Mercury critical concentrations to Enchytraeus crypticus (Annelida: Oligochaeta) under normal and extreme conditions of moisture in tropical soils - Reproduction and survival.

Soil provides many ecosystem services that are essential to maintain its quality and healthy development of the flora, fauna and human well-being. Environmental mercury levels may harm the survival and diversity of the soil fauna. In this respect, efforts have been made to establish limit values of mercury (Hg) in soils to terrestrial fauna. Soil organisms such as earthworms and enchytraeids have intimate contact with trace metals in soil by their oral and dermal routes, reflecting the potentially adverse effects of this contaminant. The main goal of this study was to obtain Hg critical concentrations under normal and extreme conditions of moisture in tropical soils to Enchytraeus crypticus to order to assess if climate change may potentiate their acute and chronic toxicity effects. Tropical soils were sampled from of two Forest Conservation Units of the Rio de Janeiro State - Brazil, which has been contaminated by Hg atmospheric depositions. Worms were exposed to three moisture conditions, at 20%, 50% and 80% of water holding capacity, respectively, and in combination with different Hg (HgCl2) concentrations spiked in three types of tropical soil (two natural soils and one artificial soil). The tested concentrations ranged from 0 to 512mg Hg kg-1 dry weight. Results indicate that the Hg toxicity is higher under increased conditions of moisture, significantly affecting survival and reproduction rate.

A TME study with the fungicide pyrimethanil combined with different moisture regimes: effects on enchytraeids.

Today's ecosystems are influenced by different factors that could evolve into stressors. Effects of pesticides, especially in agricultural areas, may interact with environmental factors, such as soil moisture fluctuation caused by global climate change. In this contribution, two semi-field studies conducted in Germany and Portugal with terrestrial model ecosystems are presented. Their aim was to assess the effects of the fungicide pyrimethanil under different soil moisture levels on Enchytraeidae. In Portugal a no observed effect concentration design was chosen, using two concentration levels: the maximum application rate (MAR) according to the safe use registration within the European Union and five times the MAR (1.82 and 9.09 mg/kg dry soil, respectively). Both concentrations did neither affect the total enchytraeid abundance nor single populations. In Germany an ECx design (effect concentration) was conducted, using 11 concentrations. In general, 14 EC50 values for different combinations of single species, moisture level and sampling date were determined. The strongest effects were found in dry soil, particularly for Fridericia connata (EC50: 3.48 mg/kg dry soil after 8 weeks of exposure). The advantages and challenges of these test designs are discussed with regard to the registration process of pesticides in the European Union. In any case, enchytraeids are suitable test organisms in such higher tier studies for the combined evaluation of chemical and climatic stressors due to their usually high diversity and abundances and their close contact with the soil solution.

Analysis of heat and cold tolerance of a freeze-tolerant soil invertebrate distributed from temperate to Arctic regions: evidence of selection for extreme cold tolerance.

Tolerance to thermal extremes is critical for the geographic distributions of ectotherm species, many of which are probably going to be modified by future climatic changes. To predict species distributions it is important to understand the potential of species to adapt to changing thermal conditions. Here, we tested whether the thermal tolerance traits of a common freeze-tolerant potworm were correlated with climatic conditions and if adaptation to extreme cold constrains the evolutionary potential for high temperature tolerance. Further, we tested if evolution of thermal tolerance traits is associated with costs in other fitness traits (body size and reproduction). Lastly, we tested if slopes of temperature-survival curves (i.e., the sensitivity distribution) are related to tolerance itself. Using 24 populations of the potworm, Enchytraeus albidus Henle (Enchytraeidae), collected from a wide range of climatic conditions, we established a common garden experiment in which we determined high and low temperature tolerance (using survival as endpoint), average reproductive output and adult body size. Heat tolerance was not related to environmental temperatures whereas lower lethal temperature was about 10 °C lower in Arctic populations than in populations from temperate regions. Reproduction was not related to environmental temperature, but was negatively correlated with cold tolerance. One explanation for the trade-off between cold tolerance and reproduction could be that the more cold-hardy populations need to channel energy to large glycogen reserves at the expense of less energy expenditure for reproduction. Adult body size was negatively related to environmental temperature. Finally, the slopes of temperature-survival curves were significantly correlated with critical temperature limits for heat and cold tolerance; i.e., slopes increased with thermal tolerance. Our results suggest that relatively heat-sensitive populations possess genetic variation, leaving room for improved heat tolerance through evolutionary processes, which may alleviate the effects of a warmer future climate in the Arctic. On the other hand, we observed relatively narrow sensitivity distributions (i.e., less variation) in the most heat tolerant populations. Taken together, our results suggest that both cold and heat tolerance can only be selected for (and improved) until a certain limit has been reached.

