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SUMMARYMytilid bivalves employ a set of threads (the byssus) to attach themselves to both hard and soft substrates. In this study, we measured the mechanical properties of byssal threads from two semi-infaunal mytilids (Geukensia demissa Dillwyn and Modiolus modiolus Linnaeus) and two epifaunal mytilids (Mytilus californianus Conrad and Mytilus edulis Linnaeus). We compared material properties with and without the assumption that changes of length and area during tensile testing are insignificant, demonstrating that previous researchers have overestimated extensibility values by 30% and may also have underestimated strength values. We detected significant differences in thread properties among tested mytilid species, contrary to previous findings. Threads from semi-infaunal species were significantly thinner than those from epifaunal species, perhaps to allow the production of a greater number of threads, which form a dense network within the substrate. Geukensia demissa threads were weaker than those of the other species, and had a significantly lower stiffness at failure. Modiolus modiolus threads were significantly stiffer than M. edulis threads but also significantly less extensible, suggesting a trade-off between stiffness and extensibility. The only thread property that did not show significant differences across species was toughness -even when byssal threads differ in strength or stiffness, they seem to absorb similar amounts of energy per unit volume prior to failure. This study reveals notable differences between the byssal thread properties of different mytilid bivalves and provides a reliable and thorough methodology for future comparative studies. Supplementary material available online at

In their 2010 book, Biology's First Law, D. McShea and R. Brandon present a principle that they call ''ZFEL,'' the zero force evolutionary law. ZFEL says (roughly) that when there are no evolutionary forces acting on a population, the population's complexity (i.e., how diverse its member organisms are) will increase.Here we develop criticisms of ZFEL and describe a different law of evolution; it says that diversity and complexity do not change when there are no evolutionary causes.

SUMMARYThe byssus is the set of proteinaceous threads widely used by bivalves to attach themselves to the substrate. Previous researchers have focused on a single byssate family, the Mytilidae. However, the properties of byssal threads from species outside this family are of interest -first, because evolutionary patterns are only detectable if species from a range of taxa are examined, and second, because recent biomimetic research efforts would benefit from a wider range of 'mussel glue' exemplars. In the present study, we measured the mechanical properties of the byssal threads of two species outside the Mytilidae, the pen shell Atrina rigida Lightfoot and the flame 'scallop' Ctenoides mitis Lamarck. The mechanical properties of their byssal threads were significantly different from those of mytilids. For instance, the byssal threads of both species were significantly weaker than mytilid threads. Atrina rigida threads were significantly less extensible than mytilid threads, while C. mitis threads exhibited the highest extensibility ever recorded for the distal region of byssal threads. However, there were also interesting similarities in material properties across taxonomic groups. For instance, the threads of A. rigida and Modiolus modiolus Linnaeus both exhibited a prominent double-yield behavior, high stiffness combined with low extensibility, and similar correlations between stiffness and other thread properties. These similarities suggest that the thread properties of some semi-infaunal species may have evolved convergently. Further research on these patterns, along with biochemical analysis of threads which exhibit unusual properties like double-yield behavior, promises to contribute to both evolutionary biology and materials engineering. Supplementary material available online at

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