Search

SCHEROTHECA ALBINA MARCHÁN & DECAËNS SP. NOV. (FIGS 2 A, 3 G, H, 4) Table 3. Continued . 1214.31311.310.211.216.311.710.710.6 0 Zoobank registration: urn: lsid: zoobank. org:act: BDD2FFC5-C311-4CA5-B6C8-227910470E04. Type material: Holotype. France • adult; Occitanie, Hérault, le Soulié; latitude/longitude: 43.531 / 2.658; elevation: 853 masl; 06-May-2021;T.Decaëns, D.Fernández Marchán leg.; BOLD Sample ID: DFM-0572; deposited in the Musée national d’Histoire naturelle, Paris. Paratypes: France • five adult specimens, one subadult specimen; same data as holotype (four specimens); BOLD Sample ID: DFM-0573, DFM-0574, DFM-0575, DFM-0576; Occitanie, Hérault, Saint-Etienne-d’Albagnan; latitude/longitude: 43,543/2,86; elevation: 250 m asl; 14-Apr-2021; T. Decaëns, D. Fernández Marchán leg. (two specimens); BOLD Sample ID: DFM-0025, DFM-0026; specimens DFM-0573, DFM-0575, DFM-0576 deposited in Eco&Sols collection, Montpellier; DFM-0025, DFM-0026, DFM-0574 deposited at Musée national d’Histoire naturelle, Paris. Etymology: The species name is derived from the lack of cutaneous pigmentation which characterize it by opposition to the usually pigmented continental species of Scherotheca. Treated as adjectival. Diagnosis: Scherotheca albina can be distinguished from Sc. rhodana (Bouché, 1972) (the most similar species) by the position of the clitellum (XXVII–XLI vs. (1/2 XXVI) 1/2 XXVII–1/2 XLII (1/2 XLIII)) and tubercula pubertatis (XXXI–1/n XLI vs. 1/n XXXII– XLI), by its smaller body size and by the absence of cutaneous pigmentation (Table 2). COI uncorrected average pairwise distances and topology of multilocus molecular phylogenetic trees supports the status of Sc. albina as independent from other morphologically related species. Description External morphology: Body pigmentation absent or faint brown in the anterior part of the body (Fig. 2a). Beige with whitish clitellum in fixed specimens (Fig. 3g, h). Average length 15.15 cm (13.16–17.2 cm, N = 2 adults); average diameter 0.64/0.73/ 0.59 cm (preclitellar/clitellar/postclitellar); body subcylindrical in cross-section, ventrally flattened; average number of segments 249 (244–253, N = 2 adults). Average weight (fixed specimens), 4.88 g (4.37–5.39 g, N = 2 adults). Prostomium epilobous, closed. Longitudinal furrows in segments I and II. Nephridial pores ‘en solfège’ (irregularly distributed). Spermathecal pores at intersegmental furrows 9/10–12/13(13/14). Male pores in segment XV, surrounded by a moderately developed porophore. Female pores in segment XIV. Clitellum saddle-shaped in segments XXVII–XLI. Tubercula pubertatis in segments XXXI–1/n XLI. Setae closely paired. Internal anatomy: Septa 5/6–10/11 thickened and muscular. Lateral hearts in segments VI–XI. Calciferous glands in segments X– XIV, with diverticula in segment X. Crop in segments XV–XVI, gizzard in segments XVII–XIX. Typhlosole pinnate. Male sexual system holandric, testes and funnels (not enclosed in testes sacs, but with sperm present) located ventrally in segments X and XI. Four pairs of reniform seminal vesicles in segments IX, X, XI and XII, the first two pairs being smaller. Ovaries and female funnels in segment XIII, ovarian receptacles (ovisacs) in segment XIV. Four or five pairs of small globular spermathecae in segments IX, X, XI, XII (XIII) (intersegments 9/10– 12/13(13/14)), simple. Nephridial bladders U-shaped, reclinate in segment XXX. Distribution and ecology: Scherotheca albina was found in the vicinities of the Regional Park of Haut-Languedoc, Hérault, Occitanie (Fig. 4), in managed grasslands surrounded by forests.

SCHEROTHECA HAYMOZI (ZICSI, 1977) COMB.NOV. Allolobophora haymozi Zicsi, 1977. Aporrectodea haymozi Zicsi, 1977: in Zicsi and Csuzdi, 1999. Aporrectodea haymoziformis Zicsi and Csuzdi, 1999. Remarks: The original descriptions of Al. haymozi and Ap. haymoziformis differ slightly in the position of the clitellum (XXXIII–1/2 XL, XL vs. 1/2 XXXIII, XXXIII, XXXIV–XL, XLI), tubercula pubertatis (XXXV, XXXVI–XL vs. 1/2 XXXIV, XXXIV–XL, 1/2 XLI) and spermathecae (XII, XIII, XIV vs. XI, XII, XIII, XIV). However, the results of the molecular phylogenetic analysis and COI genetic distances clearly indicate that these morphological differences correspond to intraspecific variability between different populations, as specimens fitting both descriptions showed a small genetic divergence. These justify considering Ap. haymoziformis as a junior synonym of Al. haymozi. The phylogenetic position of the species further justifies its placement within the genus Scherotheca (Fig. 1)., Published as part of Marchán, Daniel Fernández, Novo, Marta, Domínguez, Jorge, Silva, Alberto Da, Martínez, Alejandro, Gerard, Sylvain, Hedde, Mickaël & Decaëns, Thibaud, 2023, The best is yet to come: six new species within a largebodied earthworm genus (Scherotheca, Lumbricidae) in a densely sampled region (south-western France), pp. 802-820 in Zoological Journal of the Linnean Society 198 (3) on page 816, DOI: 10.1093/zoolinnean/zlad002, http://zenodo.org/record/8147364

Acanthodrilidae Claus, 1880 The Acanthodrilidae is the second most speciose earthworm family – 746 species and 48 subspecies in 68 genera. Csuzdi (1996) divided it into three subfamilies: Acanthodrilinae, Octochaetinae and Benhamiinae, but the molecular work of James and Davidson (2012) provided evidence that suggested raising the Behamiinae to family level (Benhamiidae). Hence, we consider Benhamiidae as a separate family here. Diplocardinae Michaelsen 1899 has been proposed, but differently defined. The African, Central and South American, and Austral-Asian octochaetine worms surely belong to Acanthodrilinae (Buckley et al. 2011; James & Davidson 2012). Various other subfamilies were proposed by Blakemore (2013), but to date there has been limited acceptance of these subfamilies due to unresolved taxonomic issues and disagreement, and pending further molecular work within the Acanthodrilidae (see Fragoso & Rojas 2016, 2019). The endemic taxa of Acanthodrilinae can be found throughout the southern continents (South America, South Africa, Madagascar, Australia, New Zealand, New Caledonia and the Antarctic Islands), as well as in the USA, Mexico and the Caribbean Islands (Fragoso & Rojas 2016, 2019; Buckley et al. 2011). However, they are completely missing from Europe and the Northern part of Asia (the Palearctic Region). This widely distributed Gondwanan subfamily includes 41 genera and ca. 580 species (updated from Fragoso & Rojas 2016). From continental Central America and the Caribbean Islands and Mexico, we know 14 genera (Balanteodrilus, Borgesia, Diplocardia, Diplotrema, Exxus, Kaxdrilus, Larsonidrilus, Lavellodrilus, Mayadrilus, Protozapotecia, Neotrigaster, Trigaster, Zapatadrilus, Zapotecia) and around 120 species (Fragoso & Rojas 2016, 2018, 2019; Cervantes et al. 2016). Further endemic genera are found in Eastern African countries and in Madagascar (Howascolex and Vazimbascolex, with four and one species, respectively; Csuzdi et al. 2016; Hong et al. 2019). In USA and Mexico there are over 50 species of Diplocardia, and in Mexico, the following additional native genera can be found: Balanteodrilus (3 species), Diplotrema (6 species), Kaxdrilus (3 species), Larsonidrilus (2 species), Lavellodrilus (3 species), Mayadrilus (1 species), Microscolex (3 species), Protozapotecia (4 species), and Zapotecia (2 species) (Reynolds 2020; Fragoso & Rojas 2016, updated with new species). In Central America and the Caribbean, there are the following native genera: Balanteodrilus (1 sp.), Diplotrema (4 species), Kaxdrilus (4 species), Parachilota (2 species), Protozapotecia (3 species) and Zapotecia (1 sp.), while in South America, we find mostly Chilota (26 species), Diplotrema (9 species), Microscolex (10 species) and Yagansia (21 species) in the Andean and Pampean regions of Chile and Argentina (Fragoso & Rojas 2016, 2018; Cervantes & Fragoso 2018). Exxus is of unknown provenance, but suspected to be Caribbean or Central American. In Africa, the Acanthodrilinae are distributed mainly in South Africa comprising 5 genera (Chilota, Eodriloides, Microscolex, Parachilota and Udeina) and ca. 110 species (Plisko & Nxele 2015). In Australia, native acanthodrilinae include six native genera (Diplotrema, Kayarmacia, Microscolex, Neodiplotrema, Rhododrilu s and Torresiella) and approximately 120 species (expanded from Fragoso & Rojas 2016). However, the genus Diplotrema is in need of further work (particularly molecular) within the Acanthodrilinae, which will probably end up splitting off the Neotropical species present in Mexico from those in Oceania (Fragoso & Rojas 2019). At least two Acanthodrilinae species – Microscolex dubius (Fletcher), and M. phosphoreus (Dugés), probably originally from Southern South America (Argentina) – are cosmopolitan, although they tend to be more common in subtropical and temperate climate regions (Rota et al. 2018b). These species were described from specimens collected outside their native range, and have been widely reported from North American and European countries (e.g., Portugal, Spain, France, Italy, Hungary, Serbia, Bulgaria, Albania and Greece), Australia, New Zealand and South Africa. The species M. phosphoreus is a well-known widespread bioluminescent worm, but recent work has shown the need for more detailed analysis of its taxonomy (both morphology and genetics) and questioned the identity of samples deposited throughout the world, due to the presence of cryptic morphs that may be different species (Rota et al. 2018b). However, this high morphological plasticity and deep genetic differentiation can also be due to its well-known parthenogenetic reproduction mode (as in many peregrine species) (Gates 1972)., Published as part of Misirlioğlu, Mete, Reynolds, John Warren, Stojanović, Mirjana, Trakić, Tanja, Sekulić, Jovana, James, Samuel W., Csuzdi, Csaba, Decaëns, Thibaud, Lapied, Emmanuel, Phillips, Helen R. P., Cameron, Erin K. & Brown, George G., 2023, Earthworms (Clitellata, Megadrili) of the world: an updated checklist of valid species and families, with notes on their distribution, pp. 417-438 in Zootaxa 5255 (1) on pages 419-420, DOI: 10.11646/zootaxa.5255.1.33, http://zenodo.org/record/7745110, {"references":["Csuzdi, Cs. (1996) Revision der Unterfamilie Benhamiinae Michaelsen, 1897 (Oligochaeta: Acanthodrilidae). Mitteilungen aus dem Zoologischen Museum in Berlin, 72, 347 - 367. https: // doi. org / 10.1002 / mmnz. 19960720219","James, S. W. & Davidson, S. K. (2012) Molecular phylogeny of earthworms (Annelida: Crassiclitellata) based on 28 S, 18 S and 16 S gene sequences. Invertebrate Systematics, 26 (2), 213 - 229. https: // doi. org / 10.1071 / IS 11012","Buckley, T. R., James, S., Allwood, J., Bartlam, S., Howitt, R. & Prada, D. (2011) Phylogenetic analysis of New Zealand earthworms (Oligochaeta: Megascolecidae) reveals ancient clades and cryptic taxonomic diversity. Molecular Phylogenetics and Evolution, 58, 85 - 96. https: // doi. org / 10.1016 / j. ympev. 2010.09.024","Blakemore, R. J. (2013) The major megadrile families of the world reviewed again on their taxonomic types (Annelida: Oligochaeta: Megadrilacea). Opuscula Zoologica, Budapest, 44 (2), 107 - 127.","Fragoso, C. & Rojas, P. (2016) Lavellodrilus notosetosus sp. nov. (Annelida, Crassiclitellata, Acanthodrilidae): a new Mexican earthworm with uncommon characters, revealed by a preliminary revision of subfamily Acanthodrilinae. Zootaxa, 4154, 101 - 138. https: // doi. org / 10.11646 / zootaxa. 4154.2.1","Fragoso, C. & Rojas, P. (2019) More new Diplotrema earthworm species from southern Mexico (Annelida, Crassiclitellata, Acanthodrilidae, Acanthodrilinae). Zootaxa, 4688 (4), 483 - 502. https: // doi. org / 10.11646 / zootaxa. 4688.4.2","Cervantes, G., Fragoso, F., Monteros, A. E., Sanchez-Ramos, G., Lara-Villalon, M., Yanez-Pacheco, M. J., Lazaro-Castellanos, J. O. & James, S. W. (2016) New species of the earthworm genus Zapatadrilus (Clitellata, Acanthodrilidae) from northern Mexico. Zootaxa, 4189 (2), 311 - 326. https: // doi. org / 10.11646 / zootaxa. 4189.2.7","Csuzdi, Cs., Razafindrakoto, M. & Hong, Y. (2016) The second species of the endemic Malagasy earthworm genus Howascolex Michaelsen, 1901; Howascolex farafangana sp. n. (Clitellata, Megadrili). African Invertebrates, 57 (2), 83 - 91. https: // doi. org / 10.3897 / AfrInvertebr. 57.10048","Hong, Y., Razafindrakoto, M., Blanchart, E., Szederjesi, T. & Csuzdi, Cs. (2019) New acanthodrilid species from Madagascar (Clitellata, Acanthodrilidae). Acta Zoologica Academiae Scientiarum Hungaricae, 65 (3), 215 - 233. https: // doi. org / 10.17109 / AZH. 65.3.215.2019","Reynolds, J. W. (2020) Earthworms in American ecoregions. LAP LAMBERT Academic Publishing, Mauritius. 433 pp.","Cervantes, G. & Fragoso, C. (2018) Protozapotecia acaxetlensis and Protozapotecia oyametlensis, two new earthworm species (Crassiclitellata: Acanthodrilidae) from Mexican central mountains. Zootaxa, 4496, 431 - 439. https: // doi. org / 10.11646 / zootaxa. 4496.1.32","Brown, G. G., James, S. W., Csuzdi, C., Lapied, E., Decaens, T., Reynolds, J. W., Misirlioglu, M., Stovanic, M., Trakic, T., Sekulic, J., Phillips, H. & Cameron, E. (2023) A checklist of megadrile earthworm (Annelida: Clitellata) species and subspecies of the world. Available from: Zenodo. https: // doi. org / 10.5281 / zenodo. 7301848","Plisko, J. D. & Nxele, T. C. (2015) An annotated key separating foreign earthworm species from the indigenous South African taxa (Oligochaeta: Acanthodrilidae, Eudrilidae, Glossoscolecidae, Lumbricidae, Megascolecidae, Microchaetidae, Ocnerodrilidae and Tritogeniidae). African Invertebrates, 56 (3), 663 - 708. https: // doi. org / 10.5733 / afin. 056.0312","Rota, E., Martinsson, S., Erseus, C., Petushkov, V. N., Rodionova, N. S. & Omodeo, P. (2018 b) Green light to an integrative view of Microscolex phosphoreus (Duges, 1837) (Annelida: Clitellata: Acanthodrilidae). Zootaxa, 4496 (1), 175 - 189. https: // doi. org / 10.11646 / zootaxa. 4496.1.13","Gates, GE. (1972) Burmese earthworms. An introduction to the systematics and biology of Megadrile Oligochaetes with special reference to Southeast Asia. Transactions of the American Philosophical Society, 62, 1 - 326. https: // doi. org / 10.2307 / 1006214"]}

Arecoidae James, Csuzdi & Brown, 2023 Fam. nov. Type genus: Areco Righi, Ayres & Bittencourt, 1978 Diagnosis: Crassiclitellata with quadrangular body, and dorsal post-clitellar groove. Four pairs of closelypaired, regularly distributed hook-shaped setae. Dorsal pores present. Genital pores microscopic. Long muscular esophagus, but no gizzard. No calciferous glands, no prostates or copulatory pouches. Intestine begins in XXXVII, typhlosole present. Pretesticular spermathecae, ovaries in XIII. Moniliform hearts in segments VIII to XI. No subneural vessel. Nephridia holoic, without bladders. Metandric, with one pair of testicles and seminal funnels in segment XI. Seminal vesicles in XII to XV. Etymology: This new family name derives from the genus name Areco which with the species name reco is derived from “arecoreco”, originally in the Tupi indigenous language, meaning to confuse or to mix something up (Drumond 1952). Considerations: The intriguing aqua-megadrile (sensu Jamieson 1988) species Areco reco was described by Righi et al. (1978) from four specimens collected at the Reserva Ducke near Manaus, Amazonas (Brazil) in 1975. Type specimens are deposited in the Instituto Nacional de Pesquisas da Amazonia (INPA) collection in Manaus. The authors placed the species tentatively in Sparganophilidae, but emphasized that the exact position of Areco was uncertain. They claimed similarity with Glyphydrilocrius by shape of body and organization of digestive system, and with Sparganophilus by presence of pretesticular spermathecae, though this is a feature shared with several other South American families (e.g., Glossoscolecidae, Rhinodrilidae). Furthermore, Sparganophilus and Glyphydrilocrius are holandric, while Areco is metandric. Additional Areco specimens were collected at the type locality by S. James and S. Tapia-Coral in 2012 (Erséus et al. 2020), and molecular phylogenomic analyses revealed it to be sister to Rhinodrilidae, and far from Sparganophilidae and other semi-aquatic families (Erséus et al. 2020). Nonetheless, Arecoidae shows important genetic divergence and major morphological differences with Rhinodrilidae, lacking gizzard and calciferous glands, so we propose here the erection of this new monotypic family to accommodate its only species and genus.