Description of Hemienchytraeus wuhanensis sp. nov. (Annelida, Clitellata, Enchytraeidae) from central China, with comments on species records of Hemienchytraeus from China.

Hemienchytraeus wuhanensis sp. nov. is described from hardwood forest soil in Wuhan, China. This moderately sized enchytraeid species of 6-9 mm body length is characterized by: (1) an oesophageal appendage with tertiary branches, (2) three pairs of secondary pharyngeal gland lobes in V, VI, VII, (3) five pairs preclitellar nephridia, from 5/6 to 9/10, (4) dorsal vessel originating in clitellar segments, (5) a girdle-shaped clitellum, (6) a relatively small male reproductive apparatus without seminal vesicle, and (7) spermathecae that extend to VI-VII. DNA barcodes of paratype specimens of the new species are provided. Previous species records of Hemienchytraeus from China are critically discussed.

Segmented worms (Phylum Annelida): a celebration of twenty years of progress through Zootaxa and call for action on the taxonomic work that remains.

Zootaxa has been the leading journal on invertebrate systematics especially within Annelida. Our current estimates indicate annelids include approximately 20,200 valid species of polychaetes, oligochaetes, leeches, sipunculans and echiurans. We include herein the impact of Zootaxa on the description of new annelid species in the last two decades. Since 2001, there have been over 1,300 new annelid taxa published in about 630 papers. The majority of these are polychaetes (921 new species and 40 new genera) followed by oligochaetes (308 new species and 10 new genera) and leeches (21 new species). The numerous papers dealing with new polychaete species have provided us a clear picture on which polychaete families have had the most taxonomic effort and which authors and countries have been the most prolific of descriptions of new taxa. An estimated additional 10,000+ species remain to be described in the phylum, thus we urge annelid workers to continue their efforts and aid in training a new generation of taxonomists focused on this ecologically important group.

Legacy effects of temporary grassland in annual crop rotation on soil ecosystem services.

The introduction of temporary grassland into an annual crop rotation is recognized to improve soil ecosystem services, and resulting legacies can be beneficial for the following crops. In this context, the aim of the present study was to evaluate legacy effects of introducing temporary grassland into an annual crop rotation on five ecosystem services (i) soil structure maintenance (aggregate stability), (ii) water regulation (saturated hydraulic conductivity), (iii) biodiversity conservation (microbial biomass and microbial metabolic activity, as well as microorganism, enchytraeid, springtail and earthworm communities), (iv) pathogen regulation (soil suppressiveness to Verticillium dahliae), and (v) forage production and quality. Three crop rotation schemes, maintained for twelve years, were compared in four random blocks, one being an annual crop rotation without grassland (0%), another with a medium percentage of grassland (50%, corresponding to 3 years of continuous grassland in the crop rotation), and a third one with a high percentage of grassland in the crop rotation (75%, corresponding to 6 years of continuous grassland in the crop rotation). The results showed that the grassland introduction into an annual crop rotation improved, whatever the duration of the grassland, soil structure maintenance and biodiversity conservation, while it decreased pathogen regulation and did not modify water regulation. Comparing the two crop rotations that included grassland, indicated a stronger beneficial grassland legacy effect for the higher proportion of grassland concerning soil structure maintenance and biodiversity conservation. By contrast, water regulation, pathogen regulation and forage production were not affected by the legacy of the 75% grassland during the rotation. Overall, our findings demonstrated the extent to which grassland legacies are affecting the current state of soil properties and possible ecosystem services provided. To improve ecosystem services, soil management should take legacy effects into account and consider longer timeframes to apply beneficial practices.

Global diversity of earthworms and enchytraeids (Clitellata): papers in honor of András Zicsi (1928-2015). Editorial.

This Special Volume of Zootaxa unites forty papers written in honor of the late András Zicsi (1928-2015), the eminent earthworm taxonomist. They deal with the taxonomy, systematics and distribution of earthworms and enchytraeids, the two major groups of soil-dwelling Oligochaeta. Altogether, 71 new species-group taxa are described, 60 species and subspecies of earthworms and 11 species of enchytraeids. They are from 15 countries all around the globe: Spain, Italy, Hungary, Turkey, Botswana, Mexico, Colombia, Brazil, China, Japan, Korea, Taiwan, Laos, Malaysia, Thailand, and Vietnam.

A new species of Enchytraeus (Enchytraeidae, Oligochaeta) from the profundal of Lake Van, the world's largest soda Lake (Turkey, East Anatolia).

