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Earthworm specimens collected in southern Taiwan consisted of seven new species of Amynthas and several previously known species, mostly widely distributed peregrines. The new species are A. nanrenensis of the octothecal A . corticis species group, A. monsoonus and A . huangi of the sexthecal A . aelianus species group, and four proandric octothecal species: A . chaishanensis , A . hengchunensis , A . kaopingensis and A . ailiaoensis . Amynthas chaishanensis has dorsal intrasegmental spermathecal pores, but the other three proandric species have dorsal, lateral or ventral intersegmental spermathecal pores, respectively. The proandric species are united by several features, including the enclosure of segment xi in a sac, as in A. formosae (also proandric), octothecal with spermathecae in vi-ix, spermathecal diverticula stalks generally kinked and often enclosed in membrane, and prostatic ducts divided polytomously into numerous small ductlets, which may be grouped into bundles of two to five. In A. ailiaoensis the prostatic duct trunk contains up to seven separate lumens in the ental half, surrounded by the circular muscle of the duct, while in A . chaishanensis the prostatic duct trunk contains about 40 small lumens. Of the previously known species in the collection, Pontodrilus litoralis and Metaphire houlleti are first reported from Taiwan. Additional locations for A . incongruus and A . robustus are given, and in the latter case the material appears to be the usual male-sterile morph. Other species found are Pontoscolex corethrurus , Amynthas corticis , A . gracilis , Metaphire californica , and Polypheretima elongata . [ABSTRACT FROM AUTHOR]

Abstract: Fourteen species new to science are described from material collected at several sites in the Blue Mountains and the John Crow Mountains of eastern Jamaica, doubling the known endemic Jamaican earthworm fauna. New data on Dichogaster montecyanensis (Sims) are provided. All species are placed in the genus Dichogaster Beddard, which is here treated sensu lato, i.e. including Eutrigaster Cognetti. Eight of the new species have lost the posterior pair of prostates and the seminal grooves of the male field. These are D. bromeliocola, D. crossleyi, D. davidi, D. garciai, D. harperi, D. haruvi, D. hendrixi, and D. johnsoni. D. sydneyi n. sp. has independently lost the posterior prostates but not the seminal grooves. The new species D. altissima and D. manleyi have the conventional dichogastrine prostatic battery and male field characteristics. Three species described here, D. farri, D. garrawayi, and D. marleyi, all have a third pair of prostates in the 20th segment, no seminal grooves, dorsal paired intestinal caeca in segment lxv, and lack penial setae. [Copyright &y& Elsevier]

Abstract: In this paper we present an up-to-date list of the terrestrial earthworms of the state of São Paulo (SP), Brazil. Seventy-seven species are known, of which about 39% are exotic and 61% native, mostly of the Glossoscolecidae (55% of total) and Ocnerodrilidae (17% of total) families. Pontoscolex corethrurus (Müller, 1857) and Amynthas spp. are widespread in the state, while many native species appear to be endemic. Various large-bodied worms (minhocuçus), some intensely harvested for the bait industry, may be endangered. Considering the significance of earthworm activity for soil function, the intensity of land use transformation in Brazil, and the probable existence of new species to be described from SP and other Brazilian states, further sampling efforts and taxonomic training of Brazilian scientists are urgently needed. [Copyright &y& Elsevier]

Amazonian rainforests, once thought to be pristine wilderness, are increasingly known to have been widely inhabited, modified, and managed prior to European arrival, by human populations with diverse cultural backgrounds. Amazonian Dark Earths (ADEs) are fertile soils found throughout the Amazon Basin, created by pre‐Columbian societies with sedentary habits. Much is known about the chemistry of these soils, yet their zoology has been neglected. Hence, we characterized soil fertility, macroinvertebrate communities, and their activity at nine archeological sites in three Amazonian regions in ADEs and adjacent reference soils under native forest (young and old) and agricultural systems. We found 673 morphospecies and, despite similar richness in ADEs (385 spp.) and reference soils (399 spp.), we identified a tenacious pre‐Columbian footprint, with 49% of morphospecies found exclusively in ADEs. Termite and total macroinvertebrate abundance were higher in reference soils, while soil fertility and macroinvertebrate activity were higher in the ADEs, and associated with larger earthworm quantities and biomass. We show that ADE habitats have a unique pool of species, but that modern land use of ADEs decreases their populations, diversity, and contributions to soil functioning. These findings support the idea that humans created and sustained high‐fertility ecosystems that persist today, altering biodiversity patterns in Amazonia. [ABSTRACT FROM AUTHOR]