Biwadrilidae Jamieson, 1971 This monogeneric family includes only one species, Biwadrilus bathybates (Stephenson), known only from the Lake Biwa region in Japan. It was originally described as a Criodrilus species, and Blakemore (2008b) transferred it back to Criodrilidae based on new material and re-analysis of previously observed material. However, molecular analysis shows this family solidly placed as the sister family to Kynotidae, and therefore quite distant phylogenetically from the Criodrilidae (James & Davidson 2012), hence we maintain it within Biwadrilidae, following Jamieson (1971, 1988)., Published as part of Misirlioğlu, Mete, Reynolds, John Warren, Stojanović, Mirjana, Trakić, Tanja, Sekulić, Jovana, James, Samuel W., Csuzdi, Csaba, Decaëns, Thibaud, Lapied, Emmanuel, Phillips, Helen R. P., Cameron, Erin K. & Brown, George G., 2023, Earthworms (Clitellata, Megadrili) of the world: an updated checklist of valid species and families, with notes on their distribution, pp. 417-438 in Zootaxa 5255 (1) on page 423, DOI: 10.11646/zootaxa.5255.1.33, http://zenodo.org/record/7745110, {"references":["Jamieson, B. G. M. (1971) Glossoscolecidae. In: Brinkhurst, R. O. & Jamieson, B. G. M. (Eds.), Aquatic Oligochaeta of the World. Oliver and Boyd, Edinburgh, pp. 723 - 837.","Blakemore, R. (2008 b) Review of Criodrilidae (Annelida: Oligochaeta) including Biwadrilus from Japan. Opuscula Zoologica Budapest, 37, 11 - 22.","James, S. W. & Davidson, S. K. (2012) Molecular phylogeny of earthworms (Annelida: Crassiclitellata) based on 28 S, 18 S and 16 S gene sequences. Invertebrate Systematics, 26 (2), 213 - 229. https: // doi. org / 10.1071 / IS 11012","Jamieson, B. G. M. (1988) On the phylogeny and higher classification of the Oligochaeta. Cladistics, 4, 367 - 401. https: // doi. org / 10.1111 / j. 1096 - 0031.1988. tb 00520. x"]}

Moniligastridae Claus, 1880 Although moniligastrids are not considered part of the Crassiclitellata since their clitella have only a single layer like the Enchytraeidae, they tend to have much larger individual body size and mass than enchytraeids and are phylogenetically sister to the Crassiclitellata (James & Davidson 2012; Schmelz et al. 2021). The family is mainly distributed in the Oriental region from south India east to the Philippines and north to southern Siberia and Japan (Gates 1972; Perel 1997). It includes 185 species (and 19 subspecies), distributed among five genera (Desmogaster, Drawida, Eupolygaster, Hastirogaster, Moniligaster), of which Drawida is the most widely distributed and speciose (162 spp./sspp.). Three species, Drawida barwelli (Beddard), Drawida japonica (Michaelsen), and Drawida nepalensis Michaelsen, are cosmopolitan, but of these, D. barwelli is the most widely distributed. It is also sometimes mentioned under its synonymous names D. bahamensis (Beddard), or D. beddardi (Rosa), being reported from Africa (Csuzdi 2005), Australia, South-East Asia, Mexico, Central America, the Caribbean, and several Pacific Islands (Blakemore et al. 2014; Csuzdi et al. 2017)., Published as part of Misirlioğlu, Mete, Reynolds, John Warren, Stojanović, Mirjana, Trakić, Tanja, Sekulić, Jovana, James, Samuel W., Csuzdi, Csaba, Decaëns, Thibaud, Lapied, Emmanuel, Phillips, Helen R. P., Cameron, Erin K. & Brown, George G., 2023, Earthworms (Clitellata, Megadrili) of the world: an updated checklist of valid species and families, with notes on their distribution, pp. 417-438 in Zootaxa 5255 (1) on page 427, DOI: 10.11646/zootaxa.5255.1.33, http://zenodo.org/record/7745110, {"references":["James, S. W. & Davidson, S. K. (2012) Molecular phylogeny of earthworms (Annelida: Crassiclitellata) based on 28 S, 18 S and 16 S gene sequences. Invertebrate Systematics, 26 (2), 213 - 229. https: // doi. org / 10.1071 / IS 11012","Schmelz, R., Erseus, C., Martin, P., Van Haaren, T. & Timm, T. (2021) A proposed order-level classification in Oligochaeta (Annelida, Clitellata). Zootaxa, 5040, 589 - 597. https: // doi. org / 10.11646 / zootaxa. 5040.4.9","Gates, GE. (1972) Burmese earthworms. An introduction to the systematics and biology of Megadrile Oligochaetes with special reference to Southeast Asia. Transactions of the American Philosophical Society, 62, 1 - 326. https: // doi. org / 10.2307 / 1006214","Perel, T. S. (1997) The earthworms of the fauna of Russia: cadaster and key. Academy of Sciences of the USSR. Pub. Nauka, Moscow, 101 pp. [in Russian]","Csuzdi, Cs. (2005) Earthworms (Annelida: Oligochaeta) of Sao Tome. Journal of Natural History, 39 (33), 3039 - 3058. https: // doi. org / 10.1080 / 00222930500198403","Blakemore, R. J., Lee, S. & Seo, H. - Y. (2014) Reports of Drawida (Oligochaeta: Moniligastridae) from far East Asia. Journal of Species Research, 3 (2), 127 - 166. https: // doi. org / 10.12651 / JSR. 2014.3.2.127","Csuzdi, Cs., Pearlson, O. & Pavlicek, T. (2017) New Acanthodrilus species from New Caledonia (Clitellata, Megadrili, Acanthodrilidae). Journal of Natural History, 51, 1899 - 1912. https: // doi. org / 10.1080 / 00222933.2017.1355500"]}

Sparganophilidae Michaelsen, 1918 This family native to North America has only one genus (Sparganophilus), with 12 species and 2 subspecies (Reynolds 1980, 2008). At least one species, Sparganophilus tamesis Benham (Sparganophilus eiseni Smith, is a junior synonym of S. tamesis; see Rota et al. 2016), is cosmopolitan, and found mainly in limicole habitats, throughout the USA (within and outside of its native range), Mexico (Ikeda et al. 2020), Canada (Reynolds 2022) and in several European countries including Germany, France, England, Switzerland, Italy and Spain (Rota et al. 2018a; Bouché & Qiu 1998)., Published as part of Misirlioğlu, Mete, Reynolds, John Warren, Stojanović, Mirjana, Trakić, Tanja, Sekulić, Jovana, James, Samuel W., Csuzdi, Csaba, Decaëns, Thibaud, Lapied, Emmanuel, Phillips, Helen R. P., Cameron, Erin K. & Brown, George G., 2023, Earthworms (Clitellata, Megadrili) of the world: an updated checklist of valid species and families, with notes on their distribution, pp. 417-438 in Zootaxa 5255 (1) on page 428, DOI: 10.11646/zootaxa.5255.1.33, http://zenodo.org/record/7745110, {"references":["Reynolds, J. W. (1980) The earthworm family Sparganophilidae (Annelida, Oligochaeta) in North America. Megadrilogica, 3 (12), 189 - 204.","Reynolds, J. W. (2008) Sparganophilidae (Annelida, Oligochaeta) distribution in North America and elsewhere, revisited. Megadrilogica, 12 (9), 125 - 143.","Rota, E., Martinsson, S., Bartoli, M., Beylich, A., Graefe, U., Laini, A., Wetzel, M. J. & Erseus, C. (2016) Mitochondrial evidence supports a Nearctic origin for the spreading limicolous earthworm Sparganophilus tamesis Benham, 1892 (Clitellata, Sparganophilidae). Contributions to Zoology, 85 (1), 113 - 119. https: // doi. org / 10.1163 / 18759866 - 08501005","Ikeda, H., Callaham Jr., M. A., Shefferson, R. P., Wenk, E. S. & Fragoso, C. (2020) A comparison of latitudinal species diversity patterns between riverine and terrestrial earthworms from the North American temperate zone. Journal of Biogeography, 47, 1373 - 1382. https: // doi. org / 10.1111 / jbi. 13826","Rota, E., Benelli, S., Erseus, C., Soors, J. & Bartoli, M. (2018 a) New data and hypotheses on the invasiveness, habitat selection, and ecological role of the limicolous earthworm Sparganophilus tamesis Benham, 1892. Fundamental and Applied Limnology, 192 / 2, 129 - 136. https: // doi. org / 10.1127 / fal / 2018 / 1147","Bouche, M. B. & Qiu, J. - P. (1998) Un nouveau Sparganophilus (Annelida: Oligochaeta) d'Europe, avec considerations paleogeographiques sur les Lumbricina. Documents Pedozoologiques et Integrologiques, 4, 178 - 180."]}

