Folia Fac. Sci. Nat. Univ. Masaryk. Brun., Biol., 110 (2007): 123-133

Newsletter on No. 10: Proceedings of the 7 th International Symposium on Enchytraeidae; May 25-28, 2006, Brno, Czech Republic Edited by J. SCHLAGHAMERSKÝ

The small (Enchytraeidae, Tubificidae, Aeolosomatidae) of a South Moravian floodplain forest with disturbed water regime

JI ŘÍ SCHLAGHAMERSKÝ

Department of Botany and Zoology; Faculty of Science; Masaryk University, Kotlá řská 2; 611 37 Brno; Czech Republic; e-mail: [email protected]

ABSTRACT

The community of small annelids was studied in a hardwood floodplain forest on the River Dyje in South Moravia (Czech Republic). The site has not been flooded since 1972 due to river regulation. Four samplings between Oct. 2001 and July 2005 yielded data on density, vertical distribution and community structure in soil. Additionally tree humus under bark was sampled. Densities in soil ranged between 5 000 and 16 500 ind./m 2 with 38-69 % present in the upper 3 cm of soil. The community was dominated by the enchytraeid genera Fridericia (40 % of individuals) and (20 % of individuals). In total 28 enchytraeid species, Rhyacodrilus falciformis (Tubificidae) and Aeolosoma cf. niveum (Aeolosomatidae) were found; some identifications remained preliminary. Achaeta pannonica , Fridericia benti and Stercutus niveus are reported from the Czech Republic for the first time. Marionina argentea showed significant preference for moist terrain depressions. Tree humus hosted a species-poor assemblage with varying dominances of Enchytraeus buchholzi s.l. and Henlea nasuta .

Keywords: tree humus, soil zoology, fluvisol, South Moravia, Czech Republic

INTRODUCTION

Little has been published on enchytraeid assemblages in floodplains and even less on those in floodplain forests. The most comprehensive report on assemblages of small annelids in floodplain forests has been presented by BEYLICH & GRAEFE (2007) and is based on four hardwood and one softwood (in Central European terminology) floodplain forest sites on the Middle Elbe (Germany). Additionally, these authors present 124 JI ŘÍ SCHLAGHAMERSKÝ data on a high number of sites in other floodplain habitats. Enchytraeids were also studied in fen habitats on peat soil in the Biebrza river basin in Poland, including an alder carr as the only forest habitat (NOWAK & PILIPIUK, 1997; STERZYŃSKA & PILIPIUK, 1999). A brief paper by GRAEFE (1998) on communities of wet soils, including three Vega-Gley soils under grasslands on the river Elbe (Germany), neither includes a complete species list nor quantitative data. Further data on enchytraeids in floodplain grasslands in Germany and the Netherlands were published by MÖLLER (1967, 1971), VAN VLIET & DIDDEN (2003) and by PLUM & FILSER (2005). One of the ecosystem research projects conducted under the International Biological Programme (IBP) and later the Man and the Biosphere (MaB) programme of the UNESCO had focused on the floodplain forests in South Moravia, Czech Republic. Unfortunately, regarding soil fauna proper, only earthworms were studied within that project (ZAJONC, 1985). Later, earthworms were studied again in the principal research plot of the UNESCO projects, as well as at other sites in South Moravian floodplains (PIŽL, 1998, 1999). The present study provides quantitative and qualitative data on the assemblage of small annelids existing today in the principal research plot that had been intensively studied within the IBP and MaB projects. This assemblage includes not only a rather high number of enchytraeid species, but also representatives of the Tubificidae and Aeolosomatidae. These two families are generally considered aquatic but some species occur in soil as well, although published records are scarce (cf. MELLIN, 1988; SCHLAGHAMERSKÝ, 1998; SCHLAGHAMERSKÝ, 2005). Attention is also paid to the micro-habitat under bark of dead trees usually neglected in soil-zoological studies. As described in more detail below, the study site has undergone substantial changes regarding its water regime, in particular its flood exposure. Since the early 1970s the site has developed into a much drier habitat and it is a question of definition if the term floodplain forest still applies to the present state. However, even under natural river dynamics, floodplains include sites of different flood exposure and the conditions on individual sites develop over time. As most other European “floodplain” forests also have a disturbed water regime, the present study can still be considered as providing valuable information on enchytraeids and other small annelids of floodplain forests. The study site is situated within the Lower Morava Biosphere Reserve (the recently enlarged former Pálava Biosphere Reserve) for which a unique inventory of invertebrates has been published (ROZKOŠNÝ & VA ŇHARA, 1995a, 1995b, 1996; OPRAVILOVÁ et al., 1999). However, enchytraeids were one of the few taxa not covered by this inventory. Generally, our knowledge of the enchytraeid fauna in the Czech Republic is rather limited, reliable data being restricted to a small number of sites. By providing information on the community structure of the terrestrial small annelid assemblage in a South Moravian hardwood floodplain forest, the present contribution should partially fill in the outlined gaps in our knowledge on the small annelid fauna of European floodplain forests in general, of the Czech Republic and of the Lower Morava Biosphere Reserve in particular.

