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Taxonomical problems in enchytraeids (Oligochaeta) from Spitsbergen
... In Arctic soils, enchytraeids are a particularly important decomposer group. Whereas there are relatively many faunistic records from the Svalbard archipelago and its main island Spitsbergen, there are still few ecological studies on enchytraeids from this area and the High Arctic in general (Stephenson, 1922(Stephenson, , 1924(Stephenson, , 1925Nurminen, 1965Nurminen, , 1966Birkemoe and Dózsa-Farkas, 1994;Birkemoe, 1995;Dózsa-Farkas, 1999;Sømme and Birkemoe, 1997;Bauer, 2002aBauer, , 2002bBirkemoe et al., 2000;Coulson et al., 2013aCoulson et al., , 2013bÁ vila-Jiménez et al., 2019;Devetter et al., 2021). Several studies have looked into the potential effect of nutrient enrichment of tundra soil close to bird cliffs, but none at its effect on enchytraeids (but see Schlaghamersky and Devetter, 2019). ...
... Didden, 1991;Nowak, 2004) but also in some wetlands (Schenková et al., 2018). In the checklists of Dózsa-Farkas (1999) and Coulson and Refseth (2004) E. buchholzi was not listed for Svalbard. Nor was it listed in the update published by Coulson (2007) or in the online version of the Checklist of the terrestrial and freshwater invertebrate fauna of Svalbard dated 6 June 2013 (Coulson, 2013). ...
... Individual terrestrial sampling sites have been shown to host assemblages of up to 11 species on the island of Edgeøya (Ávila-Jiménez et al., 2019), but assemblages of 8 species and above all included the primarily aquatic, marine-littoral Lumbricullus nielseni Nurminen, 1965 and were from samples taken close to the shoreline. Dózsa-Farkas (1999) found up to 9 species at sites along the coast of the island of Nordaustlandet, again the highest species richness was found in wet sites and included one or two primarily aquatic species of Lumbricillus. From Spitsbergen island not more than 6 species have been reported from a single site (habitat) so far (for a detailed account see Schlaghamersky and Devetter, 2019), except for Barentsburg with 7 species in imported soil, including the presumably introduced E. dichaetus, E. buchholzi s.l. and Cognettia glandulosa (Michaelsen, 1888). ...
... Enchytraeid life cycles vary with species and habitat. Reproduction may occur throughout the year or during a short period of time (Springett 1970;Römbke 1989;Dózsa-Farkas 1999), with corresponding variation in lifecycle length (Springett 1970;Klungland 1981;Römbke 1989). ...
... The enchytraeids were identified to species level. The determinations made by Birkemoe and Dózsa-Farkas (1994) of Henlea similis and Bryodrilus ehlersi glandulosa were corrected later by Dózsa-Farkas (1999), the two species being confused with Henlea glandulifera and Bryodrilus diverticulatus, respectively. These corrections were based on examination of a large material from Svalbard, Greenland, Siberia, and Estland, as well as criteria given by Rota et al. 1998. ...
... The enchytraeids studied here are common throughout the Arctic. Henlea glandulifera, B. parvus, B. diverticulatus and Mesenchytraeus argentatus have a northern distribution, whereas H. perpusilla and H. ventriculosa have a much wider geographic distribution (Nielsen and Christensen 1959;Nurminen 1973aNurminen , 1973bBirkemoe and Dózsa-Farkas 1994;Rota et al. 1998;Dózsa-Farkas 1999). The worms from our Arctic populations of H. perpusilla and H. ventriculosa are shorter at maturity (i.e., have fewer segments) than worms from more southerly populations (Nielsen and Christensen 1959). ...
We report the results of the first study of the population dynamics and life cycles of Arctic enchytraeid populations. Sampling was undertaken in a Salix heath in Adventdalen, Svalbard, during one summer and the succeeding spring. In addition, a Cassiope heath at a more northerly site close to Ny-Alesund, Svalbard, was sampled twice. The Arctic enchytraeids were generally smaller at maturity than their temperate-zone relatives. The three most numerous species in the Salix heath, Henlea perpusilla, Henlea glandulifera, and Bryodrilus parvus, hatched from cocoons in early summer and attained adult size early in their second summer. A few H. perpusilla and H. glandulifera reached mature size in their first summer; since the summer of investigation was unusually cold, these species may have a 1-year life cycle in warmer years. Life cycles were apparently longer in the Cassiope heath than in the Salix heath. Henlea perpusilla, H. glandulifera, and B. parvus produced eggs throughout the summer in the Salix heath, though hatching was restricted to early summer. Therefore, the hypothesis that cocoons require a cold period to hatch was tested in a laboratory experiment. When soil containing cocoons was incubated at -5 degreesC for 3 weeks, a significant increase in juveniles was demonstrated for H. perpusilla and Bryodrilus diverticulatus compared with soils kept at constant summer temperatures. This is the first time that breaking of dormancy by an external stimulus has been demonstrated in enchytraeid cocoons.