Benhamiidae Michaelsen, 1897 Native species of the Benhamiidae (351 species and 6 subspecies in 20 genera) are mainly restricted to subSaharan Africa, Oceania, Central America and northern South America (Csuzdi 2010). However, a few Dichogaster (including the type species of the genus) are also known from the Pacific islands of Fiji and Samoa (Easton 1984; James & Davidson 2012). In Africa, the native Benhamiidae are present mainly in the tropical region with 16 genera (Afrogaster, Agastrodrilus, Benhamia, Benhamiona, Dichogaster, Dudichiodrilus, Guineoscolex, Loksaia, Millsonia, Monogaster, Monothecodrilus, Omodeona, Pickfordia, Pickfordiella, Reginaldia and Wegeneriella) and ca. 250 species. From the Neotropics, i.e., the Caribbean islands, Central America, Mexico, and Northern South America we know five genera (Dichogaster, Eutrigaster, Neogaster, Omodeoscolex, and Wegeneriona) and around 100 species, mainly of Dichogaster and Eutrigaster, many of which show high endemism rates, particularly on the Caribbean islands. The diversity of the minute South-American species is certainly underestimated due to a lack of sampling, especially in Amazonia, where recent work (in French Guyana), revealed several new species, that are presently being described. Several species in the family Benhamiidae, particularly in the genus Dichogaster, are widespread anthropochores in the tropics, as their native ranges were probably originally in Central and Eastern Africa (Gates 1972; Csuzdi 2010). These species were probably widely dispersed during the centuries of commercial product exchanges between Africa and other continents, where ships could have transported soil in planted pots containing these generally parthenogenetic species. More recent transport with manure piles or agricultural equipment may also have led to the abundance of Dichogaster gracilis (Michaelsen), Dichogaster bolaui (Michaelsen), Dichogaster saliens (Beddard), and Dichogaster affinis (Michaelsen) in many no-tillage sites in Southern and Central Brazil, where they may reach abundances well over 100 individuals m-2 (Bartz et al. 2009; Bartz et al. 2014; Santos et al. 2018). Although Dichogaster spp. are generally restricted to warm and humid tropical regions, several species have been reported from greenhouses in temperate climates and at least one species (D. bolaui) was considered a domicole (adapted to human homes) species, as it was found in bathtubs, showers and sewage pipes in Hungary, Ireland, Finland, Sweden and Israel (Terhivuo 1991; Erséus et al. 1994; Rota & Schmidt 2006; Csuzdi et al. 2008). In fact, the type locality of D. bolaui is Hamburg, where it was found in fermenting bark at a factory (Michaelsen 1891). Dichogaster annae (Horst) is an epigeic species, common in vermicomposting ventures in tropical countries such as Brazil (James & Guimar„es 2010), but its distribution is probably much wider than presently known or reported, due to the lack of recognition by worm composters, and the little knowledge of its life-cycle and biology. Dichogaster modiglianii (Rosa) is an epi-endogeic species also with wide distribution (Africa, Southeast Asia, Australia, Oceania, Central America, Caribbean, Southern North America (Blakemore 2010), and was recently found in several Amazonian sites, associated with Pre-Columbian human activities (Conrado 2018). Specimens of D. bolaui were also recently reported from a remote and human-free Amazonian rainforest area in Southern French Guyana, highlighting the potential of these cosmopolitan species to establish stable populations that persist centuries after original introduction by humans (Maggia et al. 2021)., Published as part of Misirlioğlu, Mete, Reynolds, John Warren, Stojanović, Mirjana, Trakić, Tanja, Sekulić, Jovana, James, Samuel W., Csuzdi, Csaba, Decaëns, Thibaud, Lapied, Emmanuel, Phillips, Helen R. P., Cameron, Erin K. & Brown, George G., 2023, Earthworms (Clitellata, Megadrili) of the world: an updated checklist of valid species and families, with notes on their distribution, pp. 417-438 in Zootaxa 5255 (1) on page 422, DOI: 10.11646/zootaxa.5255.1.33, http://zenodo.org/record/7745110, {"references":["Easton, E. G. (1984) Earthworms (Oligochaeta) from islands of the south-western Pacific, and a note on two species from Papua New Guinea. New Zealand Journal of Zoology, 11: 111 - 128. https: // doi. org / 10.1080 / 03014223.1984.10423750","James, S. W. & Davidson, S. K. (2012) Molecular phylogeny of earthworms (Annelida: Crassiclitellata) based on 28 S, 18 S and 16 S gene sequences. Invertebrate Systematics, 26 (2), 213 - 229. https: // doi. org / 10.1071 / IS 11012","Gates, GE. (1972) Burmese earthworms. An introduction to the systematics and biology of Megadrile Oligochaetes with special reference to Southeast Asia. Transactions of the American Philosophical Society, 62, 1 - 326. https: // doi. org / 10.2307 / 1006214","Bartz, M. L. C., Brown, G. G., Pasini, A., de Oliveira Fernandes, J., Curmi, P., Dorioz, J. & Ralisch, R. (2009) Earthworm communities in organic and conventional coffee cultivation. Pesquisa Agropecuaria Brasileira, 44, 928 - 933. https: // doi. org / 10.1590 / S 0100 - 204 X 2009000800019","Bartz, M. L. C., Brown, G. G., da Rosa, M. G., Filho, O. K., James, S. W., Decaens, T. & Baretta, D. (2014) Earthworm richness in land-use systems in Santa Catarina, Brazil. Applied Soil Ecology, 83, 59 - 70. https: // doi. org / 10.1016 / j. apsoil. 2014.03.003","Santos, A., Gorte, T., Demetrio, W. C., Ferreira, T., Nadolny, H., Cardoso, G. B. X., Tonetti, C., Ralisch, R., Nunes, A. P., Coqueiro, A. C. P., Leandro, H. C. L., Wandscheer, C. A. R., Bortoluzzi, J., Brown G. G. & Bartz, M. L. C. (2018) Earthworm species in no-tillage agroecosystems and native Atlantic forests in Western Parana, Brazil. Zootaxa, 4496, 517 - 534. https: // doi. org / 10.11646 / zootaxa. 4496.1.40","Terhivuo, J. (1991) Dichogaster bolaui (Michaelsen) (Octochaetidae) - an allochthonous oligochaete invading urban sewer system. Memoranda Societatis pro Fauna et Flora Fennica, 67, 61 - 65.","Erseus, C., Omodeo, P. & Rota, E. (1994) The first records of the allochthonous species Dichogaster saliens and D. bolaui (Oligochaeta: Octochaetidae) from Sweden. Megadrilogica, 6 (2), 17 - 20.","Rota, E. & Schmidt, O. (2006) Dichogaster bolaui (Oligochaeta: Octochaetidae), anunusual invader in a swimming pool in Ireland. Journal of Natural History, 40 (3 - 4), 161 - 167. https: // doi. org / 10.1080 / 00222930600630875","Csuzdi Cs., Pavlicek, T. & Nevo, E. (2008) Is Dichogaster bolaui (Michaelsen, 1891) the first domicole earthworm species? European Journal of Soil Biology, 44, 198 - 201. https: // doi. org / 10.1016 / j. ejsobi. 2007.05.003","Michaelsen, W. (1891) Oligochaeten des Naturhistorishchen Museums in Hamburg. IV. Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten, 8, 3 - 42.","Blakemore, R. J. (2010) Cosmopolitan iarthworms - an Eco-taxonomic guide to the peregrine species of the world. 4 th ed. VermEcology, Yokohama.","Conrado, A. C. (2018) Diversidade da macrofauna edafica em sistemas florestais e agricolas amazonicos. Master's Thesis, Federal University of Parana, Curitiba, 119 pp.","Maggia, M. E., Decaens T., Lapied E., Dupont L., Roy V., Schimann H., Orivel J., Murienne J., Baraloto C., Cottenie K. & Steink D. (2021) At each site its diversity: DNA barcoding reveals remarkable earthworm diversity in tropical rainforest of French Guiana. Applied Soil Ecology, 164, 103932. https: // doi. org / 10.1016 / j. apsoil. 2021.103932"]}

Glossoscolecidae Michaelsen, 1900 In its more restricted sense, after the splitting up of the Rhinodrilidae by James (2012), the Glossoscolecidae family is distributed mainly in continental Central and South America from Panama to Northern Argentina and Uruguay. The first species described from Latin America belongs to this family, Glossoscolex giganteus Leuckart, a large earthworm from Southeastern Brazil. In fact, the genus contains a relatively large proportion of large-bodied (> 30 cm long) species (18 out of 58 known species; Feijoo & Brown 2023). So far, native species are known in the Caribbean Islands only from Guadeloupe (James & Gamiette 2016), but they may likely occur on other islands. The family includes six genera (Enantiodrilus, Fimoscolex, Glossodrilus, Glossoscolex, Holoscolex, and Righiodrilus) and 156 species (plus 9 subspecies). One widespread species, Enantiodrilus borellii Cognetti (with which both Diaguita species, D. vivianae Righi and D. michaelseni Cordero were synonymized by Moreno et al. 2005) may constitute a potentially invasive species in the tropics, considering its distribution from Argentina to Venezuela (Fragoso & Brown 2007). As the family occurs in a region with high endemicity (Lavelle & Lapied 2003), there is a large potential to find many new species and possibly genera, with further sampling efforts, particularly in the Andean region and Northern South America, but also in the Atlantic Forest region of Southeastern Brazil (Silva et al. 2017)., Published as part of Misirlioğlu, Mete, Reynolds, John Warren, Stojanović, Mirjana, Trakić, Tanja, Sekulić, Jovana, James, Samuel W., Csuzdi, Csaba, Decaëns, Thibaud, Lapied, Emmanuel, Phillips, Helen R. P., Cameron, Erin K. & Brown, George G., 2023, Earthworms (Clitellata, Megadrili) of the world: an updated checklist of valid species and families, with notes on their distribution, pp. 417-438 in Zootaxa 5255 (1) on page 424, DOI: 10.11646/zootaxa.5255.1.33, http://zenodo.org/record/7745110, {"references":["Feijoo, A. & Brown, G. G. (2023) Three new Glossoscolex (Annelida: Crassiclitellata: Glossoscolecidae) in the truncatus group from the Brazilian Atlantic Forest. Zootaxa, 5225 (1), 220 - 234. https: // doi. org / 10.11646 / zootaxa. 