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Belladrilus Righi, 1984 Type species: B. jimi Righi, 1984 Diagnosis: Small Ocnerodrilidae with no dorsal pores, one pair of prostatic pores in xvii, and one of male pores in 17/18 or with seminal grooves in 17/18-xviii. Gizzard present or not in vii, one pair of calciferous glands in ix, testes and male funnels in x. One pair of tubular prostatic glands in xvii, spermathecae without diverticula. Endemic to Latin America, found in Paraguay, Argentina and Brazil, where most (6) of the known species (9) occur (Christoffersen 2008). Belladrilus Righi, 1984 was divided into subgenera Belladrilus (Belladrilus) and Belladrilus (Santomesia) Righi, 1984 on the basis of the structure of the calciferous glands (Righi 1984). The nominate subgenus has glands with a single central cavity, while the other has glands with several approximately longitudinal canals. Based on the calciferous gland structure, the two new species fall in Belladrilus (Belladrilus). Belladrilus (Belladrilus) riparius James, Bartz & Brown, sp. nov. (Fig. 11 a-c) Holotype. BRSP0922, one adult, Boracéia Biological Station of the University of S„o Paulo (USP), S„o Paulo state, Brazil, S23.654003°, W 45.888603°, 840 m asl., in wet margins of a river; 21 April 2004, G.G. Brown and S.W. James, colls. Paratypes. BRSP0202, 19 adults of which 4 were dissected, same data as holotype. Etymology. The species is named for its collection location at the edge of a river. Description. Dimensions 47-62 mm by 1.4-1.5 mm at segment x, 1.3-1.5 mm at clitellum, 1.2-1.3 mm at xxx, body cylindrical, segments 125-134. Setae closely paired throughout, CD above lateral; setal formula AA:AB:BC: CD = 3:1:4:1 at x, 4.5:1:7:1 at xxx, DD Remarks. There are five previously-known species in the nominate subgenus, B. (B.) jimi Righi, 1984, B. (B.) pocaju Righi, 1984, B. (B.) arua Righi, 1984, B. (B.) otarion Righi, 1995, and B. (B.) vaucheri Zicsi, 1995. The new species described here and B. (B.) otarion are distinguished from the others by having spermathecal pores in 7/8, spermathecae in viii, and seminal vesicles in ix and xi. However, the new species differs from that species and all others in the subgenus by having prostatic pore papillae elevated with seminal grooves on the posterior faces of the papillae, nephridia commencing in xii or xiii; prostates extending posteriorly past xxx, and sessile glands associated with the entry of the vas deferens into the body wall in segment xviii. The male pores are in mid- xviii rather than at 17/18 as in B. (B.) otarion, the clitellum of B. (B.) riparius sp. nov. is annular, and it has genital markings in the male field. The following DNA barcodes are from type series specimens: GU013848, GU013849., Published as part of James, Samuel W., Bartz, Marie L. C. & Brown, George G., 2023, New Ocnerodrilidae genera, species and records from Brazil (Annelida: Crassiclitellata), pp. 235-269 in Zootaxa 5255 (1) on pages 252-254, DOI: 10.11646/zootaxa.5255.1.22, http://zenodo.org/record/7744817, {"references":["Righi, G. (1984) 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","Christoffersen, M. L. (2008) A catalogue of the Ocnerodrilidae (Annelida, Oligochaeta) from South America, Italian Journal of Zoology, 75 (1), 97 - 107. https: // doi. org / 10.1080 / 11250000701695250","Righi, G. (1995) A new earthworm (Ocnerodrilidae, Oligochaeta) from a Brazilian cave and considerations about Belladrilus. Revue Suisse de Zoologie, 102, 361 - 365. https: // doi. org / 10.5962 / bhl. part. 80468","Zicsi, A. (1995) Regenw ¸ rmer aus Bolivien (Oligochaeta). Regenw ¸ rmer aus S ¸ damerika, 23. Revue Suisse de Zoologie, 102, 585 - 608. https: // doi. org / 10.5962 / bhl. part. 80475"]}

Diplomoela James, Bartz & Brown, gen. nov. Type species: Diplomoela browni sp. nov. Etymology. The genus name is derived from the double gizzard, using the Portuguese word for gizzard, “moela.” Diagnosis: Setae lumbricine, dorsal pores present or not from 14/15, male pores on xviii, acanthodriline male field with prostatic pores in xvii, xix; female pores paired in xiv, spermathecal pores paired intrasegmental in viii, ix; first setae behind viii. Male sexual system proandric; seminal vesicles multilobed, post-septal in xi; prostates in xvii, xix, ducts muscular. Spermathecae adiverticulate, ectal bulbous muscular duct; ovaries free in xiii. Gizzards large in viii, ix; paired calciferous glands in ix with parallel tubular structure without central lumen; intestinal origin in xiv, no typhlosole; numerous meronephridia commence before gonadal segments. Remarks: Diplomoela has dorsal pores, two large gizzards in the space of viii and ix, and it is meronephric, with 40 to 80 nephridia per segment in mid body. This combination of characters is unique in the family, and it is the only meronephric genus in the family., Published as part of James, Samuel W., Bartz, Marie L. C. & Brown, George G., 2023, New Ocnerodrilidae genera, species and records from Brazil (Annelida: Crassiclitellata), pp. 235-269 in Zootaxa 5255 (1) on page 264, DOI: 10.11646/zootaxa.5255.1.22, http://zenodo.org/record/7744817