... Enchytraeid life cycles vary with species and habitat. Reproduction may occur throughout the year or during a short period of time (Springett 1970;Römbke 1989;Dózsa-Farkas 1999), with corresponding variation in lifecycle length (Springett 1970;Klungland 1981;Römbke 1989). ...
... The enchytraeids were identified to species level. The determinations made by Birkemoe and Dózsa-Farkas (1994) of Henlea similis and Bryodrilus ehlersi glandulosa were corrected later by Dózsa-Farkas (1999), the two species being confused with Henlea glandulifera and Bryodrilus diverticulatus, respectively. These corrections were based on examination of a large material from Svalbard, Greenland, Siberia, and Estland, as well as criteria given by Rota et al. 1998. ...
... The enchytraeids studied here are common throughout the Arctic. Henlea glandulifera, B. parvus, B. diverticulatus and Mesenchytraeus argentatus have a northern distribution, whereas H. perpusilla and H. ventriculosa have a much wider geographic distribution (Nielsen and Christensen 1959;Nurminen 1973aNurminen , 1973bBirkemoe and Dózsa-Farkas 1994;Rota et al. 1998;Dózsa-Farkas 1999). The worms from our Arctic populations of H. perpusilla and H. ventriculosa are shorter at maturity (i.e., have fewer segments) than worms from more southerly populations (Nielsen and Christensen 1959). ...
We report the results of the first study of the population dynamics and life cycles of Arctic enchytraeid populations. Sampling was undertaken in a Salix heath in Adventdalen, Svalbard, during one summer and the succeeding spring. In addition, a Cassiope heath at a more northerly site close to Ny-Ålesund, Svalbard, was sampled twice. The Arctic enchytraeids were generally smaller at maturity than their temperate-zone relatives. The three most numerous species in the Salix heath, Henlea perpusilla, Henlea glandulifera, and Bryodrilus parvus, hatched from cocoons in early summer and attained adult size early in their second summer. A few H. perpusilla and H. glandulifera reached mature size in their first summer; since the summer of investigation was unusually cold, these species may have a 1-year life cycle in warmer years. Life cycles were apparently longer in the Cassiope heath than in the Salix heath. Henlea perpusilla, H. glandulifera, and B. parvus produced eggs throughout the summer in the Salix heath, though hatching was restricted to early summer. Therefore, the hypothesis that cocoons require a cold period to hatch was tested in a laboratory experiment. When soil containing cocoons was incubated at -5°C for 3 weeks, a significant increase in juveniles was demonstrated for H. perpusilla and Bryodrilus diverticulatus compared with soils kept at constant summer temperatures. This is the first time that breaking of dormancy by an external stimulus has been demonstrated in enchytraeid cocoons.
... Four species of Annelida not previously recorded from Svalbard were collected; two enchytraeids, Cognettia glandulosa (Michaelsen, 1888) and Enchytraeus dichaetus Schmelz and Collado, 2010 and two lumbricids, Dendrodrilus rubidus (Savigny, 1826) sensu Sims and Gerard (1985) and Dendrobaena hortensis (Michaelsen, 1890) (Table 1). More than 30 species of enchytraeids have been recorded from all types of soil habitat in Svalbard (Dózsa-Farkas 1999) but the two species collected here are likely to have been recently introduced. Enchytraeus dichaetus was originally described from Mediterranean soils. ...
Forty six species of invertebrate were collected from the manure enriched imported soils below the abandoned cow sheds in the Russian mining town of Barentsburg, Svalbard. Of these, 11 (24 %) were new records for Svalbard, including Collembola, gamasid mites, Enchytraeidae and the first identified Lumbricidae. Many of the new records are species not frequently observed in the Arctic. It is hypothesized that these species arrived with the chernozem soils imported to Barentsburg for the greenhouses from central or southern European Russia, or with livestock. The observations presented here are the first records of human invertebrate introductions establishing in Sval-bard outside of dwellings. It is not believed that the majority of new species records described present an immediate threat to the ecology of Svalbard but they may, especially Deuteraphorura variabilis, establish in the nutrient enriched floral communities beneath bird cliffs characteristic of Svalbard.
... Two species, Enchytraeus dichaetus and Cognettia glandulosa, are not known from other locations in Svalbard and are considered to be recent human introductions with the imported soils ( Coulson et al. 2013). In total, more than 30 species of enchytraeids have been recorded from all types of soil habitats in Svalbard (Nurminen 1965;Dó zsa-Farkas 1999) and abundance can exceed 50 000 individuals per square metre under favourable conditions (Birkemoe 1995). Excluding the two introduced species, six species were identified, similar to species diversity at other locations around Svalbard (Dó zsa-Farkas 1999; Birkemoe et al. 2000). ...