5225.1.21","James, S. W. & Gamiette, F. (2016) New species of Dichogaster Beddard, 1888 (Clitellata: Benhamiidae) with additional records of earthworms from Guadeloupe (French West Indies). Zootaxa, 4178 (3), 391 - 408. https: // doi. org / 10.11646 / zootaxa. 4178.3.5","Moreno, A. G., Teisaire, E. S. & Roldan, I. A. (2005) Genus Enantiodrilus Cognetti, 1902 (Annelida, Oligochaeta: Glossoscolecidae). In: Pop, V. & Pop, A. (Eds.), Advances in earthworm taxonomy II (Annelida Oligochaeta). Cluj University Press, Cluj- Napoca, pp. 41 - 52.","Fragoso, C. & Brown, G. G. (2007) Ecologia y taxonomia de las lombrices de tierra en latinoamerica: El Primer Encuentro Latino- Americano de Ecologia y Taxonomia de Oligoquetos (ELAETAO 1). In: Brown, G. G. & Fragoso, C. (Eds.), Minhocas na America Latina: Biodiversidade e ecologia. Embrapa Soja, Londrina, pp. 33 - 75.","Lavelle, P. & Lapied, E. (2003) Endangered earthworms of Amazonia: an homage to Gilberto Righi. Pedobiologia, 47, 419 - 427. https: // doi. org / 10.1078 / 0031 - 4056 - 00207"]}

Rhinodrilidae Benham, 1890 Recent changes in classification (James & Davidson 2012) moved many genera from Glossoscolecidae to Rhinodrilidae (James 2012). This family now includes 38 genera and some 376 species (plus seven subspecies) including one widespread invasive species, Pontoscolex corethrurus (M̧ller), probably the most widely distributed earthworm in the world (James 2011; Taheri et al. 2018b). Native species of this family occur from Argentina (Mischis 2007) up to Mexico (Fragoso & Rojas 2014), including several on Caribbean Islands (Rodríguez et al. 2007), although none are known from Chile (Zicsi & Csuzdi 2007). Studies on the genetic diversity of several Pontoscolex species highlighted the importance of proper identification of P. corethrurus, requiring at least barcoding, to certify species status (Taheri et al. 2018a). Species belonging to P. corethrurus must conform to the genetic lineage L1, as identified in Taheri et al. (2018a) and James et al. (2019). The genus Martiodrilus with 88 known species and subspecies is the most speciose, occurring from Panama in the north to Peru in the south, and to French Guyana in the east (Fragoso & Brown 2007), with a large number of species still to be found with additional sampling efforts in Amazonia and the Andes region. The following other speciose Rhinodrilidae genera by decreasing order of species richness are: Rhinodrilus (52 spp./sspp.), Andiorrhinus (48 spp./sspp.) and Andiodrilus (40 spp./sspp.). Together with Rhinodrilus, Martiodrilus is also remarkable for the presence of several very large-bodied earthworm species (Brown & James 2007; Zicsi 2007). One other widespread species of Rhinodrilidae is worth mentioning here: Urobenus brasiliensis Benham, found in Paraguay (Brown & Fragoso, 2007) and in Brazil from Manaus in the Amazon (R̂mbke et al. 1999; Zicsi et al. 2001) to Pelotas in the Pampa biome (Santos et al. 2019). This cryptic species requires further morphological and molecular work in order to separate the many genetic lineages that have little morphological differentiation (da Silva et al. 2017). Furthermore, the genus Alexidrilus Righi, was recently synonymized with Urobenus Benham, and its two species, A. lourdesae Righi and A. littoralis Ljungstr ̂m, with U. brasiliensis (Ferreira et al. 2023)., Published as part of Misirlioğlu, Mete, Reynolds, John Warren, Stojanović, Mirjana, Trakić, Tanja, Sekulić, Jovana, James, Samuel W., Csuzdi, Csaba, Decaëns, Thibaud, Lapied, Emmanuel, Phillips, Helen R. P., Cameron, Erin K. & Brown, George G., 2023, Earthworms (Clitellata, Megadrili) of the world: an updated checklist of valid species and families, with notes on their distribution, pp. 417-438 in Zootaxa 5255 (1) on page 428, DOI: 10.11646/zootaxa.5255.1.33, http://zenodo.org/record/7745110, {"references":["James, S. W. & Davidson, S. K. (2012) Molecular phylogeny of earthworms (Annelida: Crassiclitellata) based on 28 S, 18 S and 16 S gene sequences. Invertebrate Systematics, 26 (2), 213 - 229. https: // doi. org / 10.1071 / IS 11012","James, S. W. (2011) Earthworms. In: Simberlof, D. & Rejmanek, M. (Ids.), Encyclopaedia of biological invasions. University of California Press, Berkeley, pp. 177 - 183. https: // doi. org / 10.1525 / 9780520948433 - 045","Taheri, S., Pelosi, C. & Dupont, L. (2018 b) Harmful or useful? A case study of the exotic peregrine earthworm morphospecies Pontoscolex corethrurus. Soil Biology and Biochemistry, 116, 277 - 289. https: // doi. org / 10.1016 / j. soilbio. 2017.10.030","Mischis, C. (2007) Catalogo de las lombrices de tierra de la Argentina (Annelida, Oligochaeta). In: Brown, G. G. & Fragoso, C. (Eds.), Minhocas na America Latina: Biodiversidade e Ecologia, Embrapa, Londrina, pp. 247 - 252.","Fragoso, C. & Rojas, P. (2014) Biodiversidad de lombrices de tierra (Annelida: Oligochaeta: Crassiclitellata) en Mexico. Revista Mexicana de Biodiversidad, Suplemento 85, S 197 - S 207. https: // doi. org / 10.7550 / rmb. 33581","Rodriguez, C., Borges, S., Martinez, M. A., Fragoso, C., James, S. W. & Gonzalez, G. (2007) Biodiversidad y ecologia de las lombrices de tierra em las islas caribenas. In: Brown, G. G. & Fragoso, C. (Eds.), Minhocas na America Latina: Biodiversidade e Ecologia, Embrapa, Londrina, pp. 79 - 98.","Zicsi, A. & Csuzdi, C. (2007) An annotated checklist of the earthworms of Chile (Oligochaeta). Earthworms from South America 43. In: Brown, G. G. & Fragoso, C. (Eds.), Minhocas na America Latina: Biodiversidade e Ecologia, Embrapa, Londrina, pp. 235 - 245.","Taheri S., James, S. W., Roy, V., Decaens, T., Williams, B. W., Anderson, F., Rougerie, R., Chang, C. - H., Brown, G. G., Cunha, L., Stanton, D. W. G., Da Silva, E., Chen, J-H., Lemmon, A. R., Moriarty Lemmon, E., Bartz, M., Baretta, D., Barois, I., Lapied, E., Coulis, M. & Dupont, L. (2018 a) Complex taxonomy of the ' brush tail' peregrine earthworm Pontoscolex corethrurus. Molecular Phylogenetics and Evolution, 124, 68 - 70. https: // doi. org / 10.1016 / j. ympev. 2018.02.021","James, S. W., Bartz, M. L. C., Stanton, D. W. G., Conrado, A. C., Dupont, L., Taheri, S., da Silva, E., Cunha, L. & Brown, G. G. (2019) A neotype for Pontoscolex corethrurus (M ¸ ller, 1857) (Clitellata). Zootaxa, 4545 (1), 124 - 132. https: // doi. org / 10.11646 / zootaxa. 4545.1.7","Fragoso, C. & Brown, G. G. (2007) Ecologia y taxonomia de las lombrices de tierra en latinoamerica: El Primer Encuentro Latino- Americano de Ecologia y Taxonomia de Oligoquetos (ELAETAO 1). In: Brown, G. G. & Fragoso, C. (Eds.), Minhocas na America Latina: Biodiversidade e ecologia. Embrapa Soja, Londrina, pp. 33 - 75.","Brown, G. G. & Fragoso, C. (2007) Listado de las especies de lombrices de tierra de America Central Continental (Guatemala, Belice, Honduras, El Salvador, Nicaragua, Costa Rica, Panama), las Guyanas (Surinam, Guyanne Francaise, Guyana), Venezuela y Paraguay. In: Brown, G. G. & Fragoso, C. (Eds.), Minhocas na America Latina: Biodiversidade e ecologia. Embrapa Soja, Londrina, pp. 422 - 452.","Zicsi, A., Rombke, J. & Garcia, M. (2001) Regenwurmer (Oligochaeta) aus der Umgebung von Manaus (Amazonien). Regenw ¸ rmer aus S ¸ damerika 32. Revue Suisse de Zoologie, 108 (1), 153 - 164. https: // doi. org / 10.5962 / bhl. part. 79624","Santos, A., Maia, L. S., Ferreira, T., Demetrio, W. C., Nadolny, H. S., Ribeiro, L. V., Schiedeck, G., Costa, F. A., Bartz, M. L. C. & Brown, G. G. (2019) Minhocas como biondicadoras da qualidade do solo em ecossistemas na Embrapa Clima Temperado. Embrapa Clima Temperado, Documentos 482, Pelotas, 39 pp.","da Silva, E., Rosa, M. G., Schuhli, G. S., James, S. W., Decaens, T., Bartz, M. L. C., Nadolny, H., Feijoo, A. & Brown, G. G. (2017) O potencial do DNA barcoding para a identificac \" o e conservac \" o de especies de minhocas brasileiras. In: Moreira, F. M. S. & Kasuya, M. C. M. (Eds.), Fertilidade e biologia do solo - Integracao e tecnologia para todos, Vol. 2. Lavras, SBCS, pp. 549 - 570.","Ferreira, T., James, S. W., Bartz, M. L. C., Lima, A. C. R., Dudas, R. & Brown, G. G. (2023) Distribution and diversity of earthworms in different land use systems in Rio Grande do Sul, Brazil. Zootaxa, 5225 (1), 389 - 398. https: // doi. org / 10.11646 / zootaxa. 5225.1.31"]}