Pauqueba uirassu James, Bartz & Brown, sp. nov. (Fig. 14 a-g) Holotype. BRBADNA0325, one adult, Serra Bonita Private Reserve (RPPN Serra Bonita), Camac „, Bahia state, Brazil, S15.38485° W39.57453°, 908 m asl., in bromeliad leaf axils of primary forest; 11 November 2014, M.L.C. Bartz, H. Nadolny, E. da Silva and G.G. Brown, colls. Paratype. BRBADNA0322, adult amputee, Serra Bonita Private Reserve (RPPN Serra Bonita), Camac „, Bahia state, S15.384846°, W39.574535°, in forest floor litter; 11 November 2014, G.G. Brown, coll. Etymology. The species is named for the Instituto Uiraçu, the manager of the reserve at Serra Bonita. The term comes from the Brazilian native indigenous language Tupi, meaning large bird, and is the Tupi name given to the harpy eagle, Harpia harpyja (Linnaeus, 1758), which occurs in the native forest at Serra Bonita. Description. Dimensions> 56-61 mm by 2.7-3.2 mm at segment x, 3.0- 3.5 mm at clitellum, 3.0- 3.4 mm at xxx, body slightly ovate flattened, segments>109, 131+32 regenerates. Setae AB paired, CD unpaired; first setae in iii, setal formula AA:AB:BC:CD:DD = 2.5:1:4:3:5 at x, 3:1:4:3.5:4.5 at xxx. Prostomium broad epilobous open but furrow 1/2 missing, so could be tanylobous from length of prostomial grooves; segments lacking secondary annulations. Deep dorsal chestnut brown pigment in alcohol preservation, bright blue-purple in life; ventral face of head segments brown, AA zone of ventrum otherwise unpigmented. No dorsal pores, spermathecal pores 8/ 9 in A, wide slits with papilla in the opening. Ovipores in xiv in B midway between equator and 13/14; male pores at anterior ends of short seminal grooves in xviii, secondary prostatic pores equatorial in xix (1) or 19/20 (1); seminal grooves in AB but with T-junction at equator of xix or 19/20. Setae AB present in xvii, xix, absent in xviii. Clitellum in xiv-xix, ½ xx, annular; genital markings absent (Fig. 14 a-c). Septa 5/6-12/13 slightly muscular, numerous trans-septal muscles originating near gut in v-xii, inserting on body wall in vi-xiii, respectively; septa and entire peritoneum in anterior segments with red pigment. Alimentary canal with slight muscularity of esophagus in vi, vii but no gizzard; esophagus valvular in xii, intestinal origin in xiii, laterally pouched in xiii-xvi; no typhlosole. Calciferous glands spherical to ovate, paired in ix and x, with honeycomb-like internal structure composed of wide tubes (Fig. 14d, e); glands of ix, x with blood vessel from anterior end to septum 9/10, 10/11 respectively; blood vessel from posterior end of glands to supra-esophageal vessel. Gland attachment at dorso-lateral face of esophagus. Holonephric, first nephridia in segment iv, in intestinal segments small, confined to BC, tubular throughout. Vascular system with ventral trunk, single dorsal trunk, lateral vessels in v-ix, esophageal hearts in x-xi. Supraesophageal vessel in ix-xi. Ovaries, funnels free in xiii; paired large spermathecae in ix, consisting of duct with ectal third or less muscular, internally fluted, bearing sperm masses, duct not clearly marked off from ampulla externally; internally the fluting stops abruptly.Ampulla a flat ovate sac tightly bound to lateral body wall by short muscle or connective tissue fibers (Fig. 14f). Male sexual system holandric, testes and funnels free in x, xi; vas deferens non-muscular, superficial on body wall from 10/11 to xviii where it enters small bulb on body wall; seminal vesicles acinous in xi, xii. Slender tubular prostates coiled twice; once in a tight spiral and secondly the primary coil is wrapped around a band of muscle and mesentery with blood vessels, thus very long glands are densely packed under intestine within xv-xx. Prostatic ducts muscular, widening ectally to union with coelomic muscular copulatory pouch in xix (Fig. 14g).; primary prostatic pore on papilla within pouch. Transverse muscle fibers in xviii-xx crossing male field area. Remarks: comparisons with P. bonita sp. nov. are given under the description of that species. DNA barcodes obtained for this species are in the supplemental information as BRBADNA0322 (paratype), BRBADNA0325 (holotype); and are distinct from the other Pauqueba species. What is apparently a third species of this genus, of similar coloring and size, was represented only by a partially clitellate anterior fragment with a distinctive DNA barcode (Supplemental Information: BRBADNA 0333), collected at the Veracel Station near Porto Seguro (see Silva et al. 2023). This worm has only paired transverse slits on xix for a male field, and internally a coelomic copulatory pouch with the vas deferens terminating on an anterior bulbous appendage of the pouch. Its intestinal nephridia have a dense white cellular mantling from segment xx posteriorly., Published as part of James, Samuel W., Bartz, Marie L. C. & Brown, George G., 2023, New Ocnerodrilidae genera, species and records from Brazil (Annelida: Crassiclitellata), pp. 235-269 in Zootaxa 5255 (1) on pages 258-260, DOI: 10.11646/zootaxa.5255.1.22, http://zenodo.org/record/7744817, {"references":["Silva, E., Santos, A., Nadolny, H. S., Bartz, M. L. C., James, S. W. & Brown, G. G. (2023) Earthworms from natural and managed ecosystems in Southern Bahia, Brazil. Zootaxa, 5255 (1), 270 - 282. https: // doi. org / 10.11646 / zootaxa. 5255.1.23"]}

Kerriona apadaserraensis James, Bartz & Brown, sp. nov. (Fig. 6 a-d) Holotype. BRRS0020, one adult, Aparados da Serra National Park, Cambará do Sul, Rio Grande do Sul state, Brazil, S29.17234°, W50.11646°, 932 m asl, in wetland; 27 January 2012, S.W. James, S.K. Davidson, G.G. Brown and H. Yong, colls. Paratype. BRRS0402, one preclitellate individual, same data as holotype. Etymology. The species name is a contraction of the name of the park where the earthworms were collected: Aparados da Serra. Description. Dimensions 116-118 mm by 3.5-3.6 mm at segment x, 4 mm at clitellum, 3.6-4 mm at xxx, body cylindrical, segments 128-144. Setae closely paired throughout, CD sub-lateral; setal formula AA:AB:BC: CD = 3:1:3.5:1 at x, 3.3:1:4:1 at xxx, DD> ½ circumference. Setae AB of last 20 or so segments enlarged with 90 degree bend, nearly hooked. Prostomium epilobous open; segments lacking secondary annulations. Unpigmented, no dorsal pores, spermathecal pores within 7/8, 8/9 approximately centered in AB; with tumid lips wider than AB. Ovipores in xiv just behind 13/ 14 in B line; male pores equatorial in xviii within seminal grooves; grooves in B line slightly curved and concave laterally; prostatic pores on (or near?) elevated turrets at ends of grooves, lateral to B in xvii-xix; each turret consists of a short column capped with a round flat surface broader than a column, and small crossed furrows on each cap; small pore could be at intersection of these furrows. Setae AB present unmodified in xviii, absent in xvii, xix. Clitellum ½ xiii-xx, saddle in xiv-xx, no other genital markings (Fig. 6a). Anterior septa 5/6 nearly fused into pharyngeal musculature, this musculature in broad bands from iii, iv back to body wall of vii and merge with septum 6/7; septa 6/7/8/9 very strong and elongate, 9/10/11/12 decreasingly muscular; 12/13 and remaining all thin. Alimentary canal lacking gizzard; esophagus elongate, folded in v, shorter but still elongate vi-viii; valvular in xii, intestinal origin in ½ xii; no typhlosole; prominent blood sinuses in intestinal wall in xii-xxii; intestinal walls of xii-xix thicker with longitudinal corrugations and deep segmental pouches. Calciferous glands large, paired in ix, partitioned into multiple lumens by 4 or 5 complete dividing lamina; shorter lamina filling space of each partition (Fig. 6b); each gland with blood vessel from anterior end to extra-esophageal vessel, blood vessel from duct of gland to supra-esophageal vessel. Holonephric, first nephridia in segment viii, tubular throughout from B to above D with upper part above CD apparently a bladder, or thin peritoneal mantle attached to uppermost tubule. Vascular system with ventral trunk, single dorsal trunk in iv-viii, dorsal vessel perforated in ix, forming O-rings in x-xii, xiii; otherwise single; lateral vessels in v-ix, latero-esophageal hearts x-xi. In intestinal segments, small segmental vessels from dorsal vessel to body wall normally with branches within one intestinal segment, but those of xiv receive branches from xiii-xv. Large ovaries, with funnels in xiii; paired spermathecae in viii, ix, consisting of a large muscular duct widening ectally, with 6 inner folds; narrowing conical ampulla smaller in length and volume than duct (Fig. 6c). Male sexual system proandric, testes and funnels in x enclosed in thin membranous sac not composed of septa, vas deferens superficial on body wall from 10/11, joins body wall in xviii where vas deferens terminates in a small muscular bulb; seminal vesicles in ix large laterally placed on body wall, large and post-septal in xi. Slender tubular prostates folded under intestine within xvii-xxxiv, gland diameter about same as ducts; prostatic ducts 2 to 3 segments long, muscular, entering muscular domes of low copulatory bursae; bursal chambers entirely intraparietal containing the bases of the prostatic pore turret bodies. No penial setae. Transverse muscle bands of male field in xvii, xviii, xix with more in the first and third of these segments. Remarks. Compared to other unpigmented Kerriona with turrets in the prostatic pore openings, Kerriona apadaserraensis sp. nov. has the least extensive doubling of the dorsal vessel, because the vessel is not doubled except to a limited extent in segments ix, x-xii, and xiii. The calciferous glands have more (4 or 5 vs. 1 or 2) complete partitions than the other species, and the spermathecal ampullae are smaller than the ducts. These characters distinguish it from other Kerriona. Unfortunately, no DNA barcode data is available for this species.