... ) (Table 1). More than 30 species of enchytraeids have been recorded from all types of soil habitat in Svalbard (Dózsa-Farkas 1999) but the two species collected here are likely to have been recently introduced. Enchytraeus dichaetus was originally described from Mediterranean soils. ...
Forty six species of invertebrate were collected from the manure enriched imported soils below the abandoned cow sheds in the Russian mining town of Barentsburg, Svalbard. Of these, 11 (24 %) were new records for Svalbard, including Collembola, gamasid mites, Enchytraeidae and the first identified Lumbricidae. Many of the new records are species not frequently observed in the Arctic. It is hypothesized that these species arrived with the chernozem soils imported to Barentsburg for the greenhouses from central or southern European Russia, or with livestock. The observations presented here are the first records of human invertebrate introductions establishing in Sval-bard outside of dwellings. It is not believed that the majority of new species records described present an immediate threat to the ecology of Svalbard but they may, especially Deuteraphorura variabilis, establish in the nutrient enriched floral communities beneath bird cliffs characteristic of Svalbard.
Alpine freshwater biodiversity is at risk of extinction due to climate change and some species could remain undescribed before they go extinct. These species are not yet included in red lists of protection by law, such as annelids in alpine habitats. Within this context, we studied the annelid fauna in 20 sites located between 1223 and 2703 m asl, belonging to different habitat types (kryal, glacio-rhithral, non-glacial streams, springs, littoral lake zone), in three glaciated catchments of the Italian Alps (Noce Bianco, Careser and Conca-Carè Alto; Trentino Province). The aim of this study was to describe the annelid communities of the different habitat types collected with different sampling methods (kick sampling and drift) and investigate the relationships between species richness and abundance with altitude. Between the years 2000 and 2005, we collected 4,765 individuals in 418 samples. One genus of Polychaeta (Aeolosoma) and 36 species of Oligochaeta were identified, distributed in four families: Enchytraeidae (28), Lumbriculidae (4), Naididae (3 and Tubificinae juveniles) and Haplotaxidae (1). Five species were new to Italy: Cernosvitoviella carpatica, Cernosvitoviella cf. crassoductus, Henlea brucei, Henlea glandulifera and Mesenchytraeus sanguineus. As expected, Enchytraeidae prevailed in all sites with the genera Cernosvitoviella and Cognettia the most abundant and frequent. The regression and cluster analyses and the generalized linear mixed models we performed, highlighted that the differences in species richness and abundance among sites are explainable partly by altitude but mainly by habitat type. A clear longitudinal species turnover was evident only in the Noce Bianco and Careser glacier-fed streams, where taxonomic diversity (tested by Seriation analysis) and abundance increased with increasing distance from the glacier front and decreasing altitude. Also expected, the harsh kryal habitat was the poorest in species and individuals, with a dominance of semi-aquatic and terrestrial enchytraeid taxa. Drift was found to be more effective than kick sampling in collecting oligochaetes, especially in glacier-fed streams where the use of the pond net is generally hindered by their high and highly variable discharge. In conclusion, this study provides new data on oligochaete fauna in alpine freshwaters, useful to implement the prevision models for Alpine biodiversity up to date not including annelid fauna.
The taxonomic status of the enchytraeid species Bryodrilus parvus Nurminen, 1970 and Bryodrilus librus (Nielsen and Christensen, 1959) was tested with respect to their possible conspecifity. B. parvus, considered as a typical northern Hol-arctic species, was also found in Hungarian lowlands. The species B. librus (Nielsen and Christensen, 1959), distributed in Northern and Central Europe, is very close morphologically to B. parvus. In this paper we demonstrate that B. parvus and B.librus are not different species based on both morphological and molecular analyses of specimens from different regions in Europe. The valid name of the species is Bryodrilus librus (Nielsen & Christensen, 1959) following the princi-ple of priority (ICZN 1999, Art. 23.1).The species B. librus was revised and a new description is given here.
Marionina argentea (Michaelsen, 1889) is a taxon recorded throughout the Holarctic, from the warmer areas of the Mediterranean to all around the Arctic Ocean. However, following an in-depth and extensive comparative study (more than 100 specimens examined as permanent mounts, originating from the Mediterranean to Sweden), three species new to science have turned out to be hidden and confounded under its name. All of them are widely distributed in Europe. Marionina mendax sp. n. is mainly characterized by its large chaetal size, the coarse granulation of coelomocytes and the small penial bulbs. Marionina deminuta sp. n., in addition to its smaller body size and peculiar variation in the position of the genital organs (shifted cephalad by one segment in most specimens), consistently deviates from M. argentea sensu stricto in the pattern of the clitellum, the shape of the nephridia, and the relative size of the penial bulbs. Marionina mimula sp. n. differs from all the above species by having a posteriorly indented brain, a lumbricilline arrangement of the dorsal vessel, and a rosette of gland cells surrounding the spermathecal pore. The confused history and different interpretations of the diagnosis of M. argentea are reviewed and discussed. An interim revision of the morphological boundaries of the nominal species is proposed, pending re-examination of the type series of Enchytraeus argenteusMichaelsen, 1889.