Ocnerodrilidae Beddard, 1891 The widespread Ocnerodrilidae family, with 37 genera and 172 species (plus four subspecies), sister taxon to the Megascolecidae + Acanthodrilidae s.l. clade (James & Davidson 2012), was last reviewed by Fragoso & Rojas (2009) and Hernández-García et al. (2018), who provided keys to the genera known until then.Two sub-families (Ocnerodrilinae and Malabariinae) were proposed by Gates (1966) and their definitions expanded by Gates (1972) and Fragoso and Rojas (2009). Ocnerodrilinae is more speciose and wider spread, while Malabariinae includes species only from India, China and Myanmar (Fragoso & Rojas 2009), without extramural glands and internal calciferous lamellae. With the exception of several peregrine species with worldwide distribution, all endemic genera and species of Ocnerodrilinae are confined to South and Central America, sub-Saharan Africa, India (Curgiona), and the Seychelles (Maheina). A large number of new species and several new genera were recently found in Brazil (James et al. 2023; HernándezGarcía et al. 2018), and it is likely that further sampling particularly in Northern South America, and in Brazil’s Northeastern region will reveal many more. The majority (27) of the known genera occur in Latin America, while nine genera are known from Africa and five from Asia (India, China and Myanmar). In Latin America, many of the smaller and pigmented ocnerodriles, particularly in the genera Kerriona and Eukerria inhabit wetlands or bromeliads, and some of them are quite resistant to disturbance, being present in intensively plowed rice or no-tillage row-crop fields in Southern Brazil (Bartz et al. 2013; Bartz et al. 2014; Ferreira et al. 2023; Lima & Rodríguez 2007). There are nine widespread anthropochorous ocnerodriles: Eukerria eiseniana (Rosa), Eukerria kuekenthali (Michaelsen), Eukerria saltensis (Beddard), Eukerria stagnalis (Kinberg), Gordiodrilus elegans Beddard, Gordiodrilus habessinus Michaelsen, Gordiodrilus paski Stephenson, Nematogenia lacuum Beddard, and Ocnerodrilus occidentalis Eisen, that typically live in wet or saturated soils, generally close to water sources. Some of these are of uncertain origin, being possibly of South America or Africa but, in any case, are of Gondwanan or Central Pangean origin (Fragoso & Rojas 2009). The presence of many parthenogenetic morphs in some species (e.g., O. occidentalis), has resulted in extensive synonymies which must be checked (James et al. 2023), although the use of molecular methods may help unravel some of these. The species Nematogenia panamaensis Eisen was synonymized with N. lacuum by Righi (1984a), because variations between the two species were included in a group of specimens collected at the same location in Mato Grosso do Sul, Brazil. Similar to O. occidentalis further work is warranted on these poorly known ocnerodriles, particularly using more detailed genetic methods, in order to confirm the presence of potential cryptic species (Fragoso & Rojas 2009)., Published as part of Misirlioğlu, Mete, Reynolds, John Warren, Stojanović, Mirjana, Trakić, Tanja, Sekulić, Jovana, James, Samuel W., Csuzdi, Csaba, Decaëns, Thibaud, Lapied, Emmanuel, Phillips, Helen R. P., Cameron, Erin K. & Brown, George G., 2023, Earthworms (Clitellata, Megadrili) of the world: an updated checklist of valid species and families, with notes on their distribution, pp. 417-438 in Zootaxa 5255 (1) on page 427, DOI: 10.11646/zootaxa.5255.1.33, http://zenodo.org/record/7745110, {"references":["James, S. W. & Davidson, S. K. (2012) Molecular phylogeny of earthworms (Annelida: Crassiclitellata) based on 28 S, 18 S and 16 S gene sequences. Invertebrate Systematics, 26 (2), 213 - 229. https: // doi. org / 10.1071 / IS 11012","Fragoso, C. & Rojas, P. (2009) A new ocnerodrilid earthworm genus from Southeastern Mexico (Annelida: Oligochaeta), with a key for the genera of Ocnerodrilidae. Megadrilogica, 13 (9), 141 - 152.","Hernandez-Garcia, L. M., Burgos-Guerrero, J. E., Rousseau, G. X. & James, S. W. (2018) Brasilisia n. gen. and Arraia n. gen., two new genera of Ocnerodrilidae (Annelida, Clitellata, Oligochaeta) from Eastern Amazonia, Brazil. Zootaxa, 4496, 471 - 480. https: // doi. org / 10.11646 / zootaxa. 4496.1.36","Gates, G. E. (1966) Contributions to a revision of the earthworm family Ocnerodrilidae. VII-VIII. Annals and Magazine Natural History, 9 (13), 45 - 53. https: // doi. org / 10.1080 / 00222936608651637","Gates, GE. (1972) Burmese earthworms. An introduction to the systematics and biology of Megadrile Oligochaetes with special reference to Southeast Asia. Transactions of the American Philosophical Society, 62, 1 - 326. https: // doi. org / 10.2307 / 1006214","James, S. W., Bartz, M. L. C. & Brown, G. G. (2023) New Ocnerodrilidae genera, species and records from Brazil. Zootaxa, 5255 (1), 235 - 269. https: // doi. org / 10.11646 / zootaxa. 5255.1.22","Bartz, M. L. C., Pasini, A. & Brown, G. G. (2013) Earthworms as soil quality indicators in Brazilian no-tillage systems. Applied Soil Ecology, 69, 39 - 48. https: // doi. org / 10.1016 / j. apsoil. 2013.01.011","Bartz, M. L. C., Brown, G. G., da Rosa, M. G., Filho, O. K., James, S. W., Decaens, T. & Baretta, D. (2014) Earthworm richness in land-use systems in Santa Catarina, Brazil. Applied Soil Ecology, 83, 59 - 70. https: // doi. org / 10.1016 / j. apsoil. 2014.03.003","Ferreira, T., James, S. W., Bartz, M. L. C., Lima, A. C. R., Dudas, R. & Brown, G. G. (2023) Distribution and diversity of earthworms in different land use systems in Rio Grande do Sul, Brazil. Zootaxa, 5225 (1), 389 - 398. https: // doi. org / 10.11646 / zootaxa. 5225.1.31","Lima, A. C. R. & Rodriguez, C. (2007) Earthworm diversity from Rio Grande Do Sul, Brazil, with a new native criodrilid genus and species (Oligochaeta: Criodrilidae). Megadrilogica, 11 (2), 9 - 18.","Righi, G. (1984 a) On a collection of Neotropical Megadrili Oligochaeta I. Ocnerodrilidae, Acanthodrilidae, Octochaetidae, Megascolecidae. Studies on Neotropical Fauna and Environment, 19 (1), 9 - 31. https: // doi. org / 10.1080 / 01650528409360641"]}

Lumbricidae Rafinesque-Schmaltz, 1815 The Lumbricidae is the phylogenetically youngest family in the subclass of Oligochaeta. The family perhaps originated in the Palearctic but there is a well-supported clade native to Eastern North America. Overall, the Lumbricidae family comprises some 615 species and 74 subspecies belonging to 47 genera (Brown et al. 2023), although the lumbricid taxonomy is still controversial and not yet settled (Bouché 1972; Mršić 1991; Qiu & Bouché 1998; Csuzdi & Zicsi 2003; Domínguez et al. 2015; de Sosa et al. 2019; Marchán et al. 2022a; Marchán et al. 2022c). Around 33 cosmopolitan species of the family are widespread and often invasive in numerous regions (Blakemore 2009), such as North America, Australia, New Zealand, South Africa, Southern South America, and even in equatorial countries at higher elevation (e.g., Zicsi 2007). In North America there are both native and non-native species and genera. The native genera are Bimastos and Eisenoides with 15 and two species, respectively. There are 11 genera of non-native invasive lumbricids in North America (Fragoso & Rojas 2014; Reynolds 2020, 2022), and 10 genera of invasives in South America (Fragoso & Brown 2007). In Europe, centers of diversity of lumbricids are situated in the Pyrenees and the Balkan Peninsula, as well as in the Carpathians and the Caucasus (Kvavadze 1985, 1999; Rota & de Jong 2015). The biodiversity of the