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"]}

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., 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 421-422, 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","Drumond, C. (1952) Vocabulario na lingua brasilica, vol. 1 (A-H). 2 nd Ed., S \" o Paulo.","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","Erseus, C., Williams, B. W., Horn, K. M., Halanych, K. M., Santos, S. R., James, S. W., des Chatelliers, M. C. & Anderson, F. E. (2020) Phylogenomic analyses reveal a Palaeozoic radiation and support a freshwater origin for clitellate annelids. Zoologica Scripta, 49, 614 - 640. https: // doi. org / 10.1111 / zsc. 12426"]}

Eisenia andrei Bouché BRRSDNA0004, 4 adults and 1 juvenile, compost pile at Federal University of Santa Maria, Santa Maria-RS (S29.718167°, W53.708747°), 21 May 2013, G. Steffen, Z. Antoniolli, colls. BRRSDNA0022, 3 adults, in compost pile, Muçum-RS (S29.14411º, W51.86095º), 21 January 2012, G. Brown, S. James, colls., Published as part of Ferreira, Talita, James, Samuel W., Bartz, Marie Luise Carolina, Lima, Ana Claudia Rodrigues De, Dudas, Rafaela & Brown, George Gardner, 2023, Distribution and diversity of earthworms in different land use systems in Rio Grande do Sul, Brazil, pp. 399-416 in Zootaxa 5255 (1) on page 402, DOI: 10.11646/zootaxa.5255.1.32, http://zenodo.org/record/7745077

Guarani camaqua Rodríguez & Lima BRRSDNA0235, 1 adult, in a wetland by a flowing stream in the Aparados da Serra National Park, Cambará do Sul-RS, (S29.17234°, W50.11646°), 27 January 2012, S. James, G. Brown, S. Davidson, colls., Published as part of Ferreira, Talita, James, Samuel W., Bartz, Marie Luise Carolina, Lima, Ana Claudia Rodrigues De, Dudas, Rafaela & Brown, George Gardner, 2023, Distribution and diversity of earthworms in different land use systems in Rio Grande do Sul, Brazil, pp. 399-416 in Zootaxa 5255 (1) on page 401, DOI: 10.11646/zootaxa.5255.1.32, http://zenodo.org/record/7745077

Eukerria barrigosii James, Bartz & Brown, sp. nov. (Fig. 3 a-c) Holotype. BRMS0115, one adult, Miranda, Mato Grosso do Sul state, Brazil, S20.2614°, W56.4168°, 114 m asl, in flooded rice field; 27 September 2006, J.A. Barrigosi, coll. Paratypes. BRMS0013, one adult, two pre-clitellates, same data as holotype. BRMS0007, one adult, same data as holotype. Etymology. The species is named in honor of its collector, José Antonio Barrigosi, who has sent us several lots of specimens from the region. Description. Dimensions 53-73 mm by 1.2-1.4 mm at segment x, 1.0- 1.2 mm at xxx, 1.2-1.5 mm at clitellum; body cylindrical throughout, segments 84. Setae closely paired throughout, CD sub-lateral; setal formula AA:AB: BC:CD = 4:1:4.5:1 at xxx, DD> ½ circumference. Setae AB present unmodified in xvii-xix. Prostomium epilobous open; segments without secondary annulations. Unpigmented, no dorsal pores, spermathecal pores in 7/8, 8/ 9 in BC just below C. Ovipores in xiv just behind 13/ 14 in B line. Male pores in seminal grooves in xviii; prostatic pores at ends of grooves lateral to B in xvii-xix; seminal grooves, prostatic pores on longitudinal elevated pads aligned about the AB distance lateral to B in xvii-xix. Clitellum xiii- xxii, saddle-shaped, genital markings mid-ventral in BB from equator of ix to equator of x, mid-ventral in AA from equator of xv to 16/17, mid-ventral in AA from equator of xx to equator of xxii, or only to 21/22 (Fig. 3a). Anterior septa 5/6-8/9 thickly muscular, others membranous. Alimentary canal with distinct gizzard in vii; esophagous valvular in xi, intestinal origin ½ xii; no typhlosole, slight intestinal constriction at 21/22. Calciferous glands paired in ix, elongate with ascending duct, attached latero-dorsally to esophagus. Glands with thick wall composed of longitudinal tubules, central lumen present (Fig. 3b); each gland with blood vessel from anterior end to extra-esophageal vessel, blood vessel from posterior face of gland to supra-esophageal vessel. Holonephric, tubular nephridia with dense white mantle filling segmental cavities from A to mid-dorsal, first nephridia in xii. Vascular system with ventral trunk, single dorsal trunk, lateral vessels in vi-ix, large latero-esophageal hearts in x, xi. Fan-shaped ovaries, free with funnels in xiii; paired spermathecae in viii, ix, each an irregular sac ampulla on long slender duct with thicker section at ampulla junction, sperm chambers within this thicker part of the duct, no diverticulum; spermathecae of viii smaller (Fig. 3c). Male sexual system proandric, testes and funnels in x free, vas deferens superficial on body wall from 10/11 to xviii, iridescent and enlarging in xvii-xviii, terminating in small round chamber on body wall of xviii, seminal vesicles in ix with several long stalked branches, placed laterally on body wall, large and post-septal in xi; tubular prostates folded under intestine within xvii-xxiii, xxxv, xxxvii; prostatic ducts ectally enlarged, muscular, no penial setae. Remarks: The new species is geographically and morphologically close to E. subandina (Rosa, 1895) from the same region of Brazil but on the opposite side of the Pantanal (Urucum; Jamieson 1970). Eukerria barrigosii sp. nov. is diagnosed by the saddle-shaped longer clitellum ending on xxii, whereas most others terminate in xix or xx, lack of copulatory bursae at the prostatic pores, spermathecal pores below C in BC, lack of genital markings, spermathecae not in spiral form and with ducts shorter than ampulla, the first muscular septum 5/6, and the first nephridia in xii. Unfortunately, no DNA barcode data is available for this species., Published as part of James, Samuel W., Bartz, Marie L. C. & Brown, George G., 2023, New Ocnerodrilidae genera, species and records from Brazil (Annelida: Crassiclitellata), pp. 235-269 in Zootaxa 5255 (1) on pages 240-241, DOI: 10.11646/zootaxa.5255.1.22, http://zenodo.org/record/7744817, {"references":["Rosa, D. (1895) Viaggio del dottor Alfredo Borelli nella Republica Argentina e nel Paraguay - Oligocheti terricoli. Bolletino dei Musei di Zoologia ed Anatomia comparata della Universita di Torino, 10 (204), 1 - 3. https: // doi. org / 10.5962 / bhl. part. 8048","Jamieson, B. M. (1970) A taxonomic revision of the oligochaete genus Eukerria Michaelsen, 1935 (Ocnerodrilinae, Megascolecidae). Bulletin of the British Museum (Natural History), Zoology, 20, 133 - 172."]}

Perionyx excavatus Perrier BRRSDNA0020, 2 adults, in compost pile, Muçum-RS (S29.14411°, W51.86095°), 21 January 2012, G. Brown, S. James, colls., Published as part of Ferreira, Talita, James, Samuel W., Bartz, Marie Luise Carolina, Lima, Ana Claudia Rodrigues De, Dudas, Rafaela & Brown, George Gardner, 2023, Distribution and diversity of earthworms in different land use systems in Rio Grande do Sul, Brazil, pp. 399-416 in Zootaxa 5255 (1) on page 402, DOI: 10.11646/zootaxa.5255.1.32, http://zenodo.org/record/7745077

