 376 species belonging to 190 genera in 39 families, from the 8,521 adults found amongst the 20,539 collected specimens. Faunistic results include the discovery of 7 new species to the Iberian Peninsula, 3 new species to Spain and 11 putative new species to science. As largely expected by environmental features, the southern parks showed a higher proportion of Iberian and Mediterranean species than the northern parks, where the Palearctic elements were largely dominant. The analysis of approximately 3,200 DNA barcodes generated in the present study, corroborated and provided finer resolution to the morphologically based delimitation and identification of specimens in some taxonomically challenging families. Specifically, molecular data confirmed putative new species with diagnosable morphology, identified overlooked lineages that may constitute new species, confirmed assignment of specimens of unknown sexes to species and identified cases of misidentifications and phenotypic polymorphisms.

Molecular gut content analysis of different spider body parts.

Molecular gut-content analysis has revolutionized the study of food webs and feeding interactions, allowing the detection of prey DNA within the gut of many organisms. However, successful prey detection is a challenging procedure in which many factors affect every step, starting from the DNA extraction process. Spiders are liquid feeders with branched gut diverticula extending into their legs and throughout the prosoma, thus digestion takes places in different parts of the body and simple gut dissection is not possible. In this study, we investigated differences in prey detectability in DNA extracts from different parts of the spider´s body: legs, prosoma and opisthosoma, using prey-specific PCR and metabarcoding approaches. We performed feeding trials with the woodlouse hunter spider Dysdera verneaui Simon, 1883 (Dysderidae) to estimate the time at which prey DNA is detectable within the predator after feeding. Although we found that all parts of the spider body are suitable for gut-content analysis when using prey-specific PCR approach, results based on metabarcoding suggested the opisthosoma is optimal for detection of predation in spiders because it contained the highest concentration of prey DNA for longer post feeding periods. Other spiders may show different results compared to D. verneaui, but given similarities in the physiology and digestion in different families, it is reasonable to assume this to be common across species and this approach having broad utility across spiders.

Evolution of Chemosensory Gene Families in Arthropods: Insight from the First Inclusive Comparative Transcriptome Analysis across Spider Appendages.

Unlike hexapods and vertebrates, in chelicerates, knowledge of the specific molecules involved in chemoreception comes exclusively from the comparative analysis of genome sequences. Indeed, the genomes of mites, ticks and spiders contain several genes encoding homologs of some insect membrane receptors and small soluble chemosensory proteins. Here, we conducted for the first time a comprehensive comparative RNA-Seq analysis across different body structures of a chelicerate: the nocturnal wandering hunter spider Dysdera silvatica Schmidt 1981. Specifically, we obtained the complete transcriptome of this species as well as the specific expression profile in the first pair of legs and the palps, which are thought to be the specific olfactory appendages in spiders, and in the remaining legs, which also have hairs that have been morphologically identified as chemosensory. We identified several ionotropic (Ir) and gustatory (Gr) receptor family members exclusively or differentially expressed across transcriptomes, some exhibiting a distinctive pattern in the putative olfactory appendages. Furthermore, these IRs were the only known olfactory receptors identified in such structures. These results, integrated with an extensive phylogenetic analysis across arthropods, uncover a specialization of the chemosensory gene repertoire across the body of D. silvatica and suggest that some IRs likely mediate olfactory signaling in chelicerates. Noticeably, we detected the expression of a gene family distantly related to insect odorant-binding proteins (OBPs), suggesting that this gene family is more ancient than previously believed, as well as the expression of an uncharacterized gene family encoding small globular secreted proteins, which appears to be a good chemosensory gene family candidate.

A geographical distribution database of the genus Dysdera in the Canary Islands (Araneae, Dysderidae).

The ground-dweller spider genus Dysdera shows very high species richness on the oceanic archipelago of the Canary Islands, providing one of the most outstanding examples of island radiation among spiders, only paralleled by Tetragnatha spiders on the Hawaiian archipelago. A georeferenced database of the 48 Dysdera species occurring in the Canary Islands was assembled to facilitate ongoing and future research on this remarkable lineage. All species are endemic to the archipelago except for the cosmopolitan Dysdera crocata. The dataset consists of 794 distributional records documented from 1971 to 2015, each locality being represented only once per species. Distribution maps are provided for each species, along with basic diversity and distribution information. The database and geographical maps included in this article stand for the most updated, accurate and complete information on the distribution of the spider genus Dysdera in the Canary Islands.

The imprint of geologic history on within-island diversification of woodlouse-hunter spiders (Araneae, Dysderidae) in the Canary Islands.

Geological processes and ecological adaptation are major drivers of diversification on oceanic islands. Although diversification in these islands is often interpreted as resulting from dispersal or island hopping rather than vicariance, this may not be the case in islands with complex geological histories. The island of Tenerife, in the Canary Islands, emerged in the late Miocene as 3 precursor islands that were subsequently connected and reisolated by volcanic cycles. The spider Dysdera verneaui is endemic to the island of Tenerife, where it is widely distributed throughout most island habitats, providing an excellent model to investigate the role of physical barriers and ecological adaptation in shaping within-island diversity. Here, we present evidence that the phylogeographic patterns of this species trace back to the independent emergence of the protoislands. Molecular markers (mitochondrial genes cox1, 16S, and nad1 and the nuclear genes ITS-2 and 28S) analyzed from 100 specimens (including a thorough sampling of D. verneaui populations and additional outgroups) identify 2 distinct evolutionary lineages that correspond to 2 precursor islands, each with diagnostic genital characters indicative of separate species status. Episodic introgression events between these 2 main evolutionary lineages explain the observed incongruence between mitochondrial and nuclear markers, probably as a result of the homogenization of their ITS-2 sequence types. The most widespread lineage exhibits a complex population structure, which is compatible with either secondary contact, following connection of deeply divergent lineages, or alternatively, a back colonization from 1 precursor island to another.

Living on the edge: demographic and phylogeographical patterns in the woodlouse-hunter spider Dysdera lancerotensis Simon, 1907 on the eastern volcanic ridge of the Canary Islands.

The Eastern Canary Islands are the emerged tips of a continuous volcanic ridge running parallel to the northeastern African coast, originated by episodic volcanic eruptions that can be traced back to the Miocene and that, following a major period of quiescence and erosion, continued from the Pliocene to the present day. The islands have been periodically connected by eustatic sea-level changes resulting from Pleistocene glacial cycles. The ground-dwelling spider Dysdera lancerotensis Simon, 1907 occurs along the entire ridge, except on recent barren lavas and sand dunes, and is therefore an ideal model for studying the effect of episodic geological processes on terrestrial organisms. Nested clade and population genetic analyses using 39 haplotypes from 605 base pairs of mitochondrial DNA cytochrome c oxidase I sequence data, along with phylogenetic analyses including two additional mitochondrial genes, uncover complex phylogeographical and demographic patterns. Our results indicate that D. lancerotensis colonized the ridge from north to south, in contrast to what had been expected given the SSW-NNE trend of volcanism and to what had been reported for other terrestrial arthropods. The occurrence of several episodes of extinction, recolonization and expansion are hypothesized for this species, and areas that act as refugia during volcanic cycles are identified. Relaxed molecular clock methods reveal divergence times between main haplotype lineages that suggest an older origin of the northern islets than anticipated based on geological evidence. This study supports the key role of volcanism in shaping the distribution of terrestrial organisms on oceanic islands and generates phylogeographical predictions that warrant further research into other terrestrial endemisms of this fascinating region.