Global Diversification of Anelosimus Spiders Driven by Long-Distance Overwater Dispersal and Neogene Climate Oscillations.

Vicariance and dispersal events, combined with intricate global climatic history, have left an imprint on the spatiotemporal distribution and diversity of many organisms. Anelosimus cobweb spiders (Theridiidae), are organisms ranging in behavior from solitary to highly social, with a cosmopolitan distribution in temperate to tropical areas. Their evolutionary history and the discontinuous distribution of species richness suggest that 1) long-distance overwater dispersal and 2) climate change during the Neogene (23-2.6 Ma), may be major factors in explaining their distribution and diversification. Here, we test these hypotheses, and explicitly test if global Miocene/Pliocene climatic cooling in the last 8 Ma affected Anelosimus radiation in parallel in South America and Madagascar. To do so, we investigate the phylogeny and spatiotemporal biogeography of Anelosimus through a culmination of a 20-year comprehensive global sampling at the species level (69 species, including 84% of the known 75 species worldwide, represented by 268 individuals) using nucleotide data from seven loci (5.5 kb). Our results strongly support the monophyly of Anelosimus with an Oligocene ($\sim $30 Ma) South American origin. Major clades on other continents originate via multiple, long-distance dispersal events, of solitary or subsocial-but not social-lineages, from the Americas. These intercontinental dispersals were to Africa, Madagascar (twice), and SE Asia/Australasia. The early diversification of Anelosimus spiders coincides with a sudden thermal increase in the late Oligocene ($\sim $27-25 Ma), though no causal connection can be made. Our results, however, strongly support the hypothesis that global Neogene climatic cooling in the last 8 Ma drove Anelosimus radiation in parallel in South America and Madagascar, offering a rare empirical evidence for diversification of a socially diverse group driven by an interplay between long-distance dispersal and global Neogene climatic changes. [Cobweb spiders; diversification; global biogeography; long-distance dispersal; molecular phylogenetics; neogene climate changes; sociality; vicariance.].

Molecular phylogeny of moth-specialized spider sub-family Cyrtarachninae, which includes bolas spiders.

The evolutionary process of the unique web architectures of spiders of the sub-family Cyrtarachninae, which includes the triangular web weaver, bolas spider, and webless spider, is thought to be derived from reduction of orbicular 'spanning-thread webs' resembling ordinal orb webs. A molecular phylogenetic analysis was conducted to explore this hypothesis using orbicular web spiders Cyrtarachne, Paraplectana, Poecilopachys, triangular web spider Pasilobus, bolas spiders Ordgarius and Mastophora, and webless spider Celaenia. The phylogeny inferred from partial sequences of mt-COI, nuclear 18S-rRNA and 28S-rRNA showed that the common ancestor of these spiders diverged into two clades: a spanning-thread web clade and a bolas or webless clade. This finding suggests that the triangular web evolved by reduction of an orbicular spanning web, but that bolas spiders evolved in the early stage, which does not support the gradual web reduction hypothesis.

1000 spider silkomes: Linking sequences to silk physical properties.

Spider silks are among the toughest known materials and thus provide models for renewable, biodegradable, and sustainable biopolymers. However, the entirety of their diversity still remains elusive, and silks that exceed the performance limits of industrial fibers are constantly being found. We obtained transcriptome assemblies from 1098 species of spiders to comprehensively catalog silk gene sequences and measured the mechanical, thermal, structural, and hydration properties of the dragline silks of 446 species. The combination of these silk protein genotype-phenotype data revealed essential contributions of multicomponent structures with major ampullate spidroin 1 to 3 paralogs in high-performance dragline silks and numerous amino acid motifs contributing to each of the measured properties. We hope that our global sampling, comprehensive testing, integrated analysis, and open data will provide a solid starting point for future biomaterial designs.

The moth specialist spider Cyrtarachne akirai uses prey scales to increase adhesion.