Parabauba veracelensis James, Bartz & Brown, sp. nov. (Fig. 15 a-d) Holotype. BRBADNA0340, late juvenile, Estaç „o Veracel Private Reserve (RPPN Estaç„o Veracel), Porto Seguro, Bahia state, Brazil, S16.38724 ˚ W39.16957 ˚, 90 m asl., 30 cm deep in mineral soil of primary forest; 14 November 2014, M.L.C. Bartz, H. Nadolny, E. da Silva and G.G. Brown, colls. Paratype s: BRBADNA336, 337, 338, 339, 341, five late juveniles, same data as holotype, one in each vial. Etymology. The species is named for the type locality (RPPN Estaç„o Veracel), and the Veracel (Veracruz Celulose) company, the owner and manager of the reserve. Description. Dimensions 40-55 mm by 2 mm at segment x, 2 mm at xxx, body cylindrical, segments 130-153; last 20 to 30 segments tapering and shortening. Setae closely paired; first setae in ii, setal formula AA:AB:BC:CD: DD = 5:1:10:1:32 at xxx. Prostomium epilobous open; segments lacking secondary annulations. Unpigmented, no dorsal pores, spermathecal pores intersegmental 7/8/ 9 in AB, surrounded by oval lip. Ovipores not observed; male field acanthodriline, slightly depressed; prostatic pores equatorial in xvii, xix on transverse oval papillae; male pores in xviii in seminal grooves. Setae AB in xvii, xviii, xix not seen. Clitellum not developed; genital markings absent (Fig. 15a, b). Septa 5/6-9/10 strongly muscular, 10/11/12/13 slightly muscular. Alimentary canal with long well-developed gizzard in v; esophagus valvular in xii, intestinal origin in xiii (Fig. 15c, d); no typhlosole. Calciferous glands ovate oriented vertically, paired in ix and x, no central lumen, parallel large tubules or lamellae converging near duct; glands of ix with blood vessel from anterior end to extra-esophageal vessel, and blood vessel from posterior end of gland to supra-esophageal vessel; glands of x with blood vessels through duct of gland. Gland attachment at dorsolateral face of esophagus. Holonephric, first nephridia in segment v, in intestinal segments small, confined to BC, tubular throughout. Vascular system with ventral trunk, single dorsal trunk, lateral vessels in vii-ix, latero-esophageal hearts in x-xi. Supraesophageal vessel in ix-xi. Ovaries, funnels free in xiii; paired adiverticulate spermathecae in viii, ix, duct muscular with ectal bulb at body wall, tapering entally towards ampulla; ampulla ovate (Fig. 15c). Male sexual system holandric, testes and funnels free in x, xi; vas deferens superficial on body wall from 10/11 to xviii where it enters body wall; seminal vesicles acinous in xi, xii. Slender tubular prostates coiled under or alongside intestine within xiii-xx (Fig. 15d). Prostatic ducts long, muscular, of constant diameter. Remarks. So far, the only described species in the genus, it is distinguished from all other Ocnerodrilidae by the diagnostic characters of the genus. Unlike the other Ocnerodrilidae examined for this publication, the calciferous glands contained abundant white powder, presumed to be calcium carbonate granules. DNA barcode data for this species was not obtained. A different, still undescribed species of Parabauba was found by Silva et al. (2023) at the Una Biological Reserve, around 120 km (in straight line) from the Veracel station.

Genus Ocnerodrilus Eisen, 1878 Type species: Ocnerodrilus occidentalis Eisen, 1878 Diagnosis: The diagnosis of Righi (1994) is still valid after the description of the present species. It is summarized as follows: setae lumbricine, dorsal pores absent, biprostatic or quadriprostatic, with male pores near prostatic pores in 17/18 or joined to prostatic pores in xvii; Holandric, metagynic, adiverticulate spermathecae in ix. Gizzard absent, calciferous glands paired in ix with a central cavity surrounded by lengthwise lamellae of varying depths. Remarks: Spermathecae, male organs, including prostates and male pores may be absent in some species with parthenogenetic reductions. Some species have smooth inner walls of the calciferous glands. There are 29 nominal species of Ocnerodrilus, at least 12 of which are probably synonyms of O. occidentalis (Gates, 1973; Blakemore 2006): O. beddardi Eisen, 1893, O. guatemalae Eisen, 1893, O. hendriei Eisen, 1893, O. sonorae Eisen, 1893, O. calwoodi Michaelsen, 1898, O. occidentalis “ var. arizonae ” Eisen, 1900, O. mexicanus Eisen, 1900, O. comundui Eisen, 1900, O. sinensis Eisen, 1900, O. tenellulus Gates, 1945, and O. asiaticus Chen & Hsu, 1975. Gates (1973) further considered that O. sabanae Cognetti, 1904 should be considered species incertae sedis because the calciferous glands are not adequately characterized, and O. calwoodi should be placed in synonymy because it might represent the sexual lineage from which the asexual lineage(s) may have been derived. However, that was based on then-current concepts about species status and the appropriateness of naming parthenogenetic groups. Ideas have since changed and in consideration of the morphologically distinct status of O. calwoodi, it should be removed from synonymy. Fragoso & Brown (2007) also called attention to the need for further revision of Ocnerodrilus and closely related genera, considering the discovery of several new species found by Righi (1984) and Righi & FraileMerino (1987), as well as the need for additional sampling, particularly in Central America, where the supposed amphimictic population of O. occidentalis may be found., Published as part of James, Samuel W., Bartz, Marie L. C. & Brown, George G., 2023, New Ocnerodrilidae genera, species and records from Brazil (Annelida: Crassiclitellata), pp. 235-269 in Zootaxa 5255 (1) on page 262, DOI: 10.11646/zootaxa.5255.1.22, http://zenodo.org/record/7744817, {"references":["Righi, G. (1994) On new and old-known Oligochaeta genera from Parai´ba State, Brazil. Revue Suisse de Zoologie, 101, 89 - 106. https: // doi. org / 10.5962 / bhl. part. 79901","Gates, G. E. (1973) Contributions to North American earthworms (Annelida) No. 7. Contributions to a revision of the earthworm family Ocnerodrilidae IX. What is Ocnerodrilus occidentalis? Bulletin of the Tall Timbers Research Station, 14, 13 - 28.","Blakemore, R. J. (2006) Cosmopolitan earthworms - an eco-taxonomic guide to the peregrine species of the world. Kippax, Australia: VermEcology.","Righi, G. (1984) 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"]}

