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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."]}

Tritogeniidae Plisko, 2013 The family Tritogeniidae was erected by Plisko(2013) as part of important housekeeping duties within the Microchaetidae, and includes two genera, Michalakus with a single species and the others all belonging to Tritogenia. All 39 species are endemic to Southern Africa, with only one species newly described from Botswana (Nxele et al. 2018)., 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 429, DOI: 10.11646/zootaxa.5255.1.33, http://zenodo.org/record/7745110, {"references":["Plisko, J. D. (2013) A new family Tritogeniidae for the genera Tritogenia and Michalakus, earlier accredited to the composite Microchaetidae (Annelida: Oligochaeta). African Invertebrates, 54 (1), 69 - 92. https: // doi. org / 10.5733 / afin. 054.0107","Nxele, T. C., Plisko, J. D. & Ramongalo, B. K. (2018). The first proandric species of Tritogenia Kinberg, 1867 from Botswana, Tritogenia talana sp. n. (Clitellata, Crassiclitellata, Tritogeniidae). Zootaxa, 4496, 214 - 217. https: // doi. org / 10.11646 / zootaxa. 4496.1.17"]}

Tumakidae Righi, 1995 The monogeneric Tumakidae family has only three known species, Tumak hammeni Righi, Tumak amari Celis & Rangel-Ch., and Tumak congorum Celis & Rangel-Ch., found in relatively drier, lower-elevation forests in Central Colombia and the Caribbean region, along the Magdalena River valley (Righi 1995; Celis & Rangel-Ch. 2015). Given the relatively small amount of remaining forest fragments of its native habitat in Colombia, due primarily to agricultural expansion and pastures for cattle, further efforts are needed to ensure adequate conservation of the few species known. Additional sampling efforts are also needed to discover if there are any other species present along its potential distribution range., 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 429, DOI: 10.11646/zootaxa.5255.1.33, http://zenodo.org/record/7745110, {"references":["Righi, G. (1995) Colombian earthworms. In: Van der Hammen, T. & Santos, A. G. (Eds.), Studies on Tropical Andean Ecosystems, vol. 4. Cramer, Berlin, pp. 485 - 607."]}

Eudrilidae Claus, 1880 The Eudrilidae is a peculiar earthworm family with internal fertilisation (Clausen 1965; Sims 1969). It consists of 45 genera and 305 species (plus 21 subspecies) distributed in tropical Africa from south of the Sahara to the Kalahari Desert and northern South Africa (Sims 1987). There are only two species in the family regarded as widely introduced peregrines: Eudrilus eugeniae (Kinberg) and Hyperiodrilus africanus Beddard. The first species, commonly called the "African Nightcrawler" is widespread, found mainly in vermiculture outside its native range (West Africa) throughout the tropics and sub-tropics in Latin America, the Caribbean, Madagascar, Sri Lanka, India, Philippines, Malaysia, Indonesia, Vietnam and Australia (Blakemore 2015). Rarely has this species been found inhabiting soils (Blakemore 2015) and not compost beds, and its survival under these conditions probably requires high moisture and the presence of abundant litter, or organic resources, which are its major food source. The second species, H. africanus can also be found frequently in compost-beds and high organic content soils, but its distribution is more restricted, being known from Western (Ivory Coast, Nigeria, Gabon) and Central Africa (Congo, Democratic Republic of Congo, Angola) (Tondoh & Lavelle 2005) and Brazil (Righi 1972; Sousa et al. 2020)., 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":["Clausen, M. W. (1965) Description of two new species of Libyodrilus Beddard, 1891, with remarks on the genus and on the female system in the family Eudrilidae. Videnskabelige Meddelelser Dansk Naturhistorisk Forening, 128, 273 - 292.","Sims, R. W. (1969) Internal fertilization in Eudrilid earthworms with the description of a new Pareudriline genus and species (Oligochaeta) from Ghana. Journal of Zoology, 157, 437 - 447. https: // doi. org / 10.1111 / j. 1469 - 7998.1969. tb 01714. x","Sims, R. W. (1987) Review of the Central African earthworm family Eudrilidae (Oligochaeta). In: Bonvicini Pagiliai, A. M. & Omodeo, P. (Eds.), On earthworms. Mucchi, Modena, pp. 359 - 388.","Blakemore, R. J. (2015) Eco-taxonomic profile of an iconic vermicomposter - the ' African Night-crawler' earthworm Eudrilus eugeniae (Kinberg, 1867). African Invertebrates, 56 (3), 527 - 548. https: // doi. org / 10.5733 / afin. 056.0302","Tondoh, J. E. & Lavelle, P. (2005) Population dynamics of Hyperiodrilus africanus (Oligochaeta, Eudrilidae) in Ivory Coast. Journal of Tropical Ecology, 21, 493 - 500. https: // doi. org / 10.1017 / S 0266467405002506","Righi, G. (1972) Contribuic \" o ao conhecimento dos Oligochaeta brasileiros. Papeis Avulsos de Zoologia, 25, 148 - 166.","Sousa, S. C., Hernandez-Garcia, L. M. & Rousseau, G. X. (2020) A new species of Pontoscolex earthworm (Rhinodrilidae, Clitellata) from the Gurupi Biological Reserve, along with records of earthworm species from the Amazon region of Maranh \" o, Brazil. Zootaxa, 4801, 105 - 114. https: // doi. org / 10.11646 / zootaxa. 4801.1.4"]}