We described 19 new species of pheretimoid earthworms in Palawan, Philippines. Of these, eight are Pheretima belonging to the Ph. darnleiensis group, characterised by having either four or five pairs of spermathecal pores located in 5/6–8/9. These were Pheretima ingentis sp. nov., Ph. palawanensis sp. nov., Ph. atongensis sp. nov., Ph. ascensa sp. nov., Ph. marinanaensis sp. nov., Ph. brookespointensis sp. nov., Ph. multicamera sp. nov., and Ph. hamadryades sp. nov. Seven are Polypheretima, of which Polypheretima victoriaensis sp. nov., Po. irawanensis sp. nov., Po. jenniferae sp. nov., Po. puertoprincesaensis sp. nov., Po. fleischmani sp. nov., and Po. mantalingahanensis sp. nov. belong to the Po. elongata group, characterised by having a pair of genital markings in xix, successive segments in line with the male pores, paired batteries of up to 28 spermathecae in vi and/or vii, and shallow copulatory bursae, while Po. zonadimidia sp. nov. is unique among all Polypheretima species in having a saddle-shaped clitellum and in having a pair of genital markings on xiv, xv, xvii and xix in line with the male pores. Two species are Amynthas, namely Am. magnaprostata sp. nov. of the Am. sieboldi group, characterised by having three pairs of spermathecal pores in 6/7–8/9 and a holandric male system, and Am. elnidoensis sp. nov. of the Am. minimus group, characterised by having a pair of spermathecal pores in 5/6. Also, two are Metaphire, namely Me. santalourdesensis sp. nov. of the Me. merabahensis group, characterised by having three pairs of spermathecal pores at 5/6–7/8, and Me. narraensis sp. nov. of the Me. peguana group, characterised by having three pairs of spermathecal pores in 6/7–8/9 and post-clitellar genital markings at 17/18 and 18/19. With thorough biological surveys in Palawan, we expect that more species will be discovered in this biogeographic faunal region. [ABSTRACT FROM AUTHOR]

Earthworms (Annelida: Crassiclitellata) are widely distributed around the world due to their ancient origination as well as adaptation and invasion after introduction into new habitats over the past few centuries. Herein, we report a 1.2 Gb complete genome assembly of the earthworm Amynthas corticis based on a strategy combining third-generation long-read sequencing and Hi-C mapping. A total of 29,256 protein-coding genes are annotated in this genome. Analysis of resequencing data indicates that this earthworm is a triploid species. Furthermore, gene family evolution analysis shows that comprehensive expansion of gene families in the Amynthas corticis genome has produced more defensive functions compared with other species in Annelida. Quantitative proteomic iTRAQ analysis shows that expression of 147 proteins changed in the body of Amynthas corticis and 16 S rDNA sequencing shows that abundance of 28 microorganisms changed in the gut of Amynthas corticis when the earthworm was incubated with pathogenic Escherichia coli O157:H7. Our genome assembly provides abundant and valuable resources for the earthworm research community, serving as a first step toward uncovering the mysteries of this species, and may provide molecular level indicators of its powerful defensive functions, adaptation to complex environments and invasion ability. Xing Wang et al. report a 1.2 Gb complete genome assembly of the earthworm Amynthas corticis and demonstrate that it is a triploid species. Gene family evolution, proteomic iTRAQ analysis, and 16 S rDNA sequencing demonstrate the evolution of adaptive defensive functions relative to other species in Annelida. [ABSTRACT FROM AUTHOR]