The terrestrial environment of the High Arctic consists of a mosaic of habitat types. In addition to the natural habitat diversity, various human-influenced types may occur. For the resident invertebrate fauna, these anthropogenic habitats may be either unusually favourable or detrimental. In the town of Barentsburg, Svalbard, soils were imported for the greenhouses from southern Russia. These soils were subsequently discarded outside the greenhouses and have become augmented with manure from the cowsheds. Both the greenhouse and the cowsheds are now derelict. This site represents an unusually nutrient-rich location with considerable development of organic soils, in stark contrast to the naturally forming organic soils in Svalbard, which are typically thin and nutrient poor. Few previous studies have examined the soil invertebrate communities of human-disturbed or -created habitats in the Arctic. In an often nutrient-poor terrestrial environment, it is unclear how the invertebrate fauna will react to such nutrient enhancement. In these soils, 46 species of invertebrates were determined. Eleven species have not been recorded from other habitats in Svalbard and are hence likely to have been introduced. The native species assemblage in the anthropogenic soils was not atypical for many natural sites in Svalbard. Despite the enriched organic soils and highly ameliorated winter temperature conditions, the soil invertebrate fauna biodiversity does not appear to be enhanced beyond the presence of certain probably introduced species.
Survival of enchytraeids from three areas was investigated: an oak forest (Quercus pubescens) in the vicinity of Vienna, Austria (420 m a.s.l., 48°16′N/16°21′E), a subalpine meadow in Lower Austria (1580 m a.s.l., 47°43′N/15°47′E) and an Arctic tundra heath at the Kongsfjorden, Spitsbergen (Ny-Ålesund: 78°30′N/11°5′E). In the two Austrian areas snow cover protected the enchytraeids during the 1998/99 winter from frost conditions in the soil. In laboratory experiments and in the field, worms developed from cocoons as well after drought as after subzero temperature exposure. The supercooling point (SCP) of individual worms was between –3.8 and –13.6 °C. A significant correlation existed between the quantity of gut content and survival at low temperatures. Most enchytraeid species, both in the field and in the laboratory, consumed more food at temperatures above 0 °C and most worms which survived frost, had no gut content.
We wished to test the taxonomic status of the enchytraeid species Bryodrilus glandulosus (Dózsa-Farkas, 1990) – previously B. ehlersi glandulosus – and Mesenchytraeus kuehneltiDózsa-Farkas, 1991, and their eventual conspecifity with, respectively, B. ehlersiUde, 1892 and Mesenchytraeus pelicensisIssel, 1905. The two species were both described from Hungarian Sphagnum-bogs as relict species of these unique areas. In this paper we demonstrate that these species are invalid based on morphological and molecular analyses of different European specimens.The only differential character of B. glandulosus from B. ehlersiUde, 1892 was the presence of glands at the opening of the spermathecae. However, during the thorough investigation of European B. ehlersi species we realized that contrary to the original description, all B. ehlersi have these glands. Moreover, there were no differences between the ITS gene sequences of the B. ehlersi specimens collected from different places.A similar ITS study did not find any difference between M. kuehnelti and M. pelicensis either. Their morphological differentiation was based on the presence of the atrial glands but this separation has no legitimacy since this is a variable character.For these reasons we revise the species referred and give new descriptions.
The aim was to study the invasion of terrestrial enchytraeids into the two postglacial tundras, North-eastern Greenland and the Arctic Archipelago of Canada. The majority of species recorded have a wide distribution in the Northern Hemisphere and the species diversity is considerably lower than that of potential source regions indicating either strong selection or varied dispersal ability. It appears that the Archipelago is influenced mainly from North America and North-eastern Greenland from Europe while the specialized fauna of the ancient Beringian tundra is of minor importance. The two alternative scenarios: (a) survival of a prepleistocene fauna in protected refugia within the area or (b) a postglacial re-invasion from outside are discussed, but the available data do not discriminate between these two possibilities. A total of 24 terrestrial enchytraeid taxa are recorded of which 22 are identified to the species level. Bryodrilus archipelagicus and Hemifridericia bivesiculata are described as new to science and Henlea groenlandica Černosvitov is revalidated. We hesitate to conclude that the new taxa described are endemic.
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