Kazimierzidae Nxele & Plisko, 2016 This recently erected family of earthworms, which includes 25 nominal species all in the single genus Kazimierzus, is restricted to the Western Cape and Northern Cape provinces, including the Namaqualand areas, along the Atlantic region of South Africa. This is a hotspot of biodiversity (Myers et al. 2000), with considerable topographical and botanic diversity, which will probably still lead to the discovery of new earthworm species, as observed by Nxele 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":["Nxele, T. C., Plisko, J. D., Mwabvu, T. & Zishiri, T. O. (2016) A new family Kazimierzidae for the genus Kazimierzus, earlier recorded to the composite Microchaetidae (Annelida, Oligochaeta). African Invertebrates, 57 (2), 111 - 117. https: // doi. org / 10.3897 / AfrInvertebr. 57.10042","Myers, N., Mittermeier, R. A, Mittermeier, C. G, Fonseca, G. A. B. & Kent, J. (2000) Biodiversity hotspots for conservation priorities. Nature, 403, 853 - 858. https: // doi. org / 10.1038 / 35002501","Nxele, T. C., Plisko, J. D., Mwabvu, T. & Zishiri, O. T. (2017) Four new earthworm species of Kazimierzus Plisko, 2006 (Clitellata, Kazimierzidae). Zootaxa, 4353, 187 - 194. https: // doi. org / 10.11646 / zootaxa. 4353.1.12"]}

Microchaetidae Michaelsen, 1900 The family Microchaetidae is closely related to the two other Microchaetoidea families Tritogeniidae and Kazimierzidae (Plisko 2013; Nxele et al. 2016), and consists of three genera (Geogenia, Microchaetus and Proandricus) and 81 species (plus six subspecies) distributed exclusively in temperate and Mediterranean Southern Africa. They are known from Northern South Africa near the vicinity of the Limpopo River down to the southern limit of the continent, in all provinces as well as in Lesotho and Swaziland (Plisko 2013). The three genera appear to have a rather geographically separate distribution, with little overlap. None of the known species have been reported outside of this rather restricted range, although Plisko (2013) predicted that Microchaetidae would also be found in Zimbabwe, Botswana and Mozambique. Interestingly, some of the longest known earthworm species in the world, including Microchaetus vernoni Plisko (measuring up to 2.6 m when alive) belong to this family, which contains a reasonably large number of giant (very large or long) earthworm species, some with over 1,000 segments, particularly in the genus Microchaetus (Plisko 1992, 2013)., 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":["Plisko, J. D. (2013) A new family Tritogeniidae for the genera Tritogenia and Michalakus, earlier accredited to the composite Microchaetidae (Annelida: Oligochaeta). African Invertebrates, 54 (1), 69 - 92. https: // doi. org / 10.5733 / afin. 054.0107","Nxele, T. C., Plisko, J. D., Mwabvu, T. & Zishiri, T. O. (2016) A new family Kazimierzidae for the genus Kazimierzus, earlier recorded to the composite Microchaetidae (Annelida, Oligochaeta). African Invertebrates, 57 (2), 111 - 117. https: // doi. org / 10.3897 / AfrInvertebr. 57.10042","Plisko, J. D. (1992) The Microchaetidae of Natal, with description of new species of Microchaetus Rapp and Tritogenia Kinberg, and the new genus Proandricus (Oligochaeta). Annals of the Natal Museum, 33, 337 - 378."]}

