Asymmetric female genitalia and other remarkable morphology in a new genus of cobweb spiders (Theridiidae, Araneae) from Madagascar

Symmetry is such a conspicuous feature of life that asymmetries draw our immediate attention. While not uncommon in bilateral organisms in general, asymmetry in spiders is rare. Here I report the first case of antisymmetry in external female genitalia in spiders, in the new genus Asygyna (Theridiidae: Araneae) from Madagascar. In the nearly 39 000 species of spiders described to date, the external structure of the female genitalia is symmetric. In entelegyne spiders paired external copulatory openings each lead to an internal copulatory duct, whose roughly symmetrical trajectories terminate in paired receptacles, the spermathecae. In Asygyna , here exemplified by two new species, A. huberi and A. coddingtoni , laterality is evident in the internal and external female genitalia. A single copulatory opening leads (either to the left or right depending on the individual) to a single copulatory duct with a distinctly asymmetric trajectory. The duct splits terminally shortly before entering the two spermathecae. The males are symmetric, but possibly only one palp can be used in copulation with each female. If adaptive, the selective forces behind this asymmetry are perplexing, as male access to females seems reduced. However, if males are plentiful, asymmetry may benefit the female by reducing insertion times and thus shortening copulation time, and by tightening her control over which males sire her offspring. Asygyna has a range of other bizarre sex-related morphologies, including prosomal pits and a well developed stridulatory mechanism in both sexes, a male proboscis, and simplified palps. A phylogenetic analysis, including 63 taxa and 242 morphological characters, places Asygyna in Pholcommatinae, sister to the enigmatic genus Carniella .  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 211–232.     

Complete separation along matrilines in a social spider metapopulation inferred from hypervariable mitochondrial DNA region

The distribution and quantity of genetic diversity may be profoundly influenced by the emergence and dynamics of social groups. Permanent social living in spiders has resulted in the subdivision of their populations in more or less isolated colony lineages that grow, proliferate and become extinct without mixing with one another. A newly discovered hypervariable mitochondrial DNA region allowed us to examine the fine scale metapopulation structure in the social Anelosimus eximius. We sampled 39 colonies in Ecuador and French Guiana and identified 25 haplotypes. The majority of colonies contained one haplotype. Additional haplotypes occurred in approximately 15% of the colonies, and were always closely related to the common colony haplotype. Our findings confirm that colonies consist of single matrilines, with within‐colony variation explained by mutations within the matriline. We thus found no evidence of mixing of matrilines. Likewise, colonies in a cluster often shared a haplotype, implying common colony ancestry. In few cases, however, haplotypes were shared between more distant colonies, providing evidence for occasional longer distance dispersal and/or widespread colony lineages. The geographical localities of colonies were incongruent with phylogenetic trees and haplotype networks, showing that some areas contained two or more matrilines. Hence, females do not migrate into foreign colonies, but faithfully remain within their own colony lineage, even when they disperse into new areas. These results indicate that the fine scale metapopulation structure of pure matrilines is maintained over the long term and that colony turnover is not extensive or radical enough to homogenize entire geographical areas. Genetic diversity is thus preserved to some extent at the metapopulation level.     

Behavioural and biomaterial coevolution in spider orb webs

Mechanical performance of biological structures, such as tendons, byssal threads, muscles, and spider webs, is determined by a complex interplay between material quality (intrinsic material properties, larger scale morphology) and proximate behaviour. Spider orb webs are a system in which fibrous biomaterials—silks—are arranged in a complex design resulting from stereotypical behavioural patterns, to produce effective energy absorbing traps for flying prey. Orb webs show an impressive range of designs, some effective at capturing tiny insects such as midges, others that can occasionally stop even small birds. Here, we test whether material quality and behaviour (web design) co‐evolve to fine‐tune web function. We quantify the intrinsic material properties of the sticky capture silk and radial support threads, as well as their architectural arrangement in webs, across diverse species of orb‐weaving spiders to estimate the maximum potential performance of orb webs as energy absorbing traps. We find a dominant pattern of material and behavioural coevolution where evolutionary shifts to larger body sizes, a common result of fecundity selection in spiders, is repeatedly accompanied by improved web performance because of changes in both silk material and web spinning behaviours. Large spiders produce silk with improved material properties, and also use more silk, to make webs with superior stopping potential. After controlling for spider size, spiders spinning higher quality silk used it more sparsely in webs. This implies that improvements in silk quality enable ‘sparser’ architectural designs, or alternatively that spiders spinning lower quality silk compensate architecturally for the inferior material quality of their silk. In summary, spider silk material properties are fine‐tuned to the architectures of webs across millions of years of diversification, a coevolutionary pattern not yet clearly demonstrated for other important biomaterials such as tendon, mollusc byssal threads, and keratin.     

Morphological phylogeny of cobweb spiders and their relatives (Araneae, Araneoidea, Theridiidae)

This paper offers the first cladistic analysis of a wide selection of theridiid genera based on morphological data. The analysis treats 53 theridiid taxa representing 32 genera (Achaearanea, Anelosimus, Ameridion, Argyrodes, Ariamnes, Carniella, Cerocida, Chrysso, Coleosoma, Dipoena, Emertonella, Enoplognatha, Episinus, Euryopis, Faiditus, Kochiura, Latrodectus, Neospintharus, Nesticodes, Pholcomma, Phoroncidia, Rhomphaea, Robertus, Selkirkiella, Spintharus, Steatoda, Stemmops, Theridion, Theridula, Thymoites, Thwaitesia, Tidarren) and eight outgroup taxa representing the families Nesticidae (Eidmanella and Nesticus), Synotaxidae (Synotaxus, two species), Pimoidae (Pimoa), Linyphiidae (Linyphia), Tetragnathidae (Tetragnatha) and Araneidae (Argiope). The parsimony analysis of 242 morphological and behavioural characters found a single, most parsimonious tree. The monophyly of theridiids and their sister relationship with nesticids is strongly supported. The recent resurrection of Ariamnes and Rhomphaea from Argyrodes made the latter paraphyletic. However, Ariamnes and Rhomphaea are characterized by an array of characters, and Argyrodes still contains dramatically distinct clades for which names are available: Faiditus (removed from synonymy ? RS) and Neospintharus (RS). These revalidations provide a classification with greater information content and utility. These three genera, along with Ariamnes, Rhomphaea and Spheropistha, comprise the subfamily Argyrodinae. The monophyly and composition of the subfamilies Hadrotarsinae, Spintharinae, Pholcommatinae, Latrodectinae and Theridiinae are discussed. Theridion is paraphyletic and in need of revision. Anelosimus as currently circumscribed is paraphyletic, a problem resolved by revalidating Selkirkiella (RS) and Kochiura (RS). Numerous new combinations are established. The results suggest the monophyletic origin of both kleptoparasitism and araneophagy in the lineage leading to Argyrodinae, negating hypotheses that either arose from the other. Sociality evolved multiple times within the family, accounting for as much as one fourth of the origins of social behaviour among all spiders. No losses of sociality are implied. The hypothesis of maternal care as the pathway to sociality receives support. Evolution of theridiid webs is complex, with multiple modifications and loss of the basic theridiid cobweb. © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 141 , 447–626.