A review of the earthworm Amynthasmasatakae (Beddard, 1892) (Clitellata, Megascolecidae), with designation of two new synonyms.

Correct and timely identification of an invasive species during quarantine or at an early stage of invasion before establishment or spread is critical for preventing biological invasions. However, taxonomic confusion of potential invasive earthworm species caused by incorrect taxonomic treatment or reckless taxonomic work has made it difficult to properly recognize potential invasion threats. Through analyzing publicly available DNA sequences of the mitochondrial cytochrome c oxidase subunit I (COI) gene, we confirmed the validity of the specific status of Amynthasmasatakae (Beddard, 1892), a peregrine earthworm species in East Asia with the potential to spread to other regions of the world, and designated two new synonyms of A.masatakae: Amynthastralfamadore Blakemore, 2012 syn. nov. and Amynthasscaberulus Sun and Jiang, 2021 syn. nov. Additionally, the name A.triastriatususualis Dong, Jiang, Yuan, Zhao and Qiu, 2020 is nomenclaturally unavailable since it was published in an electronic journal without ZooBank registration and an explicit statement establishing a new nominal taxon. Specimens described under this unavailable name actually belong to A.masatakae. Inadequate literature review and erroneous species identities associated with sequences in GenBank have caused even more problems in the already confusing earthworm taxonomy.

Complete mitochondrial genome of the composting worm Dendrobaena veneta (Clitellata: Oligochaeta, Lumbricidae).

Dendrobaena veneta (Rosa, 1886) is widely distributed all over Europe due to its use as compost worm. The specimen presented here was collected in Tiranë district, Albania. Currently, only two species' complete or nearly complete mitochondrial genome (mitogenome) sequences have been reported in the genus Dendrobaena; D. octaedra (Savigny, 1826) and D. tellermanica Perel, 1966. In this study, the complete mitogenome of D. veneta was sequenced, assembled, and annotated. The mitogenome of D. veneta is a circular DNA molecule, consisting of 15,475 bp with an A + T content of 61.2%. It contains 13 protein-coding genes, 2 ribosomal RNA genes, 22 transfer RNA genes, and 1 non-coding region (control region). Phylogenetic analysis showed that D. veneta is clustered with the other two Dendrobaena species in the well-supported family Lumbricidae.

Earthworms (Clitellata, Megadrili) of the world: an updated checklist of valid species and families, with notes on their distribution.

In the current paper we present an updated checklist of all the megadrile earthworms (Crassiclitellata: Annelida) in the world, and notes on the distribution of families worldwide. Biogeographic responses to geological phenomena including plate tectonics, as well as to past and present climate and habitat distributions, are the main factors determining the present distribution of earthworm families. A total of ca. 5,738 species/subspecies (5,406 species and 332 unique subspecies; i.e., not counting the nomino-typical subspecies) belonging to 23 families (including one non-crassiclitellate family: Moniligastridae) are currently recognized worldwide, of which three families (Tritogeniidae and Kazimierzidae from Southern Africa and Arecoidae, a new family from Brazil described herein), 35 genera and close to 1200 new taxa (including subspecies) were described in the 21st century. Nonetheless, the large number of still undescribed species will likely increase this value to well over 8,000 species. Ten families are monospecific and/or monogeneric and have a mostly restricted distribution. On the other hand, more than 87 widespread cosmopolitan species have been catalogued, some of them with important invasive potential, belonging mainly to families Lumbricidae, Acanthodrilidae, Benhamiidae, Megascolecidae, Rhinodrilidae and Ocnerodrilidae. Taxonomic housekeeping was performed for the preoccupied Rhinodrilidae genus Tairona Righi - herein substituted by Taironina nom. nov., and Guarani camaqua Rodríguez & Lima was reinstated and removed from synonymy with Criodrilus lacuum Hoffmeister, 1845, resulting in a wider definition of the Almidae family. Furthermore, Amynthas maximalis nom. nov. is proposed herein as a substitution name for the preoccupied name Amynthas maximus Qiu & Dong, 2019, and Arecoidae is proposed herein as a new monotypic family for the aquamegadrile species Areco reco Righi, Ayres & Bittencourt, 1978.

Complete Mitochondrial genome of Eisenia nordenskioldi pallida Malevich, 1956 from Korea, with remarks on the phylogeny of the E. nordenskiodi complex (Megadrili; Lumbricidae).