The small annelids of a South Moravian floodplain forest with disturbed water regime 125

MATERIALS AND METHODS

Study Site

The research plot of 6.9 ha was set up in 1968, having been under special management since the 1950s (VYSKOT, 1976). It is situated 1.5 km north-west of the village Lednice in South Moravia (Czech Republic) at 161 m a.s.l. (48 °48´N, 16 °46´E). The climate is subhumid to arid with a mean annual air temperature of 9 °C and a mean annual precipitation of 524 mm (means for 1901-1950; QUITT, 1971); the decade preceding the study period was drier, i.e. the temperature for the vegetation period (April-November) in 1987-1996 exceeded the long-term mean for this period by 1 °C, while the total annual precipitation values were lower than the long-term mean (BAGAR & KLIMÁNEK, 1999). The soil was brown alluvial soil (fluvisol) of Vega type overlaying an uneven relief of Quaternary sediments (several metres of Holocene alluvial loam strata above 5-8 m thick Pleistocene gravel terraces). Sandy loam of heavy texture topped by mull humus formed the upper two metres of the soil profile. Percentages of humus, carbon, and nitrogen were 56.4, 32.7 and 2.4 in the F (1-2 cm), and 10, 5.8 and 0.8 in the A mu horizons, respectively (GRUNDA et al., 1991). The plot was covered by an approximately 125-year-old hardwood floodplain forest stand (in Central European terminology) of the association Fraxino pannonicae-Ulmetum. The tree layer was dominated by Quercus robur (50 %), Fraxinus angustifolia (35 %) and Tilia cordata (10 %), the herb layer by Urtica dioica, Impatiens parviflora, Pulmonaria officinalis , and Geum urbanum. The stand was part of a forest covering the south-eastern part (275 ha) of a large island located between the main river channel of the Dyje and its artificial branch (“Zámecká Dyje”, 130 m from the plot boundary). The hydrological regime of the area has been disturbed by river regulation (up-stream construction of the Nové Mlýny Water Reservoirs on the Dyje river and construction of levees along the rectified main river channel); the site had been flooded regularly, mostly in early spring, the last inundation occurring in 1972. Since the ceasing of floods the site has been drying out, which was accompanied by changes in its flora and fauna (VA ŇHARA, 1994; DROZD, 1997). Major changes regarding the soil compartment were (GRUNDA et al., 1991): a drop in the ground water table and soil moisture content; ceasing of the water table and decomposition dynamics driven by the flooding of the site; much higher dependence of the ecosystem on local precipitation; ceasing of the river-born input of silt and nutrients; increased aeration of the upper soil leading to improved conditions for aerobic microorganisms and thus a reduction of the humus content of the A mu horizon (2- 8 cm) by about a sixth; somewhat increased nitrogen content in the upper soil and thus also a decrease in the C/N ratio. Soil pH, a major factor for soil organisms, was not modified by the changes in site hydrology: in the transitional mollic-umbric humus-rich horizon (A mu : 2-8 cm) it was about 6.5 (H 2O) and 5.4 (KCl), the values for the upper organic layer and lower mineral layer being very similar. At the time of the present study measures were taken that should lead to some rewetting of the site in future.