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"]}

Eukerria ibicatuensis James, Bartz & Brown, sp. nov. (Fig. 2a, b) Holotype. BRPR1851, one adult, Ibicatú State Park, Ibicatú, Paraná state, Brazil, S22.786°, W51.488°, 331 m asl, in wet seepage area near small river; 29 April 2004, G. Brown and S. James, colls. Paratypes. BRPR0555, two adults, one of them amputated, same data as holotype. Other material. BRPR1962, one subadult, two juveniles, same data as holotype. Etymology. The species name is based on the location of the collection in the Paraná state park of Ibicatú. Description. Dimensions 68-81 mm by 2-2.3 mm at segment x, 1.9-2.4 mm at xxx, 2.3-2.6 mm at clitellum; body cylindrical throughout, segments 160, 188 (of which 18 appear to be regenerated). Setae closely paired throughout, CD sub-lateral; setal formula AA:AB:BC:CD = 4.5:1.2:4:1 at xxx, DD> ½ circumference. Setae A present unmodified in xvii, xix; AB lacking in xviii; B lacking in xvii, xix. Prostomium epilobous open; segments without secondary annulations. Unpigmented, no dorsal pores, spermathecal pores large in 7/8, 8/ 9 in CD. Ovipores in xiv faint slits just behind 13/ 14 in B line. Male pores in seminal grooves in xviii; prostatic pores at ends of grooves slightly lateral to B in xvii-xix; prostatic porophores bearing small cleft papillae at ends of seminal grooves. Seminal grooves, prostatic pores on longitudinal elevated pads aligned lateral to ½ AB in xvii-xix; pads slightly constricted at xviii equator. Clitellum xiii-xxi, saddle-shaped, only present dorsally in xii, xxi, no other genital markings (Fig. 2a). Anterior septa 5/6-8/9 thickly muscular, 9/10-11/12 less so. Alimentary canal with gizzard in vii; esophagous valvular in xi, intestinal origin in xii; no typhlosole or other differentiation of intestine. Calciferous glands blunt sagittate with ascending duct, paired in ix, attached latero-dorsally to esophagus. Glands with thick wall, irregular central lumen formed by longitudinal, radial projections of glandular tissue towards the center (Fig 2b); each gland with blood vessel from anterior end to extra-esophageal vessel, blood vessel from posterior face of gland to supraesophageal vessel. Holonephric, tubular nephridia bound within membranous covering to form flat, transparent ribbon, first nephridia in xii. Vascular system with ventral trunk, single dorsal trunk, lateral vessels in v-ix, latero-esophageal hearts in x, xi. Fan-shaped ovaries, with funnels in xiii; paired spermathecae in viii, ix, each an irregular sac ampulla on long broad duct, no diverticulum; sperm chambers visible within duct near body wall. Male sexual system proandric, testes and funnels in x free, vas deferens superficial on body wall from 10/11 to xviii terminating without enlargement, seminal vesicles in ix with several long stalked branches, placed laterally on body wall, large and post-septal in xi; tubular prostates folded under intestine within xviii-xxi; prostatic ducts ectally enlarged, muscular, enter dome-shaped circular copulatory bursae, no penial setae. Remarks. Eukerria contains two taxa in which the prostatic porophores merge antero-posteriorly to form a longitudinal bar, rather than the situation in which the porophores of a segment encroach towards one another to form a merged, or nearly merged, transverse bar across each of xvii and xix. Eukerria garmani garmani (Rosa, 1898) and E. g. argentinae Jamieson, 1970 are these two other taxa which have the longitudinal merger. Compared to the former subspecies, E. ibicatuensis sp. nov. lacks genital markings in xiii, has a larger gizzard, has more dorsally located spermathecal pores (CD vs. mid-BC), the calciferous glands are dorso-laterally attached to the esophagus (vs. ventro-laterally), the seminal vesicles of xi are larger than those of ix (vs. equal), prostates are confined to xviiixxi, prostate ducts enter small bursae (vs. not), and the spermathecal ducts are longer than the ampulla (vs. shorter). The differences from E. g. argentinae are fewer and as follows: the first nephridia are in xii (vs. xi), and there are distinct seminal grooves rather than a very glandular porophore and male field area marked by numerous transverse and longitudinal folds. Other species are excluded by the structure of the calciferous glands and positions and/or forms of genital pores. Eukerria cuca has similarly placed spermathecal pores but they are accompanied by genital markings and internal genital marking glands; it also has a simpler lumen of the calciferous glands, shorter clitellum and lesser male field development. DNA barcodes for this species are GU013976 and GU013977. Calculations based on the K2P 14% criterion (conservative) and the ASAP algorithm (Puillandre et al. 2020) both separated this species from all other known ocnerodrilid barcode sequences., Published as part of James, Samuel W., Bartz, Marie L. C. & Brown, George G., 2023, New Ocnerodrilidae genera, species and records from Brazil (Annelida: Crassiclitellata), pp. 235-269 in Zootaxa 5255 (1) on pages 238-239, DOI: 10.11646/zootaxa.5255.1.22, http://zenodo.org/record/7744817, {"references":["Jamieson, B. M. (1970) A taxonomic revision of the oligochaete genus Eukerria Michaelsen, 1935 (Ocnerodrilinae, Megascolecidae). Bulletin of the British Museum (Natural History), Zoology, 20, 133 - 172.","Puillandre, N., Brouillet, S. & Achaz, G. (2020) ASAP: assemble species by automatic partitioning. Molecular Ecology Resources, 21 (2), 609 - 620. https: // doi. org / 10.1111 / 1755 - 0998.13281"]}

Eukerria adecarvalhoi James, Bartz & Brown, sp. nov. (Fig. 1 a-c) Holotype. BRSE0014, one adult, Fazenda Haras Chamusca, Lagarto, Sergipe state, Brazil, S11.033611°, W37.648528°, 137 m asl., degraded area on the banks of a small stream; 25 May 2012, André de Carvalho coll. Paratypes. BRSE0015, two adults, same data as holotype. Other material. BRSE0006, 8 adults, 4 juveniles, same data as holotype. BRSE0010, 4 juveniles and 2 adults, same data as holotype. Etymology. The species is named after the collector of most of the specimens, by contracting his first and family names. Description. Dimensions 35-53 mm by 1.3-1.7 mm at segment x, 1.1-1.6 mm at xxx, 1-1.5 mm at clitellum; body cylindrical throughout, segments 110-140. Setae paired throughout; setal formula AA:AB:BC:CD = 2.5:1:4:1.3 at xxx, DD Septa 6/7-11/12 thin but not membranous. Alimentary canal with variable degree of pharyngeal muscles in iv, v; weak barrel-shaped gizzard in vii; esophagous valvular in xi, intestinal origin xii; no typhlosole or other differentiation of intestine. Calciferous glands ovate, paired in ix, composed of radial lamellae with very small central lumen (Fig. 1b); each gland with blood vessel from anterior end to extra-esophageal vessel, blood vessel from duct of gland to esophageal wall becoming supra-esophageal vessel. Holonephric, tubular nephridia lacking bladders, present from segment xi or xii, occupying space from mid-BC ventrally. Vascular system with ventral trunk, dorsal trunk, these connected by lateral vessels in vii, viii; lateral hearts in ix-xi; dorsal trunk in xi doubled with each side connecting to lateral heart of that segment. Ovaries, with funnels in xiii; paired spermathecae in viii, ix, each an elongate sac or with pronounced swelling at ampulla end, no diverticulum; sperm chambers visible within duct near body wall (Fig. 1c). Male sexual system proandric, testes and funnels free in x, vas deferens superficial on body wall from 10/11 to xviii; seminal vesicles small, parietal in ix, large post-septal in xi (Fig. 1c); tubular prostates with very long glandular portion folded under and along sides of intestine within xvi-xx, xxii; prostatic ducts more muscular at body wall, generally slender and long, gradually narrowing towards junction with glandular part. No penial setae. Remarks. Of the 32 nominal species, some are from far-flung locations, and might be synonyms of the peregrines E. kuekenthali (Michaelsen, 1908) or E. saltensis (Beddard, 1895). In the former group Blakemore (2006) includes E. peguana Gates, 1942 from Myanmar and E. selangorensis (Stephenson, 1931) from Malaysia. Junior synonyms of E. saltensis include at least E. gunningi Michaelsen, 1913, E. nichollsi Jackson, 1931 and E. sydneyensis Jackson, 1931 (Blakemore, 2006). Eukerria. hortensis (Stephenson, 1931), E. limosa (Stephenson, 1931), E. pascuorum (Stephenson, 1931) and E. rubra (Friend, 1916) may also be synonyms of these or other species, e.g., E. eiseniana (Rosa, 1895), but have not been adequately evaluated (Blakemore 2006). Species described after the publication of Righi (1968), which includes a key to all the known species of Eukerria (at that time) are the following: E. cuca Righi, 1984, E. emete Righi & Guerra, 1985, E. guamais Righi, 1971, E. mucu Righi, 1988, E. taisa Righi, 1983, and E. santafesina Lungstr ̂m, 1972. All of these publications were consulted in diagnosing the new species of Eukerria described here. The widely-distributed peregrine E. saltensis is epilobous, has an annular clitellum, is much more slenderbodied, the spermathecal pores are more ventral, the male field grooves are straighter and lack the oval genital markings at the ends of the grooves, as well as lacking the deep pits containing the male pores. Its calciferous gland is usually described as a solid mass with a small central lumen, and only by Gates (1942) as having radial lamellae extending into a large lumen. The same external characters distinguish E. adecarvalhoi sp. nov. from E. kuekenthali. The spermathecal pores of E. eiseniana are above the level of setal row D, its seminal grooves near the level of C, and there is no evidence of sperm production. However it appears to have radial lamellae in the calciferous glands. The present species is diagnosed by the following combination of character states: clitellum saddle shaped ending in xx, calciferous glands with parallel lamellae projecting radially inwards from the gland wall (cf. Righi 1968), spermathecal pores at the C setal row level, lacking genital markings, 2 pairs of prostate glands, prostate glands opening with copulatory bursae (interpreting the deep pits as such), double dorsal vessel in xi, and spermathecal duct shorter than the ampulla. In the new species the calciferous glands’ radial lamellae appear to meet, or almost meet, in the center of the gland, but it is difficult to tell if the sometime appearance of a small lumen is an artifact of making sections by hand, or a real feature of the gland. The specimens from BRSE006 were slightly larger and had less muscularity in the prostatic ducts but more in the pharynx area, compared to those of BRSE0010. External characters are in complete agreement between the two sites. Unfortunately, no DNA barcode data is available for this species.

