Elementary ecology texts tell us that organisms interact in three fundamental ways, generally given the names competition, predation, and mutualism. The third member has gotten short shrift (264), and even its name is not generally agreed on. Terms that may be considered synonyms, in whole or part, are symbiosis, commensalism, cooperation, protocooperation, mutual aid, facilitation, reciprocal altruism, and entraide. We use the term mutualism, defined as "an interaction between species that is beneficial to both," since it has both historical priority (311) and general currency. Symbiosis is "the living together of two organisms in close association," and modifiers are used to specify dependence on the interaction (facultative or obligate) and the range of species that can take part (oligophilic or polyphilic). We make the normal apologies concerning forcing continuous variation and diverse interactions into simple dichotomous classifications, for these and all subsequent definitions.

Earthworms belonging to the family Lumbricidae are extremely abundant in terrestrial temperate regions. They affect soil properties and nutrient cycling, thus shaping plant community composition and aboveground food webs. Some lumbricids are also model organisms in ecology and toxicology. Despite the intense research efforts dedicated to lumbricids over the last 130 years, the evolutionary relationships and taxonomic classification of these organisms are still subject to great debate. Resolution of their systematics is hampered by the structural simplicity of the earthworm body plan and the existence of cryptic species. We sampled 160 earthworm specimens belonging to 84 lumbricid species (28 genera) and 22 Lumbricoidea outgroups, sequenced two nuclear genes, four mitochondrial genes and seven mitochondrial tRNAs and examined 22 morphological characters. We then applied a combination of phylogenetic methods to generate the first robust genus-level phylogeny of the Lumbricidae. Our results show that the current Lumbricidae classification and the underlying hypotheses of character evolution must be revised. Our chronogram suggests that lumbricids emerged in the Lower Cretaceous in the holarctic region and that their diversification has been driven by tectonic processes (e.g. Laurasia split) and geographical isolation. Our chronogram and character reconstruction analysis reveal that spermathecae number does not follow a gradual pattern of reduction and that parthenogenesis arose from sexual relatives multiple times in the group; the same analysis also indicates that both epigeic and anecic earthworms evolved from endogeic ancestors. These findings emphasize the strong and multiple changes to which morphological and ecological characters are subjected, challenging the hypothesis of character stasis in Lumbricidae.

The widely studied and invasive earthworm, Lumbricus terrestris L., 1758 has been the subject of nomenclatural debate for many years. However these disputes were not based on suspicions of heterogeneity, but rather on the descriptions and nomenclatural acts associated with the species name. Large numbers of DNA barcode sequences of the cytochrome oxidase I obtained for nominal L. terrestris and six congeneric species reveal that there are two distinct lineages within nominal L. terrestris. One of those lineages contains the Swedish population from which the name-bearing specimen of L. terrestris was obtained. The other contains the population from which the syntype series of Enterion herculeum Savigny, 1826 was collected. In both cases modern and old representatives yielded barcode sequences allowing us to clearly establish that these are two distinct species, as different from one another as any other pair of congeners in our data set. The two are morphologically indistinguishable, except by overlapping size-related characters. We have designated a new neotype for L. terrestris. The newly designated neotype and a syntype of L. herculeus yielded DNA adequate for sequencing part of the cytochrome oxidase I gene (COI). The sequence data make possible the objective determination of the identities of earthworms morphologically identical to L. terrestris and L. herculeus, regardless of body size and segment number. Past work on nominal L. terrestris could have been on either or both species, although L. herculeus has yet to be found outside of Europe.

Biological invasions are increasingly recognized as a potent force altering native ecosystems worldwide. Many of the best documented cases involve the massive invasions of North America by plant and animal taxa native to Europe. In this study, we use DNA barcoding to survey the occurrence and genetic structure of two major groups of soil invertebrates in both their native and introduced ranges:Collembola and earthworms. Populations of ten species of earthworms and five species of Collembola were barcoded from both continents. Most of these species exhibited a similar genetic structure of large and stable populations in North America and Europe, a result supporting a scenario of multiple invasions. This was expected for earthworm species involved in human economic activities, but not foreseen for Collembola species de facto unintentionally introduced. This study also establishes that invasive species surveys employing DNA barcoding gain additional resolution over those based on morphology as they Electronic supplementary material The online version of this article (