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

Net primary productivity (NPP) influences soil food webs and ultimately the amount of carbon (C) inputs in ecosystems. Earthworms can physically protect organic matter from rapid mineralization through the formation of soil aggregates. Previous studies at the Oak Ridge National Laboratory (ORNL) Free Air CO 2 Enrichment (FACE) experiment showed that elevated [CO 2 ] (e[CO 2 ]) increased fine-root production and increased soil C through soil aggregation compared to ambient [CO 2 ] (a[CO 2 ]) conditions. Our first objective was to study the response of earthworms to increased leaf and root-litter inputs caused by increased atmospheric [CO 2 ] exposure. We also took advantage of the CO 2 shutdown at the ORNL FACE site to track the shift of the δ 13 C signal in leaf-litter, fine roots, earthworms, earthworm casts, and bulk soil. Densities of the most abundant endogeic earthworm, Diplocardia spp., were positively correlated with the previous-year production of leaf litter (r = 0.66, P = 0.02) and fine roots (r = 0.62, P = 0.03); and with the leaf-litter production (r = 0.63, P = 0.03) and fine-root production (r = 0.59, P = 0.05) two years before earthworms were sampled. Within two years after the CO 2 fumigation ceased, the 13 C/ 12 C ratio increased in leaf litter (P = 0.01) and in fine roots (P = 0.05), showing an ecosystem legacy effect on soil C inputs. However, the C isotopic composition of soil, endogeic earthworms and casts had not changed the two years after the CO 2 fumigation ended. The positive response of earthworms to increased root NPP, caused by elevated [CO 2 ], is consistent with the increased soil aggregate formation and increased soil C at the ORNL FACE in the e[CO 2 ] treatment. [ABSTRACT FROM AUTHOR]

Peregrine pheretimoid earthworms, commonly known as jumping worms, are members of the family Megascolecidae that have become widely established outside of their native ranges. In many parts of the world this represents a second wave of earthworm invasions, following the introduction of peregrine European earthworms in the family Lumbricidae during the colonial era. Forest ecologists, turf managers, gardeners, and other land managers are concerned about the observed or presumed negative effects of jumping worms on invaded habitats. Although research on jumping worms has accelerated in recent decades, our understanding of their ecology remains limited. We compiled techniques useful to researchers working to fill voids in our understanding. Similar past efforts have focused on tools used to study common European species. Differences in life cycle, behavior, morphology, and physiology make it difficult to transfer experiences with European earthworms to pheretimoids. For example, the loss of reproductive features in many pheretimoid populations poses a challenge for identification, and techniques for individually tagging lumbricid earthworms have been less successful for megascolecids. The active and ongoing expansion of pheretimoid populations in many areas requires increased attention on distributed methods, such as citizen-science protocols, for detecting and tracking their expansion. Finally, the desire to limit populations of pheretimoids, including those invading gardens and other environments that might be successfully restored, has exposed the lack of options for targeted, effective control of unwanted earthworms. We identify opportunities to address these voids in our methodological tool kit and encourage the adaptation of techniques previously used in the study and management of other invasive animals. [ABSTRACT FROM AUTHOR]

Three new earthworms of the Amynthas corticis-group (Clitellata: Megascolecidae) from Phouleoi Mountain, Lao PDR. Earthworm specimens were collected from Mt. Phouleoi, Viengthong District, Houaphanh Province, Lao PDR. The species belong to the pheretimoid earthworms. All three holandric species key out to the A. corticis-group in Sims & Easton (1972), having spermathecal pores in 5/68/9. The new species are: Amynthas ammalinei sp. nov., Amynthas xayi sp. nov., and Amynthas piyakarnae sp. nov. Amynthas ammalinei sp. nov. has male pores superficial on small round porophores, 0.160.31 circumference apart. Amynthas xayi sp. nov. has male pores superficial in XVIII at centers of round porophores, 0.170.29 circumference apart. Amynthas piyakarnae sp. nov. has male pores superficial on transverse elevated elongate oval pads equatorial on XVIII, pads surrounded by 34 small outer rings, 0.250.31 circumference apart.