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"]}

Kerriona davidsonae James, Bartz & Brown, sp. nov. (Fig. 4 a-c) Holotype. BRSP0139, late aclitellate juvenile, Intervales State Park, Itapeúna, Eldorado county, S„o Paulo state, Brazil, S24.305°, W48.3651°, 818 m asl., in bromeliad leaf axils; 7 February 2009, S.W. James, S.K. Davidson, M.L.C. Bartz, and G.G. Brown, colls. Paratype. BRSP0919, late aclitellate juvenile, same data as holotype. Etymology. The species is named after our colleague and collaborator Seana Davidson, a pioneer in researching the bacterial symbionts living in earthworm nephridia. Description. Dimensions 60-69 mm by 2.5-3.0 mm at segment x, 3.3 mm at xxx, 151 segments; body cylindrical throughout, posterior gradually tapered to tail, posterior-most segments shorter. Setae paired throughout, CD sub-lateral; setal formula AA:AB:BC:CD = 2.4:1:3:1.2 at xxx, DD> ½ circumference. Prostomium epilobous open; segments without secondary annulations. Unpigmented, no dorsal pores, spermathecal pores in 7/8, 8/9 at B. Ovipores not seen; male pores in xviii just lateral to B; prostatic pores adjacent to elevated penial bodies at ends of grooves in B in xvii-xix; each penial body consists of a short column capped with an ovate flat surface broader than a column, and a small rounded triangular notch cut out of the posterior margin of each cap (Fig. 4 a,b). Setae AB present unmodified in xvii-xix. Clitellum not developed, no other genital markings. Anterior septa thin but not membranous. Alimentary canal with strong pharyngeal muscles in iv-vii (Fig. 4c); lacking gizzard; esophagous valvular in xi, intestinal origin in xii; no typhlosole; intestine with lateral pouches in xiii-xx. Calciferous glands elongate an ovate, very large, paired in ix, densely packed with longitudinal partitions; each gland with blood vessel from anterior end to extra-esophageal vessel, blood vessel from posterior face of gland to esophageal wall and supra-esophageal vessel. Holonephric, tubular nephridia without bladders; beginning in vii or viii (Fig. 4c). Vascular system with ventral trunk, single dorsal trunk iv-ix, double dorsal vessel x and posteriorly (Fig. 4c), lateral vessels in vii, viii, lateral hearts in ix-xi. Ovaries, funnels in xiii; paired spermathecae in viii, ix, immature and very small (Fig. 4c). Male sexual system proandric, testes and funnels in x enclosed in sac made by fusion of septa 9/10/11, vas deferens superficial on body wall from 10/11 to enlarged muscular end at body wall in xviii; seminal vesicles in ix small laterally placed on body wall, large and post-septal in xi; slender tubular prostates folded under intestine within xvii-xxii; prostatic ducts slender, enter 0.7 mm diameter muscular domes. No penial setae. Remarks. The proandric testes enclosed in sacs is a feature of all the unpigmented Kerriona species we report on here, plus K. limae. This character excludes K. garbei, K. luederwaldti, and K. bocainensis sp. nov. in which the testes are free, dorsal vessels single, and prostatic pores unaccompanied by turrets. Kerriona garbei differs from all other Kerriona known previously or described here by having a single pair of spermathecae, and by its much larger body size. The closest relationship of K. davidsonae sp. nov. is with those species bearing the doubled dorsal vessel and small turrets at the prostatic pores, of which we describe several in the following pages. Although the specimens of K. davidsonae sp. nov. are juveniles, sperm has matured and gathered on the funnels so maturity is close at hand. They differ from K. riodaonsa sp. nov. by having more posterior start of doubling of the dorsal vessel, more widely paired setae, thicker anterior septa, a pouched intestine in xii-xix, AB setae present in all male field segments, and a muscular termination of the vas deferens. Another possible distinction is that the muscular surroundings of the prostatic duct terminations are clearly coelomic in this species and more intraparietal in K. riodaonsa sp. nov., despite the latter being more fully mature. Kerriona davidsonae sp. nov. has many more lamellae in the calciferous glands and a more extensively doubled dorsal vessel than K. apadaserraensis sp. nov., K. vilavelhaensis sp. nov., and K. clivedwardsi sp. nov. It differs from K. limae by having more widely paired setae, calciferous glands more densely packed with lamellae, and muscular copulatory pouches around the prostatic pores. DNA barcodes: HQ562639, HQ562640, HQ562641. These are separated from all other Kerriona DNA barcodes as a species-level taxon by ASAP (Puillandre et al. 2020) and the 14% criterion., Published as part of James, Samuel W., Bartz, Marie L. C. & Brown, George G., 2023, New Ocnerodrilidae genera, species and records from Brazil (Annelida: Crassiclitellata), pp. 235-269 in Zootaxa 5255 (1) on pages 241-242, DOI: 10.11646/zootaxa.5255.1.22, http://zenodo.org/record/7744817, {"references":["Puillandre, N., Brouillet, S. & Achaz, G. (2020) ASAP: assemble species by automatic partitioning. Molecular Ecology Resources, 21 (2), 609 - 620. https: // doi. org / 10.1111 / 1755 - 0998.13281"]}