Contaminants decrease adhesive strength by interfering with substrate contact. Spider webs adhering to moths present an ideal model to investigate how natural adhesives overcome contamination because moths' sacrificial layer of scales rubs off on sticky silk, facilitating escape. However, Cyrtarachninae spiders have evolved gluey capture threads that adhere well to moths. Cyrtarachne capture threads contain large glue droplets oversaturated with water, readily flowing but also prone to drying out. Here, we compare the spreading and adhesion of Cyrtarachne akirai glue on intact mothwings, denuded cuticle and glass to the glue of a common orb-weaving spider, Larinioides cornutus, to understand how C. akirai glue overcomes dirty surfaces. Videos show that C. akirai's glue spreading accelerates along the underlying moth cuticle after the glue seeps beneath the moth scales-not seen on denuded cuticle or hydrophilic glass. Larinioides cornutus glue droplets failed to penetrate the moth scales, their force of adhesion thus limited by the strength of attachment of scales to the cuticle. The large size and low viscosity of C. akirai glue droplets function together to use the three-dimensional topography of the moth's scales against itself via capillary forces. Infrared spectroscopy shows C. akirai glue droplets readily lose free-flowing water. We hypothesize that this loss of water leads to increased viscosity during spreading, increasing cohesive forces during pull-off. This glue's two-phase behaviour shows how natural selection can leverage a defensive specialization of prey against themselves and highlights a new design principle for synthetic adhesives for adhering to troublesome surfaces.

Silk structure rather than tensile mechanics explains web performance in the moth-specialized spider, Cyrtarachne.

Orb webs intercept and retain prey so spiders may subdue them. Orb webs are composed of sticky, compliant spirals of capture silk spun across strong, stiff major ampullate silk threads. Interplay between differences in the mechanical properties of these silks is crucial for prey capture. Most orb webs depend upon insects contacting several radial and capture threads for successful retention. Moths, however, escape quickly from most orb webs due to the sacrificial scales covering their bodies. Cyrtarachne orb webs are unusual as they contain a reduced number of capture threads and moths stick unusually well to single threads. We aimed to determine how the tensile properties of the capture spiral and radial threads spun by Cyrtarachne operate in retention of moth prey. A NanoBionix UTM was used to quantify the material properties of flagelliform and major ampullate threads to test if Cyrtarachne's reduced web architecture is accompanied by improvements in tensile performance of its silk. Silk threads showed tensile properties typical of less-specialized orb-weavers, with the exception of high extensibility in radial threads. Radial thread diameters were 62.5% smaller than flagelliform threads, where commonly the two are roughly similar. We utilized our tensile data to create a finite element model of Cyrtarachne's web to investigate energy dissipation during prey impact. Large cross-sectional area of the flagelliform threads played a key role in enabling single capture threads to withstand prey impact. Rather than extraordinary silk, Cyrtarachne utilizes structural changes in the size and attachment of silk threads to facilitate web function.

Supersaturation with water explains the unusual adhesion of aggregate glue in the webs of the moth-specialist spider, Cyrtarachne akirai.

Orb webs produced by araneoid spiders depend upon aggregate glue-coated capture threads to retain their prey. Moths are challenging prey for most spiders because their scales detach and contaminate the glue droplets, significantly decreasing adhesion. Cyrtarachne are moth-specialist orb-weaving spiders whose capture threads adhere well to moths. We compare the adhesive properties and chemistry of Cyrtarachne aggregate glue to other orb-weaving spiders to test hypotheses about their structure, chemistry and performance that could explain the strength of Cyrtarachne glue. We show that the unusually large glue droplets on Cyrtarachne capture threads make them approximately 8 times more adhesive on glass substrate than capture threads from typical orb-weaving species, but Cyrtarachne adhesion is similar to that of other species after normalization by glue volume. Glue viscosity reversibly changes over 1000-fold in response to atmospheric humidity, and the adhesive strength of many species of orb spiders is maximized at a viscosity of approximately 105-106 cst where the contributions of spreading and bulk cohesion are optimized. By contrast, viscosity of Cyrtarachne aggregate glue droplets is approximately 1000 times lower at maximum adhesive humidity, likely facilitating rapid spreading across moth scales. Water uptake by glue droplets is controlled, in part, by hygroscopic low molecular weight compounds. NMR showed evidence that Cyrtarachne glue contains a variety of unknown low molecular weight compounds. These compounds may help explain how Cyrtarachne produces such exceptionally large and low viscosity glue droplets, and also why these glue droplets rapidly lose water volume after brief ageing or exposure to even slightly dry (e.g. < 80% RH) conditions, permanently reducing their adhesion. We hypothesize that the combination of large glue droplet size and low viscosity helps Cyrtarachne glue to penetrate the gaps between moth scales.