Diporodrilidae Bouché, 1970 This monogeneric family with three species and two subspecies was erected by Bouché (1970) for specimens collected in the Mediterranean islands of Corsica and Sardinia, from where they are endemic. Recent sampling in Corsica revealed several new lineages and species that must still be described (Marchán et al., 2022b). The validity of this family was also re-affirmed with recent molecular (Anchor-Hybrid Enrichment) analyses, which placed it as sister to the Lumbricidae (Marchán et al., 2022c)., 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":["Bouche, M. B. (1970). Remarques sur quelques Lumbricina de France et consequences de la decouverte des nouveaux taxons Vignysinae (Subfam. nov.) et Diporodrilidae (Fam. nov.). Pedobiologia, 10 (4), 246 - 256.","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","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"]}

Komarekionidae Gates, 1974 There is only one species in this monogeneric Nearctic family: Komarekiona eatoni Gates. It is found only in the mid-Atlantic states west to southern Illinois, USA (Gates 1974; Reynolds 2020). Interestingly, the specimens east of the Smoky Mountains are amphimictic while those west of the Smokies are parthenogenetic (Reynolds 2020)., 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":["Gates, G. E. (1974) On a new species of earthworm in a southern portion of the United States. Bulletin of Tall Timbers Research Station, 15, 1 - 13.","Reynolds, J. W. (2020) Earthworms in American ecoregions. LAP LAMBERT Academic Publishing, Mauritius. 433 pp."]}

Criodrilidae Vejdovský, 1884 The Palearctic family Criodrilidae includes mainly aquatic species in a single genus, Criodrilus with two valid species, C. lacuum from Europe, and C. ghianae Qiu & Bouché, from Mediterranean Algeria. The former species, may be among the longest living of earthworms, known to achieve 46 years of age when kept in aquaria (Timm 2020). The species is widely distributed in Europe and common all around the Mediterranean, with its northernmost distribution in Latvia (Smiljkov et al. 2005; Blakemore 2008b; Atanacković et al. 2013; Valchovski 2013; Mısırlıoğlu 2017; Timm 2020). Outside its native range, C. lacuum has only been reported from potted plants in Baltimore (USA) (McKey-Fender & MacNab 1953). Previous reports of this species from Brazil (Knäpper 1976; Knäpper & Porto 1979) actually refer to G. camaqua, a species of Almidae (see above, and 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 423, DOI: 10.11646/zootaxa.5255.1.33, http://zenodo.org/record/7745110, {"references":["Timm, T. (2020) Observations on the life cycles of aquatic Oligochaeta in aquaria. Zoosymposia, 17, 102 - 120. https: // doi. org / 10.11646 / zoosymposia. 17.1.11","Smiljkov, S., Trajanovski, S. & Budzakoska-Goreska, B. (2005) Biocenotic composition of the macrozoobenthos on different habitats from the littoral region of lake Ohrid, Prilozi, 26 (2), 143 - 155.","Blakemore, R. (2008 b) Review of Criodrilidae (Annelida: Oligochaeta) including Biwadrilus from Japan. Opuscula Zoologica Budapest, 37, 11 - 22.","Atanackovic, A. D., Sporka, F., Csanyi, B., Vasiljevic, B. M., Tomovic, J. M. & Paunovic, M. M. (2013) Oligochaeta of the Danube River - a faunistical review. Biologia, 68 (2), 269 - 277. https: // doi. org / 10.2478 / s 11756 - 013 - 0155 - 9","Valchovski, H. (2013) Contribution to the knowledge for distribution of Criodrilus lacuum (Annelida: Oligochaeta: Criodrilidae) from Bulgaria. ZooNotes, 44, 1 - 3.","Misirlioglu, M. (2017) Topraksolucanlari, Biyolojileri, Ekolojileri, Zirai Yonleri, Turkiye Turleri ve Turlerin Taksonomik Ozellikleri. 2 nd Edition. Nobel Yayinlari, Ankara, 168 pp.","McKey-Fender, D. & Macnab, J. A. (1953) The aquatic earthworm Criodrilus lacuum Hoffmeister in North America (Oligochaeta Glossoscolecidae). Wasmann Journal of Biology, 113, 373 - 378.","Knapper, C. F. U. (1976) Preliminar considerations on the occurrence of Oligochaetes in the Estuary of the Guaiba RS. Estudos Leopoldenses, 38, 39 - 41.","Knapper C. F. U. & Porto, R. P. (1979) Ocorrencia de Oligoquetas nos solos do Rio Grande do Sul. Acta Biologica Leopoldensia, 1, 137 - 166.","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"]}