126 JI ŘÍ SCHLAGHAMERSKÝ

Sampling and Data Analysis

Soil samples were taken on October 8, 2001, February 25, 2002, October 2, 2002, and July 14, 2005. The original objective of the first two samplings was the comparison of modifications of the wet funnel extraction method (KOBETI ČOVÁ & SCHLAGHAMERSKÝ, 2003). Therefore soil cores were taken in triplets (8 sample units in Oct. 2001) or pairs (12 sample units in Feb. 2002), the individual cores of one sample unit (triplet or pair of soil cores) being placed in 5-10 cm distance from each other. Equal numbers of sample units were positioned randomly within two areas of 20 m2, one representing a dense nettle stand, the dominant facies of herb vegetation. Samples were taken by a cylindrical steel corer of 10 cm 2 surface area down to 8-9 cm depth in Oct. 2001 and 6 cm in Feb. 2002 (the dense soil prevented deeper sampling). Cores taken within one triplet or pair were subjected to different modifications of wet funnel extraction. As no significant differences in extraction efficiency were found, these sample units were pooled for the computation of the mean densities and standard errors presented below. These values are thus based on rather large samples when compared to the usual sampling schemes employed for enchytraeid studies. The samplings in Oct. 2002 and July 2005 were conducted in particular to clarify several species identities, to get more precise data on community structure and vertical distribution and to look at differences between distinct micro-habitats. Based on the low densities encountered previously a steel corer of 17 cm 2 surface area was used (also allowing deeper penetration) and soil cores were taken down to 12 cm depth: 8 and 10 cores were taken in Oct. 2002 and July 2005, respectively, within an area of 20 m 2 (stratified random sampling regarding herb layer and geomorphology: three soil cores were taken in a moist rill on both occassions, two cores were taken at the base of standing trees in 2005). Additionally to soil sampling, tree humus accumulated at ca. 1 m height in pockets under loose bark of an ash tree snag was collected. At all four sampling dates the soil cores were vertically subdivided into 3 cm layers, which were placed in separate containers for transport and storage. Live specimens were obtained from the soil layers and the tree humus by wet funnel extraction (see KOBETI ČOVÁ & SCHLAGHAMERSKÝ, 2003; in Oct. 2002 and July 2005 12 hours of cold extraction were followed by heating up to 43 °C surface temperature within 4 hours; tree humus was subjected to cold extraction only). Extracted specimens were kept in Petri dishes with tap water at ca. 5 °C, counted, and identified alive under a light microscope. Species records based on a few enchytraeids hand-collected at the site from under loose bark of a lying oak trunk in 1996 are also reported as these species were not found by the other methods employed. In total, 821 enchytraeid specimens and 41 specimens of other small annelids were extracted and microscoped. Mean density and vertical distribution data were computed based on all the soil samples taken per sampling, including those from the rill or tree base. For the dates on which soil cores were taken both in a moist terrain depression and on higher lying ground the Mann-Whitney-U-Test was applied to abundance data of those species that appeared to prefer one of these situations.

The small annelids of a South Moravian floodplain forest with disturbed water regime 127

RESULTS

Mean enchytraeid densities in soil (individuals/m 2; values rounded to the nearest hundered) per sampling date ranged between 5 000 ± 900 in Feb. 2002, and 16 500 ± 3 000 in Oct. 2002 (Figure 1). Densities obtained during the first two samplings (Oct. 2001, Feb. 2002) were less then half those found later and also the absolute number of specimens was lower in spite of the larger sample size. However, the density values of the two first dates are based on the sampling of the upper 9 or 6 cm only and the same applies to the percentages of individuals per soil layer. The highest numbers (38-69 %) were always found in the upper-most 3 cm of soil (Figure 1). While substantial percentages of individuals were also present in the 3-6 cm layer in Feb. and Oct. 2002, numbers decreased steeply towards the deeper layers investigated.

0% 20% 40% 60% 80% 100% 0% 20% 40% 60% 80% 100%

0-3 cm 0-3 cm

3-6 cm 3-6 cm

6-9 cm Oct. 2001 6-9 cm Feb. 2002 n = 125 n = 94 9-12 cm 7 100 ± 2 300 ind./m2 9-12 cm 5 000 ± 900 ind./m2

0% 20% 40% 60% 80% 100% 0% 20% 40% 60% 80% 100%

0-3 cm 0-3 cm

3-6 cm 3-6 cm

6-9 cm Oct. 2002 6-9 cm July 2005 n = 227 n = 273 9-12 cm 16 500 ± 3 000 ind./m2 9-12 cm 16 100 ± 3 700 ind./m2

Figure 1: Vertical distribution of enchytraeids in the upper 12 cm of soil at the four sampling dates (sampling in Oct. 2001 down to 9 cm and in Feb. 2002 down to 6 cm only), total densities ± SE and numbers of individuals (n) recovered by extraction are given.

