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

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

Genetic, epigenetic and microbiome characterisation of an earthworm species (Octolasion lacteum) along a radiation exposure gradient at Chernobyl.

The effects of exposure to different levels of ionising radiation were assessed on the genetic, epigenetic and microbiome characteristics of the "hologenome" of earthworms collected at sites within the Chernobyl exclusion zone (CEZ). The earthworms Aporrectodea caliginosa (Savigny, 1826) and Octolasion lacteum (Örley, 1881) were the two species that were most frequently found at visited sites, however, only O. lacteum was present at sufficient number across different exposure levels to enable comparative hologenome analysis. The identification of morphotype O. lacteum as a probable single clade was established using a combination of mitochondrial (cytochrome oxidase I) and nuclear genome (Amplified Fragment Length Polymorphism (AFLP) using MspI loci). No clear site associated differences in population genetic structure was found between populations using the AFLP marker loci. Further, no relationship between ionising radiation exposure levels and the percentage of methylated loci or pattern of distribution of DNA methylation marks was found. Microbiome structure was clearly site dependent, with gut microbiome community structure and diversity being systematically associated with calculated site-specific earthworm dose rates. There was, however, also co-correlation between earthworm dose rates and other soil properties, notably soil pH; a property known to affect soil bacterial community structure. Such co-correlation means that it is not possible to attribute microbiome changes unequivocally to radionuclide exposure. A better understanding of the relationship between radionuclide exposure soil properties and their interactions on bacterial microbiome community response is, therefore, needed to establish whether these the observed microbiome changes are attributed directly to radiation exposure, other soil properties or to an interaction between multiple variables at sites within the CEZ.

Coordinated community structure among trees, fungi and invertebrate groups in Amazonian rainforests.

Little is known regarding how trophic interactions shape community assembly in tropical forests. Here we assess multi-taxonomic community assembly rules using a rare standardized coordinated inventory comprising exhaustive surveys of five highly-diverse taxonomic groups exerting key ecological functions: trees, fungi, earthworms, ants and spiders. We sampled 36 1.9-ha plots from four remote locations in French Guiana including precise soil measurements, and we tested whether species turnover was coordinated among groups across geographic and edaphic gradients. All species group pairs exhibited significant compositional associations that were independent from soil conditions. For some of the pairs, associations were also partly explained by soil properties, especially soil phosphorus availability. Our study provides evidence for coordinated turnover among taxonomic groups beyond simple relationships with environmental factors, thereby refining our understanding regarding the nature of interactions occurring among these ecologically important groups.

Effect of gamma radiation on the production of bystander signals from three earthworm species irradiated in vivo.

The effect of gamma radiation delivered over 24 h on the induction of bystander signals of three earthworm species exposed in vivo was investigated: A. chlorotica, A. caliginosa, and E. tetraedra. Worms were exposed to external gamma irradiation (Co-60 source) for 24 h and samples of head, body, and clitellum were dissected from exposed and control worms and placed in culture medium for 24 h at 19 C. The harvested medium was filtered and assayed for expression of bystander signals using both clonogenic and mitochondrial reporter assays. Different responses were observed in the different species and in the different tissues. A. chlorotica worm-treated reporters show insignificant mitochondrial response for all sections, yet a significant clonogenic reduction in survival for body sections. A. caliginosa worm-treated reporters show a significant mitochondrial response for some sections and insignificant mitochondrial response and insignificant reduction in clonogenic survival for the rest. E. tetraedra worms from a control site show significant evidence of bystander signalling, measured by mitochondrial response in reporter cells, for all sections while those harvested from a contaminated site show insignificant changes in baseline signalling when exposed to the challenge dose. In vivo exposure of earthworm species shows evidence of bystander signalling using two different reporter assays. This effect varied between the different species and tissues. There is also evidence of attenuated bystander signalling in worms harvested from a site contaminated with radiation.

Bioavailability of cobalt and silver nanoparticles to the earthworm Eisenia fetida.