Hormogastridae Michaelsen, 1900 The family Hormogastridae was recently revised by Marchán et al. (2018) using an integrative approach including ecological, morphological and molecular data, and was shown to include nine genera (Ailoscolex, Boucheona, Carpetania, Diazcosinia, Hemigastrodrilus, Hormogaster, Norana, Vignysa, Xanina) and 37 species (plus three subspecies). The genus Ailoscolex, with its single known species Ailoscolex lacteospumosus Bouché, formerly in the family Ailoscolecidae was basal to all Hormogastridae but kept within the family. The genera show rather restricted distribution and little geographic overlap, with the exception of Norana with Hormogaster and Boucheona (Marchán et al. 2018). Overall, the family is restricted to Europe and Northern Africa, being found in Southern France, the Spanish Iberian Peninsula, Corsica, Sardinia, the Tuscan Archipelago, mainland Italy, Sicily, Algeria and Tunisia. So far, no Hormogastridae have been found in Portugal. Given the restricted range of most known species, the future discovery of new taxa in this family is highly probable (see Marchán et al. 2023b)., 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":["Marchan, D. F., Fernandez R., de Sosa I., Sanchez N., Diaz Cosin D. J. & Novo M. (2018) Integrative systematic revision of a Mediterranean earthworm family: Hormogastridae (Annelida, Oligochaeta). Invertebrate Systematics, 32, 652 - 671. https: // doi. org / 10.1071 / IS 17048","Marchan, D. F., Novo, M., Dominguez, J., Sanchez, N. & Decaens, T. (2023 b) Contribution to the knowledge of the Genus Boucheona (Oligochaeta, Hormogastridae) in France with description of two new species. Zootaxa, 5255 (1), 68 - 81. https: // doi. org / 10.11646 / zootaxa. 5255.1.11"]}