The Eisenia nordenskioldi (Eisen, 1879) species complex is one of the most widely distributed earthworm taxa in northern Asia, owing to its high morphological and genetic variability. This complex is represented by smaller, slightly pigmented- or unpigmented specimens in the Korean Peninsula. Here, we report the mitogenomes of two Eisenia nordenskioldi cf. pallida specimens, No. 1 and No. 3, which are slightly pigmented and unpigmented, respectively. The total length of the mitogenomes was 15,824 bp and 15,177 bp, respectively, and contained 37 genes, comprising 13 protein-coding genes (PCGs), two rRNA genes, 22 tRNA genes, and a putative non-coding region, as in other earthworms. Nucleotide sequence comparisons of the 37 genes and the predicted protein sequences of 13 PCGs showed 96.6 % and 98.6 % similarity between the two mitogenomes, respectively. A phylogenetic analysis of 33 Crassiclitellata mitogenomes corroborated the monophyly of the Eisenia genus and recovered two highly supported subclades in the E. nordenskioldi species complex. The Korean specimens were the closest to the E. n. pallida L2 specimens collected near the type locality and probably represent the genuine E. nordenskioldi pallida Malevich, 1956.

On the molecular taxonomy of the Dendrobaena alpina species complex, with description of a new species (Crassiclitellata, Lumbricidae).

The molecular phylogenetic analysis of the D. alpina species group and related taxa revealed that this group in the present form is polyphyletic. The dark-red pigmented D. alpina alpina (Rosa, 1884) specimens from the Alps form a distinct clade together with D. alpina alteclitellata (Pop, 1938) and D. clujensis Pop, 1938 (Central European clade). The unpigmented Bulgarian specimens are now part of a clade consisting of Anatolian and Levantine species, namely D. orientalis Cernosvitov, 1940, D. pentheri (Rosa, 1905), D. orientaloides Zicsi, 1985 and D. semitica Rosa, 1893a. Consequently, the unpigmented worms from Bulgaria represent a new species described herein as Dendrobaena misirlioglui sp. nov.

The complete mitochondrial DNA sequences of two sibling species of lumbricid earthworms, Eiseniafetida (Savigny, 1826) and Eiseniaandrei (Bouché, 1972) (Annelida, Crassiclitellata): comparison of mitogenomes and phylogenetic positioning.

Composting earthworms of the genus Eisenia play an important role in soil ecosystems. However, taxonomic classification of this genus, especially the sibling species Eiseniafetida and Eiseniaandrei, is complicated because of their morphological similarity. In this study, we assessed the utility of the complete mitochondrial genome (mitogenome) for identification and differentiation of the two species. The complete mitogenomes of E.andrei and E.fetida were 15,714 and 16,560 bp, respectively. They contained 37 genes, comprising 13 protein-coding genes (PCGs), two rRNA genes, 22 tRNA genes, and a putative non-coding region, as observed in other earthworms. Sequence comparisons based on the complete nucleotide sequences excluding the non-coding region showed 85.8% similarity, whereas the predicted amino acid sequences of the 13 PCGs were 92.7% similar between the two species. In particular, distinct features were found in the non-coding regions of the mitogenomes. They include a control region associated with putative mitogenome replication and an extended sequence. The extended sequence showed significant differences between the two species and other known earthworm species, suggesting its potential as a feasible molecular marker for species identification. Phylogenetic analysis of the 36 mitogenomes of earthworm species corroborated the monophyly of the genus Eisenia and the taxonomic distinctness of the sibling species pair, E.fetida and E.andrei.

Integrative taxonomy of some Iranian peregrine earthworm species using morphology and barcoding (Annelida: Megadrili).

Despite the biological and economic importance of earthworms, the taxonomic status and evolutionary relationships of most lumbricid genera are still under debate. Further complications arise from the recognition that earthworms also show a high cryptic diversity. Past and current field studies of Iranian earthworm fauna have resulted in the identification of a total number of 28 earthworm species. However, many specimens do not fully fit into their original descriptions, making the species assignation very difficult. In this study, we evaluated the genetic diversity using mitochondrial markers as a tool to assess the species occurrence of some problematic species in Iran. Four species with high morphological variation were selected: Aporrectodea caliginosa (Savigny, 1826), Aporrectodea trapezoides (Dugès, 1828), Dendrobaena byblica (Rosa, 1893) and Dendrobaena veneta (Rosa, 1886). Morphological identification was contrasted with the molecular information generated through COI and 16S barcoding and the COI and 16S sequences stored in the Genbank. The results of this first integrative taxonomic analysis revealed that D. veneta consisted of two separated clades and that a number of species assigned to D. byblica showed very close relationships with those belonging to the genus Philomontanus. The lack of taxonomic expertise and identification characters providing a clear and unambiguous identification of earthworms highlights the urgent need of new tools to identify species unequivocally. Therefore, it is concluded that more taxonomical studies are needed to clarify the diagnostic characters and taxonomic status of the species belonging to two genera, Aporrectodea and Dendrobaena (Lumbricidae), in Iran.