We describe three new species of Eukerria, six of Kerriona, one of Ocnerodrilus and two of Belladrilus (Clitellata: Ocnerodrilidae) from Brazil, mostly from wet habitats including bromeliad leaf axils, plus some from forest soils in Bahia state. We also provide new morphological and distribution data on Kerriona luederwaldti Michaelsen, 1924. We describe the new genus Pauqueba and its two new species, both from bromeliad leaf axils and forest floor litter in the state of Bahia. It has two pairs of calciferous glands, the balantine reduction of the male terminalia but spermathecae in ix, and no gizzard. The new genus Parabauba found in deeper (30 cm) soil layers is distinctive for the combination of two pairs of calciferous glands, a large gizzard in v, acanthodriline male terminalia, and no typholosole. The new genus Diplomoela from a mesic soil in Bahia is meronephric, unusually large for the Ocnerodrilidae, and has two well-developed gizzards.

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

The southernmost state of Brazil, Rio Grande do Sul, includes both the Pampa and Atlantic Forest biomes, and covers an area of around 284 thousand km2. Since the last checklist for the state, published in 2012, many additional sites have been sampled. Hence, we provide here a full list of earthworm species and their distribution in different municipalities and land use systems in the state. Data were obtained from the literature, museums, and other earthworm collections and recent sampling efforts. Records for 19 additional species are reported here for the first time, raising the total to 78 species/subspecies currently known from Rio Grande do Sul, from the families Acanthodrilidae (5), Benhamiidae (3), Lumbricidae (13), Megascolecidae (11), Ocnerodrilidae (22), Almidae (2), Glossoscolecidae (18) and Rhinodrilidae (5). Alexidrilus Righi, 1971 is synonymized with Urobenus Benham, 1886, as both species in the genus (A. littoralis Ljungström, 1972 and A. lourdesae Righi, 1971) have similar morphology to the type species Urobenus brasiliensis Benham, 1886. Most of the native species/subspecies in the state (39), including an important number of new species, were found in native vegetation with lesser disturbance, such as swamps, native forests and pastures. On the other hand, the three peregrine and 27 exotic species (especially Pontoscolex corethrurus, Eukerria spp., lumbricids, megascolecids, Dichogaster spp., and Microscolex spp.) dominated in disturbed sites, such as compost piles, urban areas, rice and other crops, and cultivated pastures. Considering the geographic extension and the many unsampled sites in the state, further collection efforts are necessary and will certainly reveal new species, as evidenced here.

The earthworms (Crassiclitellata) of the Northern Atlantic Forest region are poorly known, and many new species are expected to be found in the region. Hence, quantitative and qualitative samples were taken in the Atlantic Forest region of southern Bahia State, Brazil, at eight sites to assess species presence and abundance in different land uses (forests, pastures, plantations). Earthworms were also qualitatively collected at an additional eight sites. Only one to two species per site were found in quantitative samples, while qualitative samples generally resulted in higher species recoveries, with highest richness observed at the native Atlantic Forest of Veracel in Porto Seguro. Pontoscolex corethrurus (Müller, 1857) was the dominant species at all sites and in most land use systems, probably favored by human migration and agricultural activities; it was only absent in one primary forest site where native Ocnerodrilidae species predominated. In total at least 14 species were found of which eight were new, belonging to the genus Rhinodrilus and three new genera in the Ocnerodrilidae family (Pauqueba, Parabauba and Diplomoela). Contrary to sites in Southern and Southeastern Brazil, where many native species belong to the Glossoscolecidae family, in southern Bahia, Ocnerodrilidae species tend to be more prevalent, occupying various habitats. More intensive sampling of these habitats, including surface litter, in and under fallen logs and dead tree trunks, under rocks, in bromeliad leaf tanks and deeper soil layers is essential in order to properly characterize earthworm communities in Brazilian Atlantic Forests.