Megascolecidae Rosa, 1891 The Megascolecidae is the most speciose family of earthworms (2,208 spp. and 127 subspecies) and includes 85 genera. The most speciose genera are Amynthas (713 spp./ssp.), Metaphire (242 spp./sspp.), Pheretima (171 spp./ sspp.), and Megascolex (104 sp. /sspp.). The available molecular studies all support its monophyly and show just slight differentiation inside the family. Therefore, given the contradictory taxonomic divisions (e.g., Jamieson et al. 2002; Blakemore 2013) here we do not distinguish subordinate family rank taxa. The Megascolecidae show a predominantly Australasian distribution being most speciose in Australia (ca. 45 genera and 400 species), New Zealand (8 genera and 50 spp.), Eastern and Southeastern Asia (over 1,500 spp.) (Sims 1980; Jamieson 2000). But interestingly, the North American megascolecids seem to be basal to several modern megascolecid groups such as the Asian Amynthas and Australian Perionychella, Megascolides, and Diporochaeta (Buckley et al. 2011). These include Arctiostrotus (7 species), Argilophilus (9 species), Chetcodrilus (3 species), Kincaidodrilus (1 species), Macnabodrilus (2 species), Nephrallaxis (2 species) and Toutellus (4 species), present mostly in the Pacific Northwestern States of the United States and Southern Canada (near Vancouver), but also in California (Reynolds, 2020, 2022). At least 27 species in the genera Amynthas and Metaphire, as well as Perionyx excavatus Perrier, Pithemera bicincta (Perrier), Polypheretima elongata (Perrier), Polypheretima taprobanae (Beddard) and Pontodrilus litoralis (Grube) are probably some of the most widely distributed earthworms in the world (Blakemore, 2009). In fact, many of these species were described from specimens collected outside their native ranges in Asia: for instance, Metaphire californica (Kinberg) from San Francisco, Amynthas gracilis (Kinberg) from Rio de Janeiro, Amynthas corticis (Kinberg) from Hawaii, and P. elongata from Peru. Therefore, they had been already extensively transported by humans from Asia to other continents even before they were recorded from their home ranges. The presence of parthenogenetic morphs, and wide plasticity in terms of soil and habitat preferences in several of these species means that they are excellent invaders, particularly in subtropical, tropical and even temperate regions (Brown et al. 2006; Chang et al. 2017). Some species, such as Pi. bicincta may have been extensively transported with Polynesians that sailed throughout the Pacific Ocean, as well as in flotsam or sailing ship ballast (Blakemore 2007; James 2011). Several Amynthas and Metaphire species have invaded cold-temperate regions, including the continental USA, causing extreme alterations in the native ecosystems (Chang 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 426, DOI: 10.11646/zootaxa.5255.1.33, http://zenodo.org/record/7745110, {"references":["Jamieson, B. G. M., Tillier, S., Tillier, A., Justine, J. - L., Ling, E., James, S., MacDonald, K. & Hugall, A. F. (2002) Phylogeny of the Megascolecidae and Crassiclitellata (Annelida, Oligochaeta): combined versus partitioned analysis using nuclear (28 S) and mitochondrial (12 S, 16 S) rDNA. Zoosystema, 24, 707 - 734.","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.","Sims, R. W. (1980) A classifications and the distribution of earthworms, suborder Lumbricina (Haplotaxida: Oligochaeta). Bulletin of the British Museum Natural History (Zoology), 39 (2), 103 - 124.","Jamieson, B. G. M. (2000) The native earthworms of Australia (Megascolecidae Megascolecinae). Science Publishers, Enfield, USA (CD-ROM).","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","Reynolds, J. W. (2020) Earthworms in American ecoregions. LAP LAMBERT Academic Publishing, Mauritius. 433 pp.","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","Brown, G. G., James, S. W., Pasini, A., Nunes, D. H., Benito, N. P., Martins, P. T. & Sautter, K. D. (2006) Exotic, peregrine, and invasive earthworms in Brazil: Diversity, distribution, and effects on soils and plants. Caribbean Journal of Science, 42, 339 - 358.","Chang, C. - H., Snyder, B. & Szlavecz K. (2017) Asian pheretimoid earthworms in North America north of Mexico: An illustrated key to the genera Amynthas, Metaphire, Pithemera, and Polypheretima (Clitellata: Megascolecidae). Zootaxa, 4179 (3), 495 - 529. https: // doi. org / 10.11646 / zootaxa. 4179.3.7","Blakemore, R. J. (2007) Origin and means of dispersal of cosmopolitan Pontodrilus litoralis (Oligochaeta: Megascolecidae). European Journal of Soil Biology, 43, S 3 - S 8. https: // doi. org / 10.1016 / j. ejsobi. 2007.08.041","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","Chang, C. - H., Bartz, M L. C., Brown, G., Callaham Jr., M. A., Cameron, E. K., Davalos, A., Dobson, A., G ˆ rres, J. H., Herrick, B. M., Ikeda, H., James, S. W., Johnston, M. R., McCay, T. S., McHugh, D., Minamiya, Y., Nouri-Aiin, M., Novo, M., Ortiz- Pachar, J., Pinder, R. A., Ransom, T., Richardson, J. B., Snyder, B. A. & Szlavecz, K. (2021) The second wave of earthworm invasions in North America: biology, environmental impacts, management and control of invasive jumping worms. Biological Invasions, 23, 3291 - 3322. https: // doi. org / 10.1007 / s 10530 - 021 - 02598 - 1"]}

Kynotidae Jamieson, 1971 This endemic family appears to be restricted to Madagascar (Razafindrakoto et al. 2017) and includes a single genus (Kynotus) and 22 species. The description of K. verticillatus Perrier, was not adequate and it cannot be assigned to any other named species or any other genus or species. It was described as Acanthodrilus “ pas du tout développé et paraissant réduit aux quatre pénis formés de soies courbes caractéristiques jusqu'ici des vers de ce genre ” (Perrier 1872)., 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":["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.","Razafindrakoto, M., Csuzdi, C., James, S. & Blanchart, E. (2017) New earthworms from Madagascar with key to the Kynotus species (Oligochaeta: Kynotidae). Zoologischer Anzeiger, 268, 126 - 135. https: // doi. org / 10.1016 / j. jcz. 2016.08.001"]}