In total 28 enchytraeid species, Rhyacodrilus falciformis (Tubificidae) and Aeolosoma cf. niveum (Aeolosomatidae) were present (Table 1). The most dominant genera were Fridericia (40 % of individuals, represented by at least 12 species) and Enchytraeus (20 % of individuals, 2 species). At the last two sampling dates, Marionina argentea was found at the site, showing a preference for the moister situation in terrain depressions (two-sided Mann-Whitney-U-Test, p < 0.05). Rhyacodrilus falciformis also seemed to prefer this microhabitat but this was not statistically significant (just surpassing the significance level [ α = 0.05] for the one-sided Mann-Whitney-U-Test). The other distinct 128 JI ŘÍ SCHLAGHAMERSKÝ

Table 1: Species list of small annelid taxa with numbers of individuals broken down according to sampling date and microhabitat (soil vs. tree humus) for the Lednice floodplain forest research plot (South Moravia, Czech Republic), new records for the Czech Republic underlined, total numbers of individuals and species given for enchytraeids and the other taxa separated by “+”.

micro-habitat soil tree humus Oct. Feb. Oct. July May Oct. July Sampling Date 2001 2002 2002 2005 1996 2002 2005 Enchytraeidae Achaeta pannonica Graefe, 1989 1 Achaeta sp. 1 1 1 Achaeta sp. 2 1 (Achaeta spp.) 23 4 8 5 Bryodrilus ehlersi Ude, 1892 1 Buchholzia appendiculata (Buchholz, 1862) 1 8 1 3 Buchholzia fallax Michaelsen, 1887 3 Buchholzia spp. 4 6 31 Cernosvitoviella cf. minor Dozsa-Farkas, 1990 1 2 Cernosvitoviella spp. 16 Enchytraeus buchholzi s.l. Vejdovsky, 1879 4 6 17 5 1 37 Enchytraeus cf. lacteus N. & Ch., 1961 1 7 (Enchytraeus spp.) 7 11 25 60 2 21 Enchytronia cf. christenseni Dózsa-Farkas, 1970 2 Enchytronia parva N. & Ch., 1959 1 1 (Enchytronia spp.) 1 15 1 Fridericia cf. auritoides Schmelz, 2003 3 Fridericia benti Schmelz, 2002 3 Fridericia bisetosa (Levinsen, 1884) 1 1 10 Fridericia connata Bretscher, 1902 2 1 1 Fridericia galba (Hoffmeister, 1843) 4 2 5 8 Fridericia isseli Rota, 1994 1 3 Fridericia nemoralis Nurminen, 1970 5 4 5 Fridericia paroniana Issel, 1904 1 Fridericia perrieri (Vejdovský, 1878) 8 2 Fridericia ratzeli (Eisén, 1904) 2 3 3 3 Fridericia semisetosa Dózsa-Farkas, 1970 2 3 26 9 Fridericia sp. 1 3 (Fridericia spp .) 35 27 31 63 3 Henlea nasuta Eisen, 1878 5 2 27 (Henlea spp.) 1 Marionina argentea Michaelsen, 1889 33 13 Marionina communis N. & Ch., 1959 15 Marionina spp. 6 Mesenchytraeus cf. gaudens Cognetii, 1903 6 Stercutus niveus Michaelsen, 1888 15 6 6 (Enchytraeidae – not identified) 15 6 16 14 5 1 Tubificidae Rhyacodrilus falciformis Bretscher, 1901 5 20 12 Aeolosomatidae Aeolosoma cf. niveum Leidy, 1865 4 Total number of individuals 125+5 94 225+24 273+12 7 38 59 Total number of species: 13+1 11 15+2 20+1 2 3 1 The small annelids of a South Moravian floodplain forest with disturbed water regime 129

microhabitat investigated was tree humus. Henlea nasuta was found almost exclusively in this situation. Here it was frequent and eudominant in the Oct. 2002 sample. In July 2005 only Enchytraeus buchholzi s.l. was found in tree humus (sampled on the same tree snag at the same height), being very abundant. A single specimen of H. nasuta was extracted from a soil sample collected on the base of this tree snag on that date and another one from soil under an Impatiens parviflora stand. Mesenchytraeus cf. gaudens (6 not fully mature specimens) and Bryodrilus ehlersi (1 specimen) had been hand- collected under the loose bark of a fallen oak trunk in May 1996 and not found in later samplings. One adult Achaeta specimen lacking bottle-shaped glands was unambiguously identified as Achaeta pannonica , a new species for the Czech Republic. Another two species are published here from the Czech Republic for the first time: Fridericia benti , a recently described species (SCHMELZ, 2003) hitherto hidden in the F. bulbosa species complex (I found several specimens also near Černotín on the Be čva river, central Moravia, in 2004; unpublished) and Stercutus niveus (I found the species also in an old- growth beech forest, the Sidonie National Nature Reserve, White Carpathians Protected Landscape Area, eastern Moravia, in 2001; unpublished).