Integrative approach of taxonomy drives us to the discovery of three new species of the genus Eutyphoeus Michaelsen 1900, namely Eutyphoeus phawngpuiensis sp. nov., Eutyphoeus serei sp. nov. and Eutyphoeus tawi sp. nov. from protected forests of Mizoram, India. E. phawngpuiensis sp. nov. belongs to the Eutyphoeus hastatus group, and is characterized by holandric male organs, having spermathecal pores at a or ab, bidiverticulate spermathecae, and lack of genital marking glands. While, E. tawi sp. nov. has spermathecal pores at b or lateral to b, without lateral intestinal caeca. E. serei sp. nov. belongs to the Eutyphoeus gigas group, and is distinguished by having metandric male organs, lacking female pore of right side, and having spoon shaped tip of penial setae without ornamentation. Molecular characterization using coi gene marker also supports the findings. The maximum intraspecific distances and distance to the nearest neighbour of E. phawngpuiensis sp. nov., E. serei sp. nov. and E. tawi sp. nov. were recorded 3.5%, 9.37%; 2.35%, 14.83%; and 2.28%, 12.86% respectively. The key to Eutyphoeus species of India was also updated.

Two new species of the genus Pheretima are described from Mt. Amuyao and Mt. Lamagan, Mountain Province, Philippines, namely Pheretima riparianensis sp. nov. and Pheretima bontocensis sp. nov. Both new species belong to the P. sangirensis species group characterized by having only one pair of spermathecal pores in 7/8. Pheretima riparianensis sp. nov. has distance between spermathecal pores 12 mm; copulatory bursae openings 0.11 circumference ventrally apart. Pheretima bontocensis sp. nov. has distance between spermathecal pores 1.4 mm; copulatory bursae openings 0.10 circumference ventrally apart.

Nonillon M. Aspe and Samuel W. James (2016) Studies over the past decade have shown the Philippine native earthworm fauna to be highly diverse, with apparent high endemism among islands and localities. Here we describe 17 new species of pheretimoid earthworms collected from priority conservation sites on Mindanao and associated islands. Nine species of the subgenus Pheretima ( Pheretima ), distinguished by the absence of secretory diverticula on the coelomic surface of the copulatory bursae, were detected. Pheretima acia n. sp., P. dinagatensis n. sp., P. enormis n. sp., P. hamiguitanensis n. sp., P. lantapanensis n. sp. belong to the P. urceolata species group, with a pair of spermathecal pores in 5/6; P. timpoongensis n. sp., P. camiguinensis n. sp., P. sibucalensis n. sp., and P. apoensis n. sp. belong to the P. sangirensis group, with a pair of spermathecal pores in 7/8. Pheretima ( Parapheretima ) pandanensis n. sp. and P. ( Parapheretima ) boaensis n. sp., members of the subgenus Pheretima ( Parapheretima ), with secretory diverticula on the coelomic surface of the copulatory bursae, are the first Philippine records of Parapheretima . In the genus Amynthas , A. dinagatensis n. sp. is athecate; A. cagdianaoensis n. sp. belongs to the belongs to the A. supuensis species group, with a pair of spermathecal pores at 8/9; and A. talaandigensis n. sp. belongs to A. rimosus group, with intrasegmental spermathecae in 6, 7, 8, and 9. In the genus Polypheretima , P. bukidnonensis n. sp. belongs to the Po. elongata species group, with paired genital markings on xix and successive segments, and paired spermathecal batteries in vi and/or vii; Po. zamboangensis n. sp., either has spermathecal pores in 5/6/7 or lacks them, and has a metandric male sexual system. Pithemera nolani n. sp. is unique among congeners in having only one pair of spermathecal pores in 5/6 and in having copulatory bursae. The geological history of Mindanao, which is a collision complex between three arc systems, may have contributed to the high diversity of the earthworm fauna on Mindanao. Future molecular studies can help elucidate the evolutionary relationships and geographical distribution of species and populations, and shed light on mechanisms of earthworm diversification in the Philippines.