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 (

Relationships among, and content of, earthworm families have been controversial and unstable. Here we analyse molecular data from 14 Crassiclitellata families represented by 54 genera, the non-crassiclitellate ‘earthworms’ of the Moniligastridae, plus several clitellate outgroups. Complete 28S and 18S gene sequences and a fragment of the 16S gene analysed separately or in concatenated Bayesian analyses indicate that most previously proposed suprafamilial taxa within the Crassiclitellata are para- or polyphyletic. There is strong support for the Metagynophora, which consists of the Crassiclitellata and Moniligastridae. The most basal within-Clitellata branch leads to the small families Komarekionidae, Sparganophilidae, Kynotidae, and Biwadrilidae, found in widely separated areas. A clade composed of Lumbricidae, Ailoscolecidae, Hormogastridae, Criodrilidae and Lutodrilidae appears near the base of the tree, but Criodrilidae and Biwadrilidae are not closely related because the former is sister to the Hormogastridae + Lumbricidae clade. The Glossoscolecidae is here separated into two families, the Glossoscolecidae s.s. and the Pontoscolecidae (fam. nov.). The Megascolecidae is monophyletic within a clade including all acanthodrilid earthworms. There is strong support for the Benhamiinae (Acanthodrilidae s.l.) as sister to Acanthodrilidae + Megascolecidae, but taxon sampling within other acanthodrilid groups was not sufficient to reach further conclusions. The resulting trees support revised interpretations of morphological character evolution.’

JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact support@jstor.org.Abstract. Earthworms represent the dominant fraction of soil macroinvertebrate biomass in Kansas tallgrass prairie and thus may have a large impact on nutrient budgets. To evaluate this impact I estimated cast production and rates of nutrient processing by the native genus Diplocardia and by exotic Lumbricidae in permanent plots on two soil types at Konza Prairie Research Natural Area. Total cast production was estimated from field population estimates, soil climate data, and cast production-temperature relationships obtained in the laboratory. The organic matter and mineral N and P contents of surface casts by month were used to estimate total processing of these substances in the field. Earthworms present in the study sites were Diplocardia longiseta, D. kansensis, D. rugosa, D. singularis, D. smithii, D. verrucosa, and the European species Aporrectodea caliginosa and Octolasion cyaneum. Estimated total cast production and nutrient processing by Diplocardia spp. exceeded that of exotics in both soil types, in spite of higher populations of Lumbricidae in one soil. Total annual soil consumption by all earthworms was 4-10% of the A-horizon, depending on soil type. Organic matter equivalent to 10% of total soil organic matter in the top 15 cm or to 100-300% of plant annual belowground production appears to pass through the earthworms each year. Mineral N processed was -10-12% of annual plant N uptake, comparable to half of the input from precipitation, while the P processed was equivalent to 50% of annual uptake. In contrast to other grassland systems studied, the introduction of Lumbricidae had a negative effect on soil turnover and nutrient mineralization due to the lower throughput of Lumbricidae and their relative intolerance of summer soil temperatures.

Clitellata is a major clade of Annelida comprising nearly all freshwater and terrestrial annelids as well as several marine species. We investigated clitellate phylogenetic relationships using transcriptomes sampled from 74 taxa (64 clitellates and 10 polychaetes), including multiple representatives of nearly all major clitellate higher taxa (Branchiobdellida, Capilloventridae, Crassiclitellata, Enchytraeidae, Haplotaxidae, Hirudinida, Lumbriculida, Moniligastridae, Naididae, Parvidrilidae, Phreodrilidae, Propappidae and Randiellidae). We used a number of filtered data matrices and phylogenetic analyses to examine the effects of data partitioning, missing data and compositional and branch‐length heterogeneity and used the resulting phylogenies for divergence time estimation and ancestral habitat reconstructions. All analyses and filtering methods produced a consistent, strongly supported topology in which (a) Enchytraeidae, Hirudinida, Hirudinea (here, Branchiobdellida plus Hirudinida), Lumbriculida, Lumbriculata (Lumbriculida plus Hirudinea), Phreodrilidae and Naididae are monophyletic, (b) a Parvidrilidae + Randiellidae clade is sister to the rest of Clitellata, (c) Phreodrilidae is sister to Naididae, (d) Haplotaxidae is non‐monophyletic, with some haplotaxids grouping with Crassiclitellata + Moniligastridae, (e) the Phreodrilidae + Naididae clade is sister to all other clitellates except Parvidrilidae + Randiellidae and Capilloventridae, and (f) Lumbriculata is sister to the Crassiclitellata + Moniligastridae + Haplotaxidae (in part) clade. Ancestral habitat reconstructions and divergence time analysis suggested that the most recent common ancestor of Clitellata lived in freshwater during the Devonian (419–359 million years ago) and that all major extant clitellate lineages arose over the next ~150 million years, with multiple lineages subsequently returning to marine habitats or invading land. This study provides a phylogenetic framework for further investigation of the geological, environmental and biotic forces and genomic changes that may have impacted clitellate evolution and enabled several major habitat transitions within this group.

Araucaria angustifolia, also known as the Paraná Pine is an endangered tree species in Brazil and little is known of the diversity of soil invertebrates inhabiting these forests. Therefore, the present study was set up to evaluate the biomass and diversity of earthworms in natural and reforested Araucaria plots, impacted or not by fire, and to identify the most efficient earthworm collection method. Four study areas included: native forest with Araucaria (NF); Araucaria reforestation (R); Araucaria reforestation submitted to an accidental fire (RF); and native grass pasture with native Araucaria and submitted to an intense accidental fire (NPF). Five soil samples containing the earthworm community were taken in a 0.3 ha area in each of the forest sites, close to five Araucaria trees selected at random. Three collection methods were tested: application of dilute Formol (0.5%) to the soil surface, handsorting of small (25 × 25 cm) or large (40 × 40 cm) monoliths. Five earthworm species were found: the native Glossoscolex sp.1, Glossoscolex sp.2, Glossoscolex bondari and Urobenus brasiliensis (Glossoscolecidae), and the exotic Amynthas corticis (Megascolecidae). Formol was more efficient for collecting A. corticis, found in much higher abundance and biomass in NF than in the other areas. Larger handsorted samples were more efficient for capturing Glossoscolex species, mainly present in RF and NPF. For adequate characterization of earthworm abundance and biomass in these Araucaria forests, both the Formol and the larger monolith methods are recommended.