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 A future goal of the growing database is to include all published and unpublished data on spider traits provided by experts worldwide and to facilitate broad cross-taxon assays in functional ecology and comparative biology.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/.

Estimation of trophic niches in myrmecophagous spider predators.

Among spiders, taxonomically the most diversified group of terrestrial predators, only a few species are stenophagous and feed on ants. The levels of stenophagy and ant-specialisation vary among such species. To investigate whether stenophagy is only a result of a local specialisation both fundamental and realised trophic niches need to be estimated. Here we investigated trophic niches in three closely-related spider species from the family Gnaphosidae (Callilepis nocturna, C. schuszteri, Nomisia exornata) with different levels of myrmecophagy. Acceptance experiments were used to estimate fundamental trophic niches and molecular methods to estimate realised trophic niches. For the latter two PCR primer sets were used as these can affect the niche breadth estimates. The general invertebrate ZBJ primers were not appropriate for detecting ant DNA as they revealed very few prey types, therefore ant-specific primers were used. The cut-off threshold for erroneous MOTUs was identified as 0.005% of the total number of valid sequences, at individual predator level it was 0.05%. The fundamental trophic niche of Callilepis species included mainly ants, while that of N. exornata included many different prey types. The realised trophic niche in Callilepis species was similar to its fundamental niche but in N. exornata the fundamental niche was wider than realised niche. The results show that Callilepis species are ant-eating (specialised) stenophagous predators, catching mainly Formicinae ants, while N. exornata is an ant-eating euryphagous predator catching mainly Myrmicinae ants.

Venom of prey-specialized spiders is more toxic to their preferred prey: A result of prey-specific toxins.

In specialized predators, a variety of adaptations have evolved to such a level of specificity that they allow very effective exploitation of focal prey. Venom is an essential adaptive trait of predatory venomous species, such as spiders, yet our knowledge of spider venom is incomplete. In agreement with the prey preference hypothesis, we expected that the venom of spider specialists should be more toxic to focal than to alternative prey, because it is composed of prey-specific toxins. Here we used spiders with three types of trophic specializations: specialists that were ant-eating, termite-eating and spider-eating. We compared the efficacy of prey capture of preferred and alternative prey (measured as paralysis latency) with that of related generalists and profiled the venom of the studied species using proteomic methods. We used 22 spider species: six myrmecophagous, two termitophagous, three araneophagous and 11 euryphagous generalist species belonging to different families. We found that ten of the eleven specialist species induced significantly shorter paralysis latency in preferred prey than in alternative prey. Generalists exhibited either similar efficiency on both prey types or slightly higher efficiency on preferred prey. Multivariate analysis of proteomic profiles (peptides and proteins) revealed significant differences between trophic specializations, particularly in peptides. Specialists appear to have venom composed of unique specific compounds as revealed by the multivariate ordination and indicator analysis. These components are likely prey-specific toxins.

Metabolic specialisation on preferred prey and constraints in the utilisation of alternative prey in an ant-eating spider.

Trophic specialists are expected to possess adaptations that increase the efficiency of handling preferred prey. Such adaptations may constrain the ability to utilise alternative prey. Here we tested whether the ant-eating spider Euryopis episinoides possesses metabolic specialisations with increased efficiency in utilising preferred prey and decreased efficiency in utilising alternative prey. In addition, we investigated the contribution of genetic variation via maternal effects. We reared E. episinoides spiders from the first instar on two different diets, either ants (preferred prey) or fruit flies (alternative prey). Spider survival rate and increases in body mass were significantly higher on the ant diet. The total development time did not differ between diet groups, nor did the number of egg sacs per female or the incubation period. However, the number of eggs per egg sac and hatching success were higher on the ant diet. There was a genetic variation in several offspring traits. Our data support the hypothesis that stenophagous ant-eating E. episinoides have a metabolic specialisation on ant utilisation indicated by higher efficiency in utilising ants than fruit flies. While most individuals of E. episinoides were able to capture fruit flies, only very few spiders were able to develop and reproduce on a pure fruit fly diet, suggesting the existence of within-species genetic variation regarding the tolerance to alternative prey.

Discovery of a monophagous true predator, a specialist termite-eating spider (Araneae: Ammoxenidae).

True predators are characterised by capturing a number of prey items during their lifetime and by being generalists. Some true predators are facultative specialists, but very few species are stenophagous specialists that catch only a few closely related prey types. A monophagous true predator that would exploit a single prey species has not been discovered yet. Representatives of the spider family Ammoxenidae have been reported to have evolved to only catch termites. Here we tested the hypothesis that Ammoxenus amphalodes is a monophagous termite-eater capturing only Hodotermes mossambicus. We studied the trophic niche of A. amphalodes by means of molecular analysis of the gut contents using Next Generation Sequencing. We investigated their willingness to accept alternative prey and observed their specific predatory behaviour and prey capture efficiency. We found all of the 1.4 million sequences were H. mossambicus. In the laboratory A. amphalodes did not accept any other prey, including other termite species. The spiders attacked the lateral side of the thorax of termites and immobilised them within 1 min. The paralysis efficiency was independent of predator:prey size ratio. The results strongly indicate that A. amphalodes is a monophagous prey specialist, specifically adapted to feed on H. mossambicus.