Balkans is the largest in Europe (Griffiths et al. 2004), due to the complex geological history and the specific geographical position with several biogeographical regions, each characterized by specific ecological, climatic and geomorphological conditions, as well as a great variety of habitats (Trakić et al. 2016). For the last 2 million years, repeated glaciations have destroyed the biotas of much of northern and alpine Europe, and on the major mountain ranges such as the Carpathians, Caucasus, and Pyrenees. The present European earthworm fauna consists of the survivors in unglaciated, non-permafrosted areas, and their descendants who colonized the rest of the continent after the end of the last (W̧rm or Wisconsinan) glaciation. There is no way to know how many species were driven to local or global extinction. Repeated cycles could have contributed to inrefuge speciation by changing vegetation zonation. However, the pre-Pleistocene conditions were probably adequate to support earthworm populations throughout Europe and even into the Arctic, including the North American Arctic, as subtropical vegetation fossils from the Eocene occur on Ellesmere Island (Eberle & Greenwood 2012; Francis 1991). Today, most earthworm scientists agree that the original lumbricid fauna was significantly destroyed during the glacial period in much of Europe. Today's lumbricid fauna come from various elements, differing both in their historical age and in their origins. Namely, the present faunal elements are fragments of modified fauna from the Tertiary Period, that are thought to have originated in the Paleocene or Eocene. These are species that, due to their adaptability to specific conditions, have survived to this day. Such relic species are now present in the Balkans, the Pyrenees and the Apennine peninsula, the northwestern part of Africa (Maghreb), the southern part of France, Sardinia, Corsica, the southern parts of Switzerland and the Czech Republic, Turkey and the central parts of Asia (Omodeo 1952, 1956, 1961, 1988; Bouché 1972, 1983; Mısırlıoğlu 2017; Mršić & Šapkarev 1988; Mršić 1991; Marchán et al. 2020; Marchán et al. 2022b). In addition to relic species, there are also "modern" species that are thought to have appeared in the Miocene and later. A greater expansion of new "modern" species occurred during interglacial and postglacial periods, mainly in the Holocene (Mršić 1991). On the Balkan Peninsula there are 90 endemic species with the largest share in Dendrobaena (25) and Allolobophora (20) (Trakić et al. 2016; Popovic et al. 2022), while 39 have been discovered in the basin of the Carpathians, mostly Dacian endemics. Of these, 12 species are of the genus Octodrilus whose distribution center is located in the Apuseni Mountains in the southern part of the Carpathians (Csuzdi et al. 2011). Other genera appear to be endemic from southern France to the northern Iberian Peninsula. The most speciose is presently Scherotheca with 42 species/subspecies recognized to date, though many other species are still expected to be found (e.g., eight new species were recently described from Corsica; Marchán et al. 2023a). The ecologically important Lumbricidae have been explored using various tools by numerous researchers. Despite significant molecular research, there are still many species that do not have a stable taxonomic status, most of which relate to archaic species with unique taxonomic characteristics and disjunctive distributions (Marchán et al. 2022c). Answers to some of these questions are expected in the near future, and will certainly lead to a better understanding of many concerns regarding the origin and dynamics of the development of the Lumbricidae family as a whole., Published as part of Misirlioğlu, Mete, Reynolds, John Warren, Stojanović, Mirjana, Trakić, Tanja, Sekulić, Jovana, James, Samuel W., Csuzdi, Csaba, Decaëns, Thibaud, Lapied, Emmanuel, Phillips, Helen R. P., Cameron, Erin K. & Brown, George G., 2023, Earthworms (Clitellata, Megadrili) of the world: an updated checklist of valid species and families, with notes on their distribution, pp. 417-438 in Zootaxa 5255 (1) on page 425, DOI: 10.11646/zootaxa.5255.1.33, http://zenodo.org/record/7745110, {"references":["Brown, G. G., James, S. W., Csuzdi, C., Lapied, E., Decaens, T., Reynolds, J. W., Misirlioglu, M., Stovanic, M., Trakic, T., Sekulic, J., Phillips, H. & Cameron, E. (2023) A checklist of megadrile earthworm (Annelida: Clitellata) species and subspecies of the world. Available from: Zenodo. https: // doi. org / 10.5281 / zenodo. 7301848","Bouche, M. B. (1972) Lombriciens de France. Ecologie et Systematique. Institut National de la Recherche Agronomique, Articles de Zoologie-Ecologie animale (Numero hors-serie). pp. 67.","Mrsic, N. (1991) Monograph on earthworms (Lumbricidae) of the Balkans I, II. Slovenska Akademija Znanosti in Umetnosti, Razred za Naravoslovne Vede, Opera, Ljubljana, 31, 1 - 757.","Bouche, M. B. & Qiu, J. - P. (1998) Un nouveau Sparganophilus (Annelida: Oligochaeta) d'Europe, avec considerations paleogeographiques sur les Lumbricina. Documents Pedozoologiques et Integrologiques, 4, 178 - 180.","Csuzdi, Cs. & Zicsi, A. (2003) Earthworms of Hungary (Annelida: Oligochaeta: Lumbricidae). Hungarian Natural History Museum, Budapest, 271 pp.","Dominguez, J., Aira, M., Breinholt, J. W., Stojanovic, M., James, S. W. & Perez-Losada, M. (2015) Underground evolution: New roots for the old tree of lumbricid earthworms. Molecular Phylogenetics and Evolution, 83, 7 - 19. https: // doi. org / 10.1016 / j. ympev. 2014.10.024","de Sosa, I., Cosin, D. J. D., Csuzdi, Cs., Paoletti, M. G. & Marchan, D. F. (2019) Placing Eophila tellinii (Oligochaeta, Lumbricidae) in a molecular phylogenetic context advances the century-old controversy around the problematic genus. European Journal of Soil Biology, 94, 103 - 114. https: // doi. org / 10.1016 / j. ejsobi. 2019.103114","Marchan, D. F., Decaens, T., Dominguez, J. & Novo, M. (2022 a) Perspectives in earthworm molecular phylogeny: recent advances in Lumbricoidea and standing questions. Diversity, 14, 30. https: // doi. org / 10.3390 / d 14010030","Marchan, D. F., James, S. W., Lemmon, A. R.,. Lemmon, E. M.,. Novo, M.,. Dominguez, J., Diaz Cosin, D. & Trigo, D. (2022 c) A strong backbone for an invertebrate group: anchored phylogenomics improves the resolution of genus-level relationships within the Lumbricidae (Annelida, Crassiclitellata). Organisms Diversity & Evolution, 22, 915 - 924. https: // doi. org / 10.1007 / s 13127 - 022 - 00570 - y","Blakemore, R. J. (2009) Cosmopolitan earthworms - a global and historical perspective. In: Shain, D. H. (Ed.), Annelids as Model Systems in the Biological Sciences. John Wiley & Sons, New York, pp. 257 - 283. https: // doi. org / 10.1002 / 9780470455203. ch 14","Fragoso, C. & Rojas, P. (2014) Biodiversidad de lombrices de tierra (Annelida: Oligochaeta: Crassiclitellata) en Mexico. Revista Mexicana de Biodiversidad, Suplemento 85, S 197 - S 207. https: // doi. org / 10.7550 / rmb. 33581","Reynolds, J. W. (2020) Earthworms in American ecoregions. LAP LAMBERT Academic Publishing, Mauritius. 433 pp.","Fragoso, C. & Brown, G. G. (2007) Ecologia y taxonomia de las lombrices de tierra en latinoamerica: El Primer Encuentro Latino- Americano de Ecologia y Taxonomia de Oligoquetos (ELAETAO 1). In: Brown, G. G. & Fragoso, C. (Eds.), Minhocas na America Latina: Biodiversidade e ecologia. Embrapa Soja, Londrina, pp. 33 - 75.","Kvavadze, E. Sh. (1985) The earthworms (Lumbricidae) of the Caucasus. Metsniereba, Tbilisi, 238 pp. [In Russian, English summary]","Kvavadze, E. Sh. (1999) Caucasian earthworms (Lumbricidae) (Systematics, Faunistic, Zoogeography, Ecology, Phylogeny). Dr. Sci. Dissertation, Institute of Zoology, Georgian Academy of Sciences, Tbilisi, 313 pp. [In Georgian]","Rota, E. &. de Jong, Y. (2015) Fauna Europaea: Annelida - Terrestrial Oligochaeta (Enchytraeidae and Megadrili), Aphanoneura and Polychaeta. Biodiversity Data Journal, 3, 1 - 47. https: // doi. org / 10.3897 / BDJ. 3. e 5737","Griffiths, G. H. I., Krystufek, B. & Reed, M. (2004) Balkan Biodiversity. Patterns and processes in the European Hotspot. Kluwer, Boston / London, 357 pp. https: // doi. org / 10.1007 / 978 - 1 - 4020 - 2854 - 0","Trakic, T., Valchovski, H. & Stojanovic, M. (2016) Endemic earthworms (Oligochaeta: Lumbricidae) of the Balkan Peninsula: a review. Zootaxa, 4189, 251 - 274. https: // doi. org / 10.11646 / zootaxa. 