Distinct movement patterns generate stages of spider web-building

The geometric complexity and stereotypy of spider webs have long generated interest in their algorithmic origin. Like other examples of animal architecture, web construction is the result of several assembly phases that are driven by distinct behavioral stages coordinated to build a successful structure. Manual observations have revealed a range of sensory cues and movement patterns used during web construction, but methods to systematically quantify the dynamics of these sensorimotor patterns are lacking. Here, we apply an analytical pipeline to quantify web-making behavior of the orb-weaver Uloborus diversus. Position tracking revealed stereotyped stages of construction that could occur in typical or atypical progressions across individuals. Using an unsupervised clustering approach, we identified general and stage-specific leg movements. A Hierarchical Hidden Markov Model revealed that stages of web-building are characterized by stereotyped sequences of actions largely shared across individuals, regardless of whether these stages progress in a typical or atypical fashion. Web stages could be predicted based on action-sequences alone, revealing that web-stages are a physical manifestation of underlying behavioral phases.

Higher-form symmetries of 6d and 5d theories

We describe general methods for determining higher-form symmetry groups of known 5d and 6d superconformal field theories (SCFTs), and 6d little string theories (LSTs). The 6d theories can be described as supersymmetric gauge theories in 6d which include both ordinary non-abelian 1-form gauge fields and also abelian 2-form gauge fields. Similarly, the 5d theories can also be often described as supersymmetric non-abelian gauge theories in 5d. Naively, the 1-form symmetry of these 6d and 5d theories is captured by those elements of the center of ordinary gauge group which leave the matter content of the gauge theory invariant. However, an interesting subtlety is presented by the fact that some massive BPS excitations, which includes the BPS instantons, are charged under the center of the gauge group, thus resulting in a further reduction of the 1-form symmetry. We use the geometric construction of these theories in M/F-theory to determine the charges of these BPS excitations under the center. We also provide an independent algorithm for the determination of 1-form symmetry for 5d theories that admit a generalized toric construction (i.e. a 5-brane web construction). The 2-form symmetry group of 6d theories, on the other hand, is captured by those elements of the center of the abelian 2-form gauge group that leave all the massive BPS string excitations invariant, which is much more straightforward to compute as it is encoded in the Green-Schwarz coupling associated to the 6d theory.

Orb-weaving spiders show a correlated syndrome of morphology and web structure in the wild

Extended phenotypes are traits that exist outside the physical body of organisms. Despite their role in the lives of the organisms that express them and other organisms influenced by extended phenotypes, the consistency and covariance with morphological and behavioural traits of extended phenotypes has rarely been evaluated. We repeatedly measured an extended phenotype involved in prey acquisition (web structure) of wild orb-weaving spiders (Micrathena vigorsii), which re-build their webs daily. We related web structure to behaviours and spider body length. Web diameter and web density were repeatable among individuals, reaction to a predation threat was very marginally so, and response to a prey stimulus and web evenness were not repeatable. Larger spiders spun wider webs, had webs with increased thread spacing, and the spider possibly tended to react more slowly to a predation threat. When a spider built a relatively larger web it was also a relatively less dense and less even web. The repeatability of web construction and relationship with spider body size we found may be common features of intra-population variation in web structure in spiders. By estimating the consistency and covariances of extended phenotypes we can begin to evaluate what maintains their variation and how they might evolve.