Description of a new Central African earthworm, Petroscolex centenarius gen. et sp. nov. (Crassiclitellata, Eudrilidae), celebrating the 100th birthday of Pietro Omodeo.

Prof. Pietro Omodeo (University of Siena, Italy), the world-renowned earthworm taxonomist and evolutionary biologist, was born in Cefalù, Sicily, Italy on the 27th September, 1919. He celebrates his 100th birthday in 2019 and members of the international community of earthworm taxonomists salute him with Petroscolex centenarius gen. et sp. nov., a new megadrile taxon discovered in 1991 by him but which has not been formally described until now. The many important contributions of Omodeo to oligochaetological research are briefly mentioned.

Annotated checklist of earthworm species described by András Zicsi (Clitellata: Megadrili).

Professor András Zicsi, the renowed taxonomist and ecologist of earthworms, passed away on 22 July, 2015. He had a decisive impact on earthworm taxonomy in the second half of the 20th century, describing 10 new genus group taxa and 243 species / subspecies new to science distributed in 10 families. A complete list of the new taxa described by Prof. Zicsi is presented here together with references to the original descriptions and the valid names.

DNA barcoding of the Italian anecic Octodrilus species in rural (vineyard) and forested areas with description of Octodrilus zicsiniello sp. nov. (Clitellata, Megadrili).

DNA barcoding of 172 anecic Octodrilus specimens collected in NE Italy and bordering Croatia has been carried out. The Bayesian phylogenetic tree showed high support for almost all currently recognized species, however, some unexpected results also appeared. The clade representing Oc. pseudocomplanatus contains a highly advanced subclade, which morphologically resembles Oc. slovenicus. The highly supported Oc. tergestinus clade consists of four unresolved divergent lineages of which the first corresponds to Oc. istrianus and the second resembles Oc. mimus morphologically; the third and fourth clades show typical tergestinus characters. The widely distributed Oc. complanatus consists of three highly divergent subclades which are sister to a new species, Oc. zicsiniello sp. nov., hereunder described.

Solving a nomenclatural conundrum: Cernosvitovia crainensis (Mrsic, 1989) and Aporrectodea macvensis Sapkarev in Mrsic, 1991 (Lumbricidae).

This contribution deals with the names and authorship of two lumbricid taxa endemic to the Balkans (see Stojanovic et al., this volume). Although their validity has never been questioned, it has been unclear up to now which publication has made these two species-group names available according to the rules of the International Code of Zoological Nomenclature (Articles in "the Code," ICZN 1999). In the following, we review the somewhat intricate history of these names and explain why the correct citation and spelling of these names are "Cernosvitovia crainensis (Mrsic, 1989)" and "Aporrectodea macvensis Sapkarev in Mrsic, 1991," respectively.

Contribution to the knowledge of the earthworm fauna of Turkey with description of three new species (Clitellata: Megadrili).

During a recent survey, 29 earthworm species were collected from different parts of Turkey, 27 of which belong to the family Lumbricidae, one to Acanthodrilidae and one to Megascolecidae. Dendrobaena proved to be the most speciose genus with 12 recorded species including two species new to science, Dendrobaena pavliceki and Dendrobaena taurica spp. nov. In addition a third new species was also found belonging to the East Mediterranean genus Healyella: Healyella zicsii sp. nov. From the species recorded earlier for Turkey, D. fridericae uludagi Omodeo Rota, 1991 and He. boluana Omodeo Rota, 1989 were found for the first time since the original description. With these new records the number of earthworm species recorded for Turkey is raised to 84.

Resurrection of the earthworm species Dendrobaena fedtschenkoi (Michaelsen, 1900), a former synonym of Dendrobaena byblica (Rosa, 1893) (Clitellata: Megadrili).

Dendrobaena byblica (Rosa, 1893) is a Circum-Mediterranean earthworm species complex containing more than a dozen synonymous names. From these, only two species have been resurrected so far, D. ganglbaueri (Rosa, 1894) and Fitzingeria annectens (Rosa, 1895). Here we demonstrate that Dendrobaena fedtschenkoi (Michaelsen, 1900) described from the Pamir Mts. Tajikistan differs from D. byblica both in morphological and molecular features, and represents an independent species.