Enchytraeus polatdemiri sp. nov. (Enchytaeidae, Oligochaeta) was discovered in the framework of a sampling campaign of the benthic invertebrate fauna of the hyperalkaline Lake Van in Eastern Anatolia, Turkey, the third-largest closed lake and the largest soda lake on Earth. It was the only oligochaete species found in all samples. DNA sequencing included a fragment of the mitochondrial cytochrome c oxidase subunit I (COI) gene, and a fragment of the nuclear histone 3 (H3) gene. For comparison, specimens from laboratory cultures of E. albidus Henle, 1837, a widespread and morphologically similar species, were sequenced as well. The new species differs from E. albidus in comparatively small body size, 2 or 3 chaetae per bundle, saddle-shaped clitellum, absence of a copulatory field between the male pores and vasa deferentia usually not extending beyond the clitellum. The individual gene trees of COI and H3, as well as the combined phylogenetic analysis of both trees, recovered Enchytraeus polatdemiri sp. nov. as a monophyletic group within the genus Enchytraeus, closely related to E. albidus, but with an average p-distance for COI of 14.5 %. E. polatdemiri sp. nov. may have evolved from a local population of Enchytraeus albidus, a species well-adapted to changing salinity conditions, or from a common ancestor into an extremophile species that dwells and reproduces in the profundal of a strongly alkaline soda lake.

Soil networks become more connected and take up more carbon as nature restoration progresses.

Soil organisms have an important role in aboveground community dynamics and ecosystem functioning in terrestrial ecosystems. However, most studies have considered soil biota as a black box or focussed on specific groups, whereas little is known about entire soil networks. Here we show that during the course of nature restoration on abandoned arable land a compositional shift in soil biota, preceded by tightening of the belowground networks, corresponds with enhanced efficiency of carbon uptake. In mid- and long-term abandoned field soil, carbon uptake by fungi increases without an increase in fungal biomass or shift in bacterial-to-fungal ratio. The implication of our findings is that during nature restoration the efficiency of nutrient cycling and carbon uptake can increase by a shift in fungal composition and/or fungal activity. Therefore, we propose that relationships between soil food web structure and carbon cycling in soils need to be reconsidered.

Biosolids applied to agricultural land: Influence on structural and functional endpoints of soil fauna on a short- and long-term scale.

Biosolids have well-documented crop and soil benefits similar to other sources of organic amendment, but there is environmental concern due to biosolids-associated pollutants. The present study investigated two field sites that had received biosolids at commercial-scale rates in parallel to associated field sections which were managed similarly but without receiving biosolids (controls). The investigated endpoints were abundance and diversity of soil organisms (nematodes, enchytraeids and earthworms) and soil fauna feeding activity as measured by the bait lamina assay. Repeated sampling of one of the field sites following the only biosolids application demonstrated an enrichment effect typical for organic amendments, which was mostly exhausted after 44months. After an initial suppression, the proportion of free-living plant-parasitic nematodes tended to increase in the biosolids-amended soil over time. Yet, none of the endpoints at this site indicated significant negative effects resulting from the biosolids until 44months post application. In contrast to the repeatedly tilled first field site, the second one was left fallow after three biosolids applications, and was sampled 96months post last application. It was only at this field site that potential evidence for a long-term impact of biosolids was detected with regard to two endpoints: earthworm abundance and structure of the nematode assemblage. Agricultural management and correlation with abiotic soil parameters explained the observed difference in earthworm abundance. Yet, the development of a highly structured and mature nematode assemblage at the control but not at the biosolids-amended section of this fallow field could not be explained by such correlations nor by soil metal concentrations. Overall, the present study found only weak evidence for negative long-term impacts of biosolids applied at commercial rates on soil fauna. High-level community parameters such as the nematode structure index (SI) appeared more suitable to detect deleterious effects on soil fauna than simple abundance measurements.

Microdrile Oligochaeta in bromeliad pools of a Honduran cloud forest.

Phytotelmata, or plant-held water bodies, often house complex aquatic invertebrate communities. Microdrile oligochaetes (Clitellata, Annelida) are known to be part of that community, but specimens are rarely identified to species level. Here we report three species of Enchytraeidae and three species of Naididae from a collection sampled in phytotelms of bromeliads in Cusuco National Park, Honduras. Two species of enchytraeids are new to science. Bryodrilus hondurensis sp. nov. is distinguished from other members of the genus by the high number of ventral chaetae, up to 11 per bundle. The genus is Holarctic and this is the southernmost record so far of a Bryodrilus species. Hemienchytraeus phytotelmatus sp. nov. is distinguished by a combination of characters, among which the huge spermathecae and seminal vesicles are most conspicuous. The genus is common in tropical and subtropical soils around the world. A third species of enchytraeids in the collection, Cernosvitoviella atrata (Bretscher), is redescribed, together with three known species of Naididae, Pristina jenkinae (Stephenson), Pristina osborni (Walton) and Pristina terrena Collado & Schmelz. Presence of ingested debris and humus in the intestine of most specimens suggests that the collected animals live and reproduce in the phytotelms. We provide a list of oligochaete species recorded so far from bromeliad pools in Central and South America.