Ocnerodrilus imperialis James, Bartz & Brown, sp. nov. (Fig. 16 a-d) Holotype. BRRJDNA0059, one adult, Tinguá Biological Reserve, Rio de Janeiro state, Brazil, S22.57007 ˚ W43.46864 ˚, 795 m asl., in wet stream bank sediment; 02 April 2014, G.G. Brown, S.W. James, A.C. Buch, L.F. Lourençato, and M.L.C. Bartz, colls. Paratype. BRRJ0012, 1 adult, 6 juveniles, same data as holotype. Etymology. The species is named for the Estrada Imperial, the old road from the former Brazilian imperial capital in Rio de Janeiro city to the summer location of the imperial court in Teresopolis Petrópolis. Description. Dimensions 38-56 mm by 1.3-1.4 mm at segment x, 1.2-1.3 mm at clitellum, 1.4-1.5 mm at xxx, body cylindrical, segments 84-97. Setae closely paired throughout, CD below lateral; setal formula AA:AB:BC:CD = 4:1:4:1 at x, 3.3:1:4:1 at xxx, DD> ½ circumference. Prostomium epilobous open; segments lacking secondary annulations, unpigmented, no dorsal pores, spermathecal pores wide slits with tumid lips in 8/9, centered on B. Ovipores transverse slits in xiv in B just behind 13/14; male and prostatic pores with common secondary opening in transverse slit openings within thickened porophores on xvii. Primary male and prostatic pores in xvii on floor of shallow invagination. Setae AB absent in xvii. Clitellum in ½xiii, xiv-xviii, saddle shaped reaching down to A setal line; genital markings absent (Fig. 16a, b). Septa 5/6-9/10 muscular, 11/12/13 less so. Alimentary canal without gizzard; esophageal glands in v-viii, esophagus with long valve in xii-xiii, intestinal origin in xiv, no typhlosole. Calciferous glands oblong, paired in ix, dorso-laterally connected to esophagus, with solid walls surrounding irregular central lumen (Fig. 16c); each gland with blood vessel from anterior end to extra-esophageal vessel, and blood vessel from posterior end of gland to supra-esophageal vessel. Holonephric, first nephridia in segment v, tubular throughout within BC-D level of segments. Vascular system with ventral trunk, single dorsal trunk, lateral vessels in v-ix, latero-esophageal hearts in x-xi. Ovaries and funnels free in xiii; paired spermathecae in ix, consisting of short narrow muscular duct, sharply demarcated ovate ampulla (Fig. 16c). Male sexual system holandric, testes and funnels enclosed in very delicate membranous sacs in x, xi, vas deferens superficial on body wall from 10/11 to xvii where it enters body wall at anterior edge of prostatic duct; seminal vesicles in xii extending into xiii, seminal vesicles with stalked attachment to posterior face of 11/12. Tubular prostates folded under intestine within xii-xix; prostatic ducts slightly muscular, about one segment long; glandular portions three times the diameter of the ducts (Fig. 16d), no copulatory bursae, or other thickened ends of prostatic ducts or vasa deferentia. Transverse muscle bands in xvii-xix. Remarks. Ocnerodrilus imperialis sp. nov., unlike the most common species O. occidentalis Eisen, 1878 is fully hermaphroditic with male funnel iridescence and large volumes of sperm enclosed in segments x and xi. Gates (1973) characterized the entire genus as having an intestinal origin in xii, but Righi (1994) does not include that character in his diagnosis of Ocnerodrilus. The present species has a distinct, long esophageal valve in xii and xiii, with the intestinal origin in xiv. Therefore, one can dismiss all species known to Gates as being senior names of O. imperialis sp. nov. That leaves 5 species, all described by Righi and a collaborator, from South American and Costa Rican specimens: O. alox Righi & Merino, 1987, O. andinus Righi, 1981, O. ibemi Righi, 1968, O. ipu Righi, 1975, and O. potyuara Righi, 1994. The new species is distinct from O. alox, another apparently amphimictic species, which has an intestinal origin in xii, spermathecal pores in 7/8 and seminal vesicles in ix and xii. Ocnerodrilus andinus is very small (10 mm vs. 41-56), the intestinal origin is in ½xii, and the calciferous gland walls are very simple, without blood sinuses and longitudinal structure. The male field of O. ibemi consists of a roughly hexagonal elevated glandular area around the male/prostatic pores, which is not present in O. imperialis sp. nov .. The two also differ in the intestinal origin. In Righi’s (1994) key, O. ibemi is said to have internal longitudinal folds or trabeculae in the calciferous glands; this character is not described in Righi (1968). Such folding is minimal if present, in O. imperialis sp. nov.. A darkly pigmented species O. ipu has two pairs of spermathecae and seminal vesicles in ix and xii, among other differences from O. imperialis sp. nov.. Compared to O. potyuara, the new species lacks seminal vesicles in ix, the seminal vesicles are stalked, not sessile on septum 11/12; it has a more posterior intestinal origin, the male and prostatic porophores have pronounced slits rather than being rounded papillae, and the prostates are folded under the gut anterior to the prostatic segments. A nomenclatural note: the name is spelled “potyura” in the abstract (p. 89) but “potyuara” elsewhere, including a species list for the genus (p. 93), several points about morphological diversity within Ocnerodrilus (p. 94), a key to the species of Ocnerodrilus (p. 96), the species description itself (p. 96), figure legend (p. 97), and remarks on the species and comparisons to others (p. 98) (Righi, 1994). The DNA barcode for the holotype is in the Supplemental Information: BRRJDNA0059.

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"]}

Rhinodrilus sp. nov. BRBA0009 (BRBADNA0330), 1 adult minhocuçu in aestivation (Figure 1A); at 60 cm depth in native forest at the Estaç „o Ecológica Pau Brasil, Porto Seguro, BA (S16.38596 W39.17433); 113 m a.s.l., 13-11-2014, M.L.C. Bartz and G.G. Brown, colls. BRBA0012, 1 adult minhocuçu; found dead on the side of the road after being run over by a motorcycle (Figure 1B); the specimen had likely wandered onto the road in the early morning after a rain-shower, from the native forest on the roadside at the RPPN Estaç„o Veracel, Porto Seguro, BA (S16.387653 W39.170093); 90 m a.s.l., 14-11-2014, G.G. Brown, coll. BRBA0013 (BRBADNA0345), 1 juvenile; topsoil (0– 20cm) of native forest at the RPPN Estaç „o Veracel, Porto Seguro, BA (S16.38724 W39.16957); 90 m a.s.l., 14-11- 2014, M.L.C. Bartz and G.G. Brown, colls.

Andiorrhinus (Amazonidrilus) duseni (Michaelsen) BRRS0403, one adult in the surface soil horizon of pasture next to secondary vegetation, Machadinho-RS (S27.61914°, W51.73108°), 17 August 2016, W. Maschio, coll., Published as part of Ferreira, Talita, James, Samuel W., Bartz, Marie Luise Carolina, Lima, Ana Claudia Rodrigues De, Dudas, Rafaela & Brown, George Gardner, 2023, Distribution and diversity of earthworms in different land use systems in Rio Grande do Sul, Brazil, pp. 399-416 in Zootaxa 5255 (1) on page 402, DOI: 10.11646/zootaxa.5255.1.32, http://zenodo.org/record/7745077