This repository contains additional data associated to Tedersoo et al. (2022) (https://doi.org/10.1111/gcb.16398).This repository contains those data necessary to rerun the superecoregion design and the endemicity analyses as well as vector maps on the endemicity and OTU richness of individual superecoregions. Further data associated with the same publication, regarding the vulnerability analysis, can be found here: 10.5281/zenodo.6983158. input.zip - Input data needed to run the analysis scripts provided [here](https://github.com/Mycology-Microbiology-Center/Fungal_Endemicity_and_Vulnerability/superecoregions_and_endemicity). The entire folder is to be copied in the working directory to run the pipeline. output.zip - Further data needed to run the analysis scripts provided [here](https://github.com/Mycology-Microbiology-Center/Fungal_Endemicity_and_Vulnerability/superecoregions_and_endemicity). The entire folder is to be copied in the working directory to run the pipeline. maps_otu_numbers_and_se_characteristics - vector maps of the superecoregions and the number of OTUs and endemic OTUs per region for different functional groups maps_traits_endemism - vector maps of endemicity indices per region f different functional groups 01_super_Ecoregions_numbers.pdf - vector map of the superecoregions used in the study numbered from 1 - 174. legend_01_super_ecoregions_numbers.csv - legend with superecoregion names for 01_super_Ecoregions_numbers.pdf. super_eoregions_results - Superecoregions with number of OTUs, number of endemic OTUs and endemicity indices in R data format, for further analyses and visualization super_ecoregions.shp - Superecoregions with number of OTUs, number of endemic OTUs and endemicity indices as shape file, for further analyses and visualization &nbsp

A growing body of work examines the direct and indirect effects of climate change on ecosystems, typically by using manipulative experiments at a single site or performing meta-analyses across many independent experiments. However, results from single-site studies tend to have limited generality. Although meta-analytic approaches can help overcome this by exploring trends across sites, the inherent limitations in combining disparate datasets from independent approaches remain a major challenge. In this paper, we present a globally distributed experimental network that can be used to disentangle the direct and indirect effects of climate change. We discuss how natural gradients, experimental approaches, and statistical techniques can be combined to best inform predictions about responses to climate change, and we present a globally distributed experiment that utilizes natural environmental gradients to better understand long-term community and ecosystem responses to environmental change. The warming and (species) removal in mountains (WaRM) network employs experimental warming and plant species removals at high- and low-elevation sites in a factorial design to examine the combined and relative effects of climatic warming and the loss of dominant species on community structure and ecosystem function, both above- and belowground. The experimental design of the network allows for increasingly common statistical approaches to further elucidate the direct and indirect effects of warming. We argue that combining ecological observations and experiments along gradients is a powerful approach to make stronger predictions of how ecosystems will function in a warming world as species are lost, or gained, in local communities.

Here we introduce the Soil BON Foodweb Team, a cross-continental collaborative network that aims to monitor soil animal communities and food webs using consistent methodology at a global scale. Soil animals support vital soil processes via soil structure modification, consumption of dead organic matter, and interactions with microbial and plant communities. Soil animal effects on ecosystem functions have been demonstrated by correlative analyses as well as in laboratory and field experiments, but these studies typically focus on selected animal groups or species at one or few sites with limited variation in environmental conditions. The lack of comprehensive harmonised large-scale soil animal community data including microfauna, mesofauna, and macrofauna, in conjunction with related soil functions, microbial communities, and vegetation, limits our understanding of biological interactions in soil systems and how these interactions affect ecosystem functioning. To provide such data, the Soil BON Foodweb Team invites researchers worldwide to use a common methodology to address six long-term goals: (1) to collect globally representative harmonised data on soil micro-, meso-, and macrofauna communities, (2) to describe key environmental drivers of soil animal communities and food webs, (3) to assess the efficiency of conservation approaches for the protection of soil animal communities, (4) to describe soil food webs and their association with soil functioning globally, (5) to establish a global research network for soil biodiversity monitoring and collaborative projects in related topics, (6) to reinforce local collaboration networks and expertise and support capacity building for soil animal research around the world. In this paper, we describe the vision of the global research network and the common sampling protocol to assess soil animal communities and advocate for the use of standard methodologies across observational and experimental soil animal studies. We will use this protocol to conduct soil animal assessments and reconstruct soil food webs at sites associated with the global soil biodiversity monitoring network, Soil BON, allowing us to assess linkages among soil biodiversity, vegetation, soil physico-chemical properties, climate, and ecosystem functions. In the present paper, we call for researchers especially from countries and ecoregions that remain underrepresented in the majority of soil biodiversity assessments to join us. Together we will be able to provide science-based evidence to support soil biodiversity conservation and functioning of terrestrial ecosystems. Made available in DSpace on 2022-04-05T05:00:35Z (GMT). No. of bitstreams: 1 George-SO-Global.pdf: 4233169 bytes, checksum: 9faad657460c5d69e137289ebd0e4488 (MD5) Previous issue date: 2022