Due to difficulties in tracing engineered nanoparticles (ENPs) in complex media, there are few data on the exposure of soil biota to ENPs. This study used neutron activated cobalt (Co NPs) and silver (Ag NPs) nanoparticles, as well as soluble cobalt and silver salts, to assess the uptake, excretion and biodistribution in the earthworm Eisenia fetida. Concentrations of cobalt in worms after four weeks exposure reached 88% and 69% of the Co ions and Co NPs concentrations in food, respectively, while corresponding values for Ag ions and Ag NPs were 2.3% and 0.4%. Both Ag ions and Ag NPs in earthworms were excreted rapidly, while only 32% of the cobalt accumulated from Co ions and Co NPs were excreted within four months. High accumulation of cobalt was found in blood and in the digestive tract. Metal characterization in the exposure medium was assessed by sequential extraction and ultrafiltration. The Co NPs showed significant dissolution and release of ions, while Ag ions and particularly Ag NPs were more inert.

Ecotoxicological assessment of TiO2 byproducts on the earthworm Eisenia fetida.

The increasing production of nanomaterials will in turn increase the release of nanosized byproducts to the environment. The aim of this study was to evaluate the behaviour, uptake and ecotoxicity of TiO(2) byproducts in the earthworm Eisenia fetida. Worms were exposed to suspensions containing 0.1, 1 and 10 mg/L of byproducts for 24 h. Size of TiO(2) byproducts showed aggregation of particles up to 700 µm with laser diffraction. Only worms exposed at 10 mg/L showed bioaccumulation of titanium (ICP-AES), increasing expression of metallothionein and superoxide dismutase mRNA (Real-time PCR) and induction of apoptotic activity (Apostain and TUNEL). TiO(2) byproducts did not induce cytotoxicity on cœlomocytes, but a significant decrease of phagocytosis was observed starting from 0.1 mg/L. In conclusion, bioaccumulation of byproducts and their production of reactive oxygen species could be responsible for the alteration of the antioxidant system in worms.

Ecotoxicological effects of an aged TiO2 nanocomposite measured as apoptosis in the anecic earthworm Lumbricus terrestris after exposure through water, food and soil.

Titanium dioxide nanoparticles seem to have a low toxicity to terrestrial organisms, though few studies are published in this area. TiO(2) used in sunscreens are nanocomposites where TiO(2) has been coated with magnesium, silica or alumina, as well as amphiphilic organics like polydimethyl siloxane (PDMS), and these coatings are modified by ageing. We assessed the ecotoxicity and propensity for bioaccumulation of an aged TiO(2) nanocomposite used in sunscreen cosmetics, and its potential effect on the frequency of apoptosis in different earthworm tissues. The earthworm Lumbricus terrestris was exposed to the TiO(2) nanocomposite for 7 days in water or 2-8 weeks in soil with the nanocomposite mixed either into food or soil at concentrations ranging from 0 to 100 mg kg(-1). Apoptosis was then measured by immunohistochemistry and Ti localized by XRF microscopy. Results showed no mortality, but an enhanced apoptotic frequency which was higher in the cuticule, intestinal epithelium and chloragogenous tissue than in the longitudinal and circular musculature. TiO(2) nanoparticles did not seem to cross the intestinal epithelium/chloragogenous matrix barrier to enter the coelomic liquid, or the cuticule barrier to reach the muscular layers. No bioaccumulation of TiO(2) nanocomposites could thus be observed.

Silver nanoparticle exposure causes apoptotic response in the earthworm Lumbricus terrestris (Oligochaeta).

In terrestrial ecotoxicology there is a serious lack of data for potential hazards posed by engineered nanoparticles (ENPs). This is partly due to complex interactions between ENPs and the soil matrix, but also to the lack of suitable toxicological end points in organisms that are exposed to ENPs in a relevant manner. Earthworms are key organisms in terrestrial ecosystems, but so far only physiological end points of low sensitivity have been used in ecotoxicity studies with ENPs. We exposed the earthworm Lumbricus terrestris to silver nanoparticles and measured their impact on apoptosis in different tissues. Increased apoptotic activity was detected in a range of tissues both at acute and sublethal concentrations (down to 4 mg/kg soil). Comparing exposure in water and soil showed reduced bioavailability in soil reflected in the apoptotic response. Apoptosis appears to be a sensitive end point and potentially a powerful tool for quantifying environmental hazards of ENPs.