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."]}

Kerriona vilavelhaensis James, Bartz & Brown, sp. nov. (Fig. 7 a-c) Holotype. BRPR0558, adult, Vila Velha State Park, Ponta Grossa, Paraná state, Brazil, S25.2519°, W50.0158°, 801 m asl., in marshy soil in a wetland; 31 January 2006, S.W. James, S.K. Davidson, G.G. Brown, N. Pinel, T. Decaëns, colls. Paratype. BRPR1853, one adult, same data as holotype. Etymology. This species is named after the type locality, the state park of Vila Velha. Description. Dimensions 37-45 mm by 1.5-1.6 mm at segment x, 1.6-1.8 mm at clitellum, 1.6-1.8 mm at xxx, body cylindrical, tapering in posterior third to narrow conical pygidium; final 20 or more segments much shorter; segments 127, 92 (80 plus 12 regenerates). Setae closely paired throughout, CD sub-lateral; setal formula AA:AB: BC:CD = 5:1:4.5:1.3 at x, 4.5:1:4:1.3 at xxx, DD> ½ circumference. Prostomium epilobous open; segments lacking secondary annulations. Unpigmented, no dorsal pores, spermathecal pores within 7/8, 8/9 at B. Ovipores in xiv just behind 13/14 lateral to B line; male pores in xviii within seminal grooves in line with B; secondary prostatic pores deep transverse slits at ends of seminal grooves in xvii, xix; grooves in B ½ xvii-½ xix. Small turrets within the prostatic pore invaginations. Setae AB present, unmodified in xviii, absent in xvii, xix. Clitellum xiv-½xx, saddle shape, slight mid-ventral elevations in xvi, xx (Fig. 7a). Anterior septa 5/6-9/10 lightly muscular. Alimentary canal lacking gizzard but esophagus faintly muscular in vii, but not enlarged compared to adjacent segments; esophagus valvular in xi, intestinal origin in xii; no typhlosole. Calciferous glands ovate, small and paired in ix (Fig. 7b), attached ventro-laterally to esophagus, with at least one longitudinal partition, and 7-8 thick lower ridges filling space of gland, but separate; each gland with blood vessel from anterior end to extra-esophageal vessel, and blood vessel from posterior face of gland to supra-esophageal vessel. Holonephric, first nephridia in segment vi, tubular throughout with apparently dorsal bladder, or thin peritoneal mantle attached to uppermost tubule bordering the possible bladder. Vascular system with ventral trunk, single dorsal trunk in iv-ix, dorsal vessel double in vii-xvi or xi-xviii, perforations in dorsal vessel from xvii or xix-xxii or scattered to xxx; lateral vessels in iv-ix, esophageal hearts in x-xi. Fan-shaped ovaries, with funnels in xiii; paired spermathecae in viii, ix, consisting of cylindrical muscular duct containing sperm chambers; ovate ampulla well-demarcated from duct (Fig. 7b). Male sexual system proandric, testes and funnels in x free, vas deferens superficial on body wall from 10/11, terminates in small bulb in xviii; seminal vesicles in ix large acinous laterally, placed on body wall, small and post-septal in xi. Large tubular prostates folded under intestine within xv, xvii-xxiv, xxvii; prostatic ducts slender, muscular, 2 or 3 segments long, enter muscular copulatory bursal domes in xvii, xix (Fig. 7c). Intra-coelomic bursae contain two wedge-shaped turrets, the posterior of which connected to a short low wall curving anteriorly. No penial setae. Remarks: In this species the copulatory bursae are clearly not only intra-parietal but are prominent in the body cavity, unlike the other Kerriona. The seminal vesicles of ix are larger than those of xi, the reverse of the common situation and that of all other known Kerriona, except for K. clivedwardsi sp. nov., which lacks seminal vesicles in ix. Calciferous gland lamellae are thicker and fewer in number than those of other Kerriona, and the extent of doubled or perforated dorsal vessel is within the range of vii to xxx. There are no DNA barcode data for this species due to repeated failures of PCR and sequencing., Published as part of James, Samuel W., Bartz, Marie L. C. & Brown, George G., 2023, New Ocnerodrilidae genera, species and records from Brazil (Annelida: Crassiclitellata), pp. 235-269 in Zootaxa 5255 (1) on page 246, DOI: 10.11646/zootaxa.5255.1.22, http://zenodo.org/record/7744817

Belladrilus (Belladrilus) browni James & Bartz sp. nov. (Fig. 12 a-c) Holotype. BRRS0054, one adult, Aparados da Serra National Park, Cambará do Sul, Rio Grande do Sul state, Brazil, S29.17234°, W50.11646°, 932 m asl., in wetland adjacent to forest; 27 January 2012, S.K. Davidson, G.G. Brown, H. Yong, and S.W. James, colls. Paratypes. BRRSDNA0245-0247, three adults, same data as holotype, one in each vial. Etymology. The species is named for George G. Brown, a co-discoverer of the worm and a leader of earthworm research in Brazil. Description. Dimensions 45-50 mm by 1.0 mm at segment x, 0.8 mm at clitellum, 0.9 mm at xxx, body cylindrical, segments 120-139. Setae closely paired throughout, CD above lateral; setal formula AA:AB:BC:CD = 4:1:5:1 at x, 5:1:5:1 at xxx, DD Remarks. Belladrilus (B.) browni sp. nov. has the largest and most elevated male pore zone of any known species in the subgenus. Like B. (B.) riparius sp. nov., it has more posterior first nephridia than the older species. On the other hand it shares the character state of seminal vesicles only in xi and spermathecae in ix with the following species: B. (B.) jimi, B. (B.) pocaju, B. (B.) arua, and B. (B.) vaucheri. With spermathecal pores in line C and differentiated spermathecal ducts vs. ampulla, it most closely resembles B. (B.) arua, which is smaller (19 vs. 45 mm, has fewer segments (84 vs. 128), male field pads L-shaped vs oval, intestinal origin in xiii vs xii, and folded prostates occupying xvii-xxiv vs. xvii-xviii. We were unable to obtain DNA barcode data for this species., Published as part of James, Samuel W., Bartz, Marie L. C. & Brown, George G., 2023, New Ocnerodrilidae genera, species and records from Brazil (Annelida: Crassiclitellata), pp. 235-269 in Zootaxa 5255 (1) on pages 254-256, DOI: 10.11646/zootaxa.5255.1.22, http://zenodo.org/record/7744817

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"]}

Urobenus brasiliensis brasiliensis Benham BRRS0049, 1 adult, surface soil horizon (0–20 cm) in lawn from the Estaç „o Experimental Cascata of Embrapa Clima Temperado, Pelotas-RS (S31.62047°, W52.52558°), 23 January 2012, G. Brown, S. James, S. Davidson, Y. Hong, G. Schiedeck, G. Schiavon, colls. BRRSDNA0118 to COFMBRRSDNA0122, 1 adult in each vial, surface soil horizon (0–20 cm) in gallery forest, Taquara-RS (S29.68597°, W50.85021°), 01 August 2014, R. Ott, G. Brown, colls. BRRSDNA0152 to BRRSDNA0155, 1 adult in each vial, surface soil horizon (0–20 cm) in native forest, Cruzeiro do Sul-RS, (S29.51696°, W51.98659°), 02 August 2014, S. James, M. Bartz, G. Cardoso, G. Brown, R. Ott, colls. BRRSDNA0169 to BRRSDNA0174, 1 adult in each vial, surface soil horizon (0–20 cm) in native pasture, Cruzeiro do Sul-RS, (S29.50020°, W51.98422°), 02 August 2014, S. James, M. Bartz, G. Cardoso, R. Ott, G. Brown, colls. BRRSDNA0184 to BRRSDNA0192, 1 adult in each vial, surface soil horizon (0–20 cm) in gallery forest, Estrela-RS, (S29.47560°, W51.95351°), 02 August 2014, R. Ott, G. Brown, colls. BRRSDNA0205, 1 adult, surface soil horizon (0–20 cm) in pasture, Estrela-RS, (S29.47242°, W51.94774°), 02 August 2014, S. James, M. Bartz, G. Cardoso, colls., Published as part of Ferreira, Talita, James, Samuel W., Bartz, Marie Luise Carolina, Lima, Ana Claudia Rodrigues De, Dudas, Rafaela & Brown, George Gardner, 2023, Distribution and diversity of earthworms in different land use systems in Rio Grande do Sul, Brazil, pp. 399-416 in Zootaxa 5255 (1) on page 402, DOI: 10.11646/zootaxa.5255.1.32, http://zenodo.org/record/7745077

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"]}