Here we introduce the Soil BON Foodweb Team, a cross-continental collaborative network that aims to monitor soil animal communities and food webs using consistent methodology at a global scale. Soil animals support vital soil processes via soil structure modification, consumption of dead organic matter, and interactions with microbial and plant communities. Soil animal effects on ecosystem functions have been demonstrated by correlative analyses as well as in laboratory and field experiments, but these studies typically focus on selected animal groups or species at one or few sites with limited variation in environmental conditions. The lack of comprehensive harmonised large-scale soil animal community data including microfauna, mesofauna, and macrofauna, in conjunction with related soil functions, microbial communities, and vegetation, limits our understanding of biological interactions in soil systems and how these interactions affect ecosystem functioning. To provide such data, the Soil BON Foodweb Team invites researchers worldwide to use a common methodology to address six long-term goals: (1) to collect globally representative harmonised data on soil micro-, meso-, and macrofauna communities, (2) to describe key environmental drivers of soil animal communities and food webs, (3) to assess the efficiency of conservation approaches for the protection of soil animal communities, (4) to describe soil food webs and their association with soil functioning globally, (5) to establish a global research network for soil biodiversity monitoring and collaborative projects in related topics, (6) to reinforce local collaboration networks and expertise and support capacity building for soil animal research around the world. In this paper, we describe the vision of the global research network and the common sampling protocol to assess soil animal communities and advocate for the use of standard methodologies across observational and experimental soil animal studies. We will use this protocol to conduct soil animal assessments and reconstruct soil food webs at sites associated with the global soil biodiversity monitoring network, Soil BON, allowing us to assess linkages among soil biodiversity, vegetation, soil physico-chemical properties, climate, and ecosystem functions. In the present paper, we call for researchers especially from countries and ecoregions that remain underrepresented in the majority of soil biodiversity assessments to join us. Together we will be able to provide science-based evidence to support soil biodiversity conservation and functioning of terrestrial ecosystems.