Molecular characterization of 'Candidatus Rickettsia vini' in Ixodes arboricola from the Czech Republic and Slovakia.

The aim of this study was to analyze the prevalence of rickettsiae in the tree-hole tick Ixodes arboricola in the Czech Republic and Slovakia. During May to September of 2009 and 2013, bird boxes belonging to three different areas were screened for ticks. In total, 454 nestlings and 109 nests of 10 hole-breeding bird species were examined. Ticks were found on Ficedula albicollis, Parus major, Cyanistes caeruleus and Sitta europaea and/or in their nests. In total, 166 ticks (17 nymphs, 10 males and 139 females) were found at 3 areas (arithmetic mean±standard error: 55.3±45.9). All ticks were tested for the presence of Rickettsia species by polymerase chain reaction targeting the rickettsial genes gltA, ompA, ompB and htrA and amplicon sequencing. All individuals except 3 nymphs were infected with 'Candidatus Rickettsia vini'. Multilocus sequence typing showed closest proximity to Rickettsia japonica and Rickettsia heilongjiangensis cluster. The presence of 'Ca. R. vini' is reported for the first time in Slovakia.

David and Goliath: potent venom of an ant-eating spider (Araneae) enables capture of a giant prey.

It is rare to find a true predator that repeatedly and routinely kills prey larger than itself. A solitary specialised ant-eating spider of the genus Zodarion can capture a relatively giant prey. We studied the trophic niche of this spider species and investigated its adaptations (behavioural and venomic) that are used to capture ants. We found that the spider captures mainly polymorphic Messor arenarius ants. Adult female spiders captured large morphs while tiny juveniles captured smaller morphs, yet in both cases ants were giant in comparison with spider size. All specimens used an effective prey capture strategy that protected them from ant retaliation. Juvenile and adult spiders were able to paralyse their prey using a single bite. The venom glands of adults were more than 50 times larger than those of juvenile spiders, but the paralysis latency of juveniles was 1.5 times longer. This suggests that this spider species possesses very potent venom already at the juvenile stage. Comparison of the venom composition between juvenile and adult spiders did not reveal significant differences. We discovered here that specialised capture combined with very effective venom enables the capture of giant prey.

Phenotypic integration in a series of trophic traits: tracing the evolution of myrmecophagy in spiders (Araneae).

Several hypotheses have been put forward to explain the evolution of prey specificity (stenophagy). Yet little light has so far been shed on the process of evolution of stenophagy in carnivorous predators. We performed a detailed analysis of a variety of trophic adaptations in one species. Our aim was to determine whether a specific form of stenophagy, myrmecophagy, has evolved from euryphagy via parallel changes in several traits from pre-existing characters. For that purpose, we studied the trophic niche and morphological, behavioural, venomic and physiological adaptations in a euryphagous spider, Selamia reticulata. It is a species that is branching off earlier in phylogeny than stenophagous ant-eating spiders of the genus Zodarion (both Zodariidae). The natural diet was wide and included ants. Laboratory feeding trials revealed versatile prey capture strategies that are effective on ants and other prey types. The performance of spiders on two different diets - ants only and mixed insects - failed to reveal differences in most fitness components (survival and developmental rate). However, the weight increase was significantly higher in spiders on the mixed diet. As a result, females on a mixed diet had higher fecundity and oviposited earlier. No differences were found in incubation period, hatching success or spiderling size. S. reticulata possesses a more diverse venom composition than Zodarion. Its venom is more effective for the immobilisation of beetle larvae than of ants. Comparative analysis of morphological traits related to myrmecophagy in the family Zodariidae revealed that their apomorphic states appeared gradually along the phylogeny to derived prey-specialised genera. Our results suggest that myrmecophagy has evolved gradually from the ancestral euryphagous strategy by integrating a series of trophic traits.

Local trophic specialisation in a cosmopolitan spider (Araneae).

Trophic specialisation can be observed in species with long-term constant exploitation of a certain prey in all populations or in a population of a species with short-term exploitation of a certain prey. While in the former case the species would evolve stereotyped or specialised trophic adaptations, the trophic traits of the latter should be versatile or generalised. Here, we studied the predatory behavioural adaptations of a presumed myrmecophagous spider, Oecobius navus. We chose two distinct populations, one in Portugal and the other in Uruguay. We analysed the actual prey of both populations and found that the Portuguese population feeds mainly on dipterans, while the Uruguayan population feeds mainly on ants. Indeed, dipterans and springtails in Portugal, and ants in Uruguay were the most abundant potential prey. In laboratory trials O. navus spiders recognised and captured a wide variety of prey. The capture efficiency of the Portuguese population measured as components of the handling time was higher for flies than for ants, while that of the Uruguayan population was higher for ants. We found phenotypic plasticity in behavioural traits that lead to increased capture efficiency with respect to the locally abundant prey, but it remains to be determined whether the traits of the two populations are genetically fixed. We conclude that O. navus is a euryphagous generalist predator which shows local specialisation on the locally abundant prey.