4189.2.3","Eberle, J. J. & Greenwood, D. R. (2012) Life at the top of the greenhouse Eocene World - A review of the Eocene flora and vertebrate fauna from Canada's high Arctic. Geological Society of America Bulletin, 124, 3 - 23. https: // doi. org / 10.1130 / B 30571.1","Francis, J. E. (1991) The dynamics of polar fossil forests: Tertiary fossil forests of Axel Heiberg Island, Canadian Arctic Archipelago. Geological Survey of Canada, Bulletin, 403, 29 - 38. https: // doi. org / 10.4095 / 131946","Omodeo, P. (1952) Particolarita della Zoogeografia dei lombrichi. Bolletino di Zoologia, 19, 349 - 369. https: // doi. org / 10.1080 / 11250005209439182","Omodeo, P. (1956) Contributo alla revisione dei Lumbricidae. Archivio Zoologico Italiano, 41, 129 - 212.","Omodeo, P. (1961) Le peuplement des Iles Mediterranees et le probleme de l'insularite. Colloques Internationaux du Centre National de la Recherche Scientifique, 94, 128 - 133.","Omodeo, P. (1988) The genus Eophila (Lumbricidae, Oligochaeta). Bolletino di Zoologia, 55, 73 - 88. https: // doi. org / 10.1080 / 11250008809386603","Bouche, M. B. (1983) The establishment of earthworm communities. In: Satchell, J. E. (Ed.), Earthworm ecology from Darwin to vermiculture. Chapman & Hall, London, U. K., pp. 431 - 448. https: // doi. org / 10.1007 / 978 - 94 - 009 - 5965 - 1 _ 38","Misirlioglu, M. (2017) Topraksolucanlari, Biyolojileri, Ekolojileri, Zirai Yonleri, Turkiye Turleri ve Turlerin Taksonomik Ozellikleri. 2 nd Edition. Nobel Yayinlari, Ankara, 168 pp.","Mrsic, N. & Sapkarev, J. (1988) Revision of the genus Allolobophora Eisen, 1874 (sensu Pop 1941) (Lumbricidae, Oligochaeta). Acta Musei Macedonici Scientiarum Naturalium, 19, 1 - 38.","Marchan, D. F., Decaens, T., Diaz Cosin, D. J., Hedde, M., Lapied, E. & Dominguez, J. (2020) French Mediterranean islands as a refuge of relic earthworm species: Cataladrilus porquerollensis sp. nov. and Scherotheca portcrosana sp. nov. (Crassiclitellata, Lumbricidae). European Journal of Taxonomy, 701, 1 - 22.","Marchan D. F., Gerard S., Hedde M., Rougerie R. & Decaens T. (2022 b) An updated checklist and a DNA barcode library for the earthworms (Crassiclitellata, Oligochaeta) of Corsica, France. Zoosystema, 44 (17), 439 - 461. https: // doi. org / 10.5252 / zoosystema 2022 v 44 a 17","Popovic, F. J., Stojanovic, M. M., Dominguez, J., Sekulic, J. M., Trakic, T. B. & Marchan, D. F. (2022) Molecular analysis of five controversial Balkanic species of Allolobophora (sensu lato) Eisen, 1873 (Lumbricidae, Clitellata) with emendation of the genus Cernosvitovia. Omodeo, 1956. Zootaxa, 5116 (3), 351 - 372. https: // doi. org / 10.11646 / zootaxa. 5116.3.3","Csuzdi, Cs., Pop, V. V. & Pop, A. A. (2011) The earthworm fauna of the Carpathian Basin with new records and description of three new species (Oligochaeta: Lumbricidae). Zoologischer Anzeiger, 250, 2 - 18. https: // doi. org / 10.1016 / j. jcz. 2010.10.001","Marchan, D. F., Dominguez, J., Hedde, M. & Decaens, T. (2023 a) The cradle of giants: insight into the origin of Scherotheca (Lumbricidae, Crassiclitellata) with description of eight new species from Corsica, France. Zoosystema, submitted."]}

Boucheona tenebrae Marchán & Novo sp. nov. urn:lsid:zoobank.org:pub: FA29B304-DD7C-4472-BDE2-618B80557709 Hormogaster praetiosa var. nigra Bouché, 1970 Etymology This species name refers to its very dark, almost black pigmentation. Material examined Holotype FRANCE • Adult; Occitanie, Aude, Quillan 19/04/2008; leg. M. Novo, R. Fernández; Locality 1 (QUILL): 42.880222 2.170000; shrubland on a pronounced slope; voucher UCMLT00393; deposited in the UCMLT. Paratypes FRANCE • 1 specimen, adult; 16/04/2012; leg. D. F. Marchán, R. Fernández, same data as holotype; voucher UCMLT00394; deposited in UCMLT. Diagnosis Pigmentation almost black with purple iridescence, length reaching 50 cm, weight reaching 27.36 gr, number of segments reaching 600. Clitellum extending over segments XIV-XXX, tubercula pubertatis as a continuous line which narrows towards the end on segments XXII-XXVII(XXVIII), first thickened septum in 6/7, multichamber disc-shaped spermathecae in intersegments 9/10 and 10/11, typhlosole composed of a maximum of 13 lamellae, first nephridial cecae in XI. Morphological description External morphology. Recently collected specimens (used for molecular information) are incomplete and damaged, thus some characters refer to the description of Hormogaster praetiosa var. nigra (Bouché, 1970) Length: 50 cm according to Bouché 1970 (living specimens?). Weight (holotype): 27.36 gr (20-25 according to Bouché 1970). Number of segments (holotype): 580 (more than 600 according to Bouché 1970). Pigmentation: Almost black with purple iridescence on living specimens, with pigmentation neatly interrupted at the level of chaetae b. Dorsally dark grey, ventrally beige on fixed specimens. Prostomium prolobic, strongly developed and with a small invagination delimiting a small prostomium-like lobe; longitudinal striation on segments I, II and III. Strongly developed cephalic keels on segments I to XIII. Closely paired chaetae; interchaetal ratio at segment 40, aa: 50, ab: 1, bc: 8, cd: 1, dd: 50. Nephridial pores in a row between chaetae b and c (very close to b). Spermathecal pores on intersegments 9/10 and 10/11 at the level of cd. Male pores open over chaetae ab at the intersegment 15/16, surrounded by heart-shaped porophores. Female pores in segment XIV at the same level as male pores. Clitellum ring shaped-shaped (only when wholly developed) extending over segments XIV-XXX. Tubercula pubertatis on XXII-XXVII(XXVIII) as a continuous line which narrows towards the end. Papillae of chaetae ab in variable positions between segments XIII, XIV, XVII-XXIII, XXVIII, XXIX. Internal anatomy. Funnel-shaped, strongly thickened septa in 7/8 to 11/12, moderately thickened in 6/7. Clearly developed hearts in VI-XI.Three shiny, strongly muscular gizzards in VI, VII and VIII. Morren's glands and intestinal gizzard not apparent. First section of the intestine slightly dilated. Typhlosole begins around segments XX with 9 lamellae, which swiftly increase to a maximum 13 lamellae and the decrease to 11 lamellae. Anterior nephridial bladders U-shaped with very close branches and no apparent caecum until segment XI. Bladders gradually flatten towards the end of the body, taking the usual elongated shape. Fraying testes and iridescent seminal funnels in X and XI. Two pairs of voluminous, grainy seminal vesicles in XI and XII. Ovaries and female funnels in XIII, ovisacs in XIV. Two pairs of multichamber disc-shaped spermathecae in intersegments 9/10 and 10/11, the posterior pair bigger. Remarks Boucheona tenebrae sp. nov. resembles B. catalaunensis, B. gallica, B. corbierensis sp. nov. and B. rosae in the position of the clitellum (XIV-XXIX,XXX) and type and position of spermathecae (multichamber disc-shaped spermathecae in intersegments 9/10 and 10/11). It differs from these close relatives in its larger size (length, weight, number of segments), its darker pigmentation (almost black vs brown-dark brown) and the position of the first thickened septum (6/7 vs 7/8). In addition, it differs from B. corbierensis sp. nov. in the position of the tubercula pubertatis (XXII-XXVII(XXVIII) vs XXII-XXIX) and maximum number of typhlosole lamellae (13 vs 15). COI uncorrected average pairwise distances and topology of multilocus molecular phylogenetic trees support the status of B. tenebrae sp. nov. as independent from other morphologically similar species. Distribution and ecology B. tenebrae sp. nov. is known from a single locality of Aude (Figure 5). It inhabits shrublands., Published as part of Marchán, Daniel F., Novo, Marta, Domínguez, Jorge, Sánchez, Nuria & Decaëns, Thibaud, 2023, Contribution to the knowledge of the genus Boucheona (Oligochaeta Hormogastridae) in France with a newly described species and a redescribed species, pp. 68-81 in Zootaxa 5255 (1) on pages 75-78, DOI: 10.11646/zootaxa.5255.1.11, http://zenodo.org/record/7744489, {"references":["Bouche, M. B. (1972) Lombriciens de France. Ecologie et systematique, Vol. 72, No. HS. INRA Editions, Paris, 671 pp."]}