Earthworms are an important soil taxon as ecosystem engineers, providing a variety of crucial ecosystem functions and services. Little is known about their diversity and distribution at large spatial scales, despite the availability of considerable amounts of local-scale data. Earthworm diversity data, obtained from the primary literature or provided directly by authors, were collated with information on site locations, including coordinates, habitat cover, and soil properties. Datasets were required, at a minimum, to include abundance or biomass of earthworms at a site. Where possible, site-level species lists were included, as well as the abundance and biomass of individual species and ecological groups. This global dataset contains 10,840 sites, with 184 species, from 60 countries and all continents except Antarctica. The data were obtained from 182 published articles, published between 1973 and 2017, and 17 unpublished datasets. Amalgamating data into a single global database will assist researchers in investigating and answering a wide variety of pressing questions, for example, jointly assessing aboveground and belowground biodiversity distributions and drivers of biodiversity change., H.R.P.P., B.K-R., and the sWorm workshops were supported by the sDiv [Synthesis Centre of the German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig (DFG FZT 118)]. H.R.P.P., O.F. and N.E. acknowledge funding by the European Research Council (ERC) under the European Union’s Horizon 2020 research and innovation programme (grant agreement no. 677232 to NE). K.S.R. and W.H.v.d.P. were supported by ERC-ADV grant 323020 to W.H.v.d.P. Also supported by iDiv (DFG FZT118) Flexpool proposal 34600850 (C.A.G. and N.E.); the Academy of Finland (285882) and the Natural Sciences and Engineering Research Council of Canada (postdoctoral fellowship and RGPIN-2019-05758) (E.K.C.); German Federal Ministry of Education and Research (01LO0901A) (D.J.R.); ERC-AdG 694368 (M.R.); the TULIP Laboratory of Excellence (ANR-10-LABX-41) (M.L); and the BBSRC David Phillips Fellowship to F.T.d.V. (BB/L02456X/1). In addition, data collection was funded by the Russian Foundation for Basic Research (12-04-01538-а, 12-04-01734-a, 14-44-03666-r_center_a, 15-29-02724-ofi_m, 16-04-01878-a 19-05-00245, 19-04-00-609-a); Tarbiat Modares University; Aurora Organic Dairy; UGC(NERO) (F. 1-6/Acctt./NERO/2007-08/1485); Natural Sciences and Engineering Research Council (RGPIN-2017-05391); Slovak Research and Development Agency (APVV-0098-12); Science for Global Development through Wageningen University; Norman Borlaug LEAP Programme and International Atomic Energy Agency (IAEA); São Paulo Research Foundation - FAPESP (12/22510-8); Oklahoma Agricultural Experiment Station; INIA - Spanish Agency (SUM 2006-00012-00-0); Royal Canadian Geographical Society; Environmental Protection Agency (Ireland) (2005-S-LS-8); University of Hawai’i at Mānoa (HAW01127H; HAW01123M); European Union FP7 (FunDivEurope, 265171; ROUTES 265156); U.S. Department of the Navy, Commander Pacific Fleet (W9126G-13-2-0047); Science and Engineering Research Board (SB/SO/AS-030/2013) Department of Science and Technology, New Delhi, India; Strategic Environmental Research and Development Program (SERDP) of the U.S. Department of Defense (RC-1542); Maranhão State Research Foundation (FAPEMA 03135/13, 02471/17); Coordination for the Improvement of Higher Education Personnel (CAPES 3281/2013); Ministry of Education, Youth and Sports of the Czech Republic (LTT17033); Colorado Wheat Research Foundation; Zone Atelier Alpes, French National Research Agency (ANR-11-BSV7-020-01, ANR-09-STRA-02-01, ANR 06 BIODIV 009-01); Austrian Science Fund (P16027, T441); Landwirtschaftliche Rentenbank Frankfurt am Main; Welsh Government and the European Agricultural Fund for Rural Development (Project Ref. A AAB 62 03 qA731606); SÉPAQ, Ministry of Agriculture and Forestry of Finland; Science Foundation Ireland (EEB0061); University of Toronto (Faculty of Forestry); National Science and Engineering Research Council of Canada; Haliburton Forest & Wildlife Reserve; NKU College of Arts & Sciences Grant; Österreichische Forschungsförderungsgesellschaft (837393 and 837426); Mountain Agriculture Research Unit of the University of Innsbruck; Higher Education Commission of Pakistan; Kerala Forest Research Institute, Peechi, Kerala; UNEP/GEF/TSBF-CIAT Project on Conservation and Sustainable Management of Belowground Biodiversity; Ministry of Agriculture and Forestry of Finland; Complutense University of Madrid/European Union FP7 project BioBio (FPU UCM 613520); GRDC; AWI; LWRRDC; DRDC; CONICET (National Scientific and Technical Research Council) and FONCyT (National Agency of Scientific and Technological Promotion) (PICT, PAE, PIP), Universidad Nacional de Luján y FONCyT (PICT 2293 (2006)); Fonds de recherche sur la nature et les technologies du Québec (131894); Deutsche Forschungsgemeinschaft (SCHR1000/3-1, SCHR1000/6-1, 6-2 (FOR 1598), WO 670/7-1, WO 670/7-2, & SCHA 1719/1-2), CONACYT (FONDOS MIXTOS TABASCO/PROYECTO11316); NSF (DGE-0549245, DGE-0549245, DEB-BE-0909452, NSF1241932, LTER Program DEB-97–14835); Institute for Environmental Science and Policy at the University of Illinois at Chicago; Dean’s Scholar Program at UIC; Garden Club of America Zone VI Fellowship in Urban Forestry from the Casey Tree Endowment Fund; J.E. Weaver Competitive Grant from the Nebraska Chapter of The Nature Conservancy; The College of Liberal Arts and Sciences at Depaul University; Elmore Hadley Award for Research in Ecology and Evolution from the UIC Dept. of Biological Sciences, Spanish CICYT (AMB96-1161; REN2000-0783/GLO; REN2003-05553/GLO; REN2003-03989/GLO; CGL2007-60661/BOS); Yokohama National University; MEXT KAKENHI (25220104); Japan Society for the Promotion of Science KAKENHI (25281053, 17KT0074, 25252026); ADEME (0775C0035); Ministry of Science, Innovation and Universities of Spain (CGL2017-86926-P); Syngenta Philippines; UPSTREAM; LTSER (Val Mazia/Matschertal); Marie Sklodowska Curie Postdoctoral Fellowship (747607); National Science & Technology Base Resource Survey Project of China (2018FY100306); McKnight Foundation (14–168); Program of Fundamental Researches of Presidium of Russian Academy of Sciences (AААА-A18–118021490070–5); Brazilian National Council for Scientific and Technological Development (CNPq 310690/2017–0, 404191/2019–3, 307486/2013–3); French Ministry of Foreign and European Affairs; Bavarian Ministry for Food, Agriculture and Forestry (Project No B62); INRA AIDY project; MIUR PRIN 2008; Idaho Agricultural Experiment Station; Estonian Science Foundation; Ontario Ministry of the Environment, Canada; Russian Science Foundation (16-17-10284); National Natural Science Foundation of China (41371270); Australian Research Council (FT120100463); USDA Forest Service-IITF.