Properties of orb weaving spider glycoprotein glue change during Argiope trifasciata web construction

An orb web’s prey capture thread relies on its glue droplets to retain insects until a spider can subdue them. Each droplet’s viscoelastic glycoprotein adhesive core extends to dissipate the forces of prey struggle as it transfers force to stiffer, support line flagelliform fibers. In large orb webs, switchback capture thread turns are placed at the bottom of the web before a continuous capture spiral progresses from the web’s periphery to its interior. To determine if the properties of capture thread droplets change during web spinning, we characterized droplet and glycoprotein volumes and material properties from the bottom, top, middle, and inner regions of webs. Both droplet and glycoprotein volume decreased during web construction, but there was a progressive increase in the glycoprotein’s Young’s modulus and toughness. Increases in the percentage of droplet aqueous material indicated that these increases in material properties are not due to reduced glycoprotein viscosity resulting from lower droplet hygroscopicity. Instead, they may result from changes in aqueous layer compounds that condition the glycoprotein. A 6-fold difference in glycoprotein toughness and a 70-fold difference in Young’s modulus across a web documents the phenotypic plasticity of this natural adhesive and its potential to inspire new materials.

Orb-weaving spiders show a correlated syndrome of morphology and web structure in the wild

Extended phenotypes are traits that exist outside the physical body of the organism. Despite their potential role in the lives of both the organisms that express them and other organisms that can be influenced by extended phenotypes, the consistency and covariance with morphological and behaviour traits of extended phenotypes is rarely evaluated, especially in wild organisms. We repeatedly measured an extended phenotype that directly influences an organism’s prey acquisition, the web structure, of wild orb-weaving spiders (Micrathena vigorsii), which re-build their webs each day. We related web structure traits to behavioural traits and body size (length). Both web diameter and web density were repeatably different among individuals, while reaction to a predation threat was slightly so, but response to a prey stimulus and web symmetry were not. There was a syndrome between morphology and web structure traits, where larger spiders spun webs that were wider, had webs with increased thread spacing, and the spider tended to react more slowly to a predation threat. When a spider built a relatively larger web it was also relatively a less dense and less symmetrical web. The repeatability of web construction and relationship with spider body size we found may be common features of intra-population variation in web structure in spiders. Individual variation along the morphology and web structure syndrome could represent variation in individual foraging strategies, or age-based correlated changes. By estimating the consistency and covariances of extended phenotypes we can begin to evaluate what maintains their variation and how they might evolve.

Division of labor increases with colony size, regardless of group composition, in the social spiderStegodyphus dumicola

Division of labor (DOL) is a pattern of work organization where individual group members specialize on different tasks. DOL is argued to have been instrumental for the success of eusocial insects, where it scales positively with group size both within and across species. Here we evaluate whether DOL scales positively with group size in a society of cooperative breeders (social spiders) and whether this pattern is impacted by the behavioral composition of the group. To do this we engineered experimental colonies of contrasting group sizes and behavioral compositions and tracked individuals participation in two colony maintenance tasks: prey capture and web construction. As with some eusocial insects, we found that larger groups exhibited DOL metrics up to 10-times greater than smaller groups, conveying that individuals specialize on particular tasks more in larger colonies. This scalar relationship did not differ by a groups behavioral composition, though groups composed of only bold spiders exhibited reduced DOL relative to all-shy or mixed groups. We also found that per capita participation in prey capture, but not web construction, decreased as a function of group size. This suggests that individuals in larger groups may save energy by reducing their involvement in some tasks. Together, our results convey that similar scalar relationships between DOL and group size can emerge both inside and outside the eusocial insects. Thus, theory developed for understanding DOL in eusocial societies may inform our understanding of group function in a larger swath of animal social diversity than is broadly appreciated.SignificanceStatementDivision of labor (DOL) has been a major area of research in the eusocial insects for decades, and is argues to underlie their ecological success. Only recently have other social arthropods, such as social spiders, been considered for studies concerning DOL. Given their smaller colony sizes, and absence of morphological castes, DOL was not thought to be an important facet of spider societies. However, we found that spider societies do indeed exhibit high degrees of DOL that is positively correlated to colony size, as seen in many eusocial insects. These findings suggest that the scalar relationship between group size and social organization seen in social insects is likely generalizable to a larger diversity of social taxa, and that cooperative breeders can show levels of division of labor equaling or exceeding those of eusocial systems evaluated to date.