Litter quality, dispersal and invasion drive earthworm community dynamics and forest soil development.

In temperate deciduous forests of eastern USA, most earthworm communities are dominated by invasive species. Their structure and functional group composition have critical impacts on ecological properties and processes. However, the factors determining their community structure are still poorly understood, and little is known regarding their dynamics during forest succession and the mechanisms leading to these changes. Earthworm communities are usually assumed to be stable and driven by vegetation. In contrast, the importance of dispersal and ecological drift is seldom acknowledged. By analyzing a 19-year dataset collected from forest stands in eastern USA, we demonstrated that on a decadal timescale, earthworm community dynamics are shaped by the interplay of selection, dispersal, and ecological drift. We highlighted that forests at different successional stages have distinct earthworm species and functional groups as a result of environmental filtering through leaf litter quality. Specifically, young forests are characterized by soil-feeding species that rely on relatively fresh soil organic matter derived from fast-decomposing litter, whereas old forests are characterized by those feeding on highly processed soil organic matter derived from slow-decomposing litter. In addition, year-to-year species gains and losses are primarily driven by dispersal from regional to local species pools, and by local extinction resulted from competition and ecological drift. We concluded that with continued dispersal of European species and the recent "second wave" of earthworm invasion by Asian species from the surrounding landscape, earthworms at the investigated forests are well-established, and will remain as the major drivers of soil development for the foreseeable future.

Molecular phylogeny and systematics of native North American lumbricid earthworms (Clitellata: Megadrili).

The family Lumbricidae is arguably the most well-known and well-studied earthworm group due to its dominance in the European earthworm fauna and its invasion in temperate regions worldwide. However, its North American members, especially the genus Bimastos Moore, 1893, are poorly understood. We revised the systematics of the genus Bimastos and tested the hypothesis of the monophyly of North American lumbricids using morphological characters and eight molecular markers. Phylogenetic analyses based on our extensive sampling of Bimastos and inclusion of Dendrodrilus and Allolobophoridella indicated a well-supported clade containing Bimastos and Eisenoides Gates, 1969, and provided the first evidence supporting that North American lumbricids are monophyletic. Assuming the available divergence time estimations and dating of land bridges are correct, it would suggest that the ancestor of this clade arrived North America through Beringia or the De Geer route during Late Cretaceous, and since then the clade has diverged from its Eurasian sister group, Eisenia. The peregrine genera Dendrodrilus and Allolobophoridella are nested within the Bimastos clade; we propose to treat them as junior synonyms of the genus Bimastos, and, contradictory to the commonly held belief of being European, they are indeed part of the indigenous North American earthworm fauna. Morphological characters, such as red-violet pigmentation, proclinate U-shaped nephridial bladders and calciferous diverticula in segment 10 further support this placement. The East Mediterranean-Levantine Spermophorodrilus Bouché, 1975 and Healyella Omodeo & Rota, 1989 are nested within the Dendrobaena sensu lato clade; therefore their close relationship with the North American Bimastos is refuted. Species fit the revised diagnosis of Bimastos are reviewed and keyed, and a new species, Bimastos schwerti sp. nov., is described.

An annotated checklist of earthworms of Greece (Clitellata: Megadrili).

The earthworm fauna of Greece is reviewed. According to the up-to-date checklist, the earthworm fauna of Greece consists of 67 species and subspecies, of which 59 taxa belong to the family Lumbricidae, three to Megascolecidae, two to Acanthodrilidae and to Ocnerodrilidae and one taxon to the family Criodrilidae. Three species are recorded for the first time from the country: Allolobophora kosowensis kosowensis Karaman, 1968, Amynthas gracilis (Kinberg, 1867) and Eukerria saltensis (Beddard, 1895). Eisenia spelaea var. athenica Cernosvitov, 1938 is proposed to be a synonym of Aporrectodea rosea (Savigny, 1826). The earthworm fauna of Greece is characterized by a large number of strict endemic species belonging to the family Lumbricidae (9 taxa), together with the occurrence of another 10 Balkanic endemic species.

Correction: Barcoding Eophila crodabepis sp. nov. (Annelida, Oligochaeta, Lumbricidae), a Large Stripy Earthworm from Alpine Foothills of Northeastern Italy Similar to Eophila tellinii (Rosa, 1888).

[This corrects the article DOI: 10.1371/journal.pone.0151799.].