DISCUSSION

The computed mean densities (per sampling date) of enchytraeids are rather low, for instance when compared to data presented for similar sites by BEYLICH & GRAEFE (2007). Insufficient sampling depth in Oct. 2001 and particularly Feb. 2002 might have led to underestimates. The high percentage of individuals in the 3-6 cm layer in Feb. 2002 indicates that a substantial portion of the populations might have been present in lower layers at that time. On the other hand, the vertical distribution at the other sampling dates shows a substantial decrease with depth and a sampling depth of 12 cm seems fully sufficient for assessing the density. The inclusion of samples taken in terrain depressions and at tree bases might have affected the density values, however, for instance the July 2005 density remains approximately the same when the two tree base samples are excluded. Taken into account the dry climate of the study area, the July 2005 density was surprisingly high and so was the percentage of individuals in the uppermost 3 cm of soil (the highest of all sampling dates). One possible explanation is that the frequency and amount of precipitation were rather high in summer 2005, particularly when compared to the preceding years. During the summer, the enchytraeid species Stercutus niveus undergoes aestivation (aneonosis) and is not extracted from soil in this inactive stage (DÓZSA-FARKAS, 1973). Indeed, no specimen was recovered at this time. This might have also led to some underestimation of the July density (not activity). Enchytraeid density and vertical distribution are known to undergo substantial changes so that mean annual densities should be based at least on year-round sampling (DIDDEN, 1993). However, the available data indicate that the mean annual density of enchytraeids lies between 5 and 20 thousand individuals per square meter (the inclusion of the other small annelids present would not change this estimate). This confirms results from other sites with only slightly acidic soil at which enchytraeids were shown to reach low densities but at the same time higher species diversity than in other habitats 130 JI ŘÍ SCHLAGHAMERSKÝ

(DIDDEN, 1993; MELLIN, 1988; SCHLAGHAMERSKÝ, 1998). The hardwood floodplain forest sites studied by BEYLICH & GRAEFE (2007) were more acidic, which might explain both their higher densities and lower species numbers. With probably more than 28 enchytraeid species, the site already comes close to the species numbers of the species-richest sites published (compare for instance with data on floodplain meadows in MÖLLER, 1971). The species number is substantially higher than the numbers (16-22) found at the four comparable hardwood floodplain forest sites (Elbholz) by BEYLICH & GRAEFE, 2007). However, besides the lower pH of the Elbholz sites, also the fact that the species numbers presented by the latter authors are based on a single sampling date per site might have led to lower values. In the present study low densities prevented the investigation of sufficiently large material of mature specimens to resolve several taxonomic problems, despite the additional samplings conducted. It was unfortunate that the revision of the genus Fridericia by SCHMELZ (2003) was only available for the identification of material from the last sampling date. The first two samplings had been primarily aimed at receiving quantitative data to compare extraction methods and the time to identify the large material to species was limited. Also the sampling of tree humus did not yield additional specimens of Mesenchytraeus cf. gaudens or Bryodrilus ehlersi, which had been collected rather accidentally in 1996 under bark of a fallen oak trunk. The occurrence on decomposing wood is typical of B. ehlersi . Mesenchytraeus species grow rather large and occur in the upper litter layer, being thus collected best when this layer is wet. However, the conditions at the site do not allow the permanent formation of a thick litter layer. Micro-habitats connected to wood decomposition are usually neglected in studies on terrestrial enchytraeids but can be important for the actual species richness of a site; their investigation also provides additional information on the ecology of individual species. The species diversity of soil-dwelling annelids in the floodplain forest is further increased by the differences between lower and higher lying ground. Three of the encountered species are considered to prefer wet soils (GRAEFE & SCHMELZ, 1999). Of these M. argentea was indeed significantly more frequent in terrain depressions, while in R. falciformis such a preference was traceable but not significant. Cernosvitoviella cf. minor was mainly found during the first two samplings when no terrain depressions were sampled. Taxonomic and faunistic notes: Probably three species of Achaeta were present. Almost all specimens had bottle-shaped glands (“setal follicles”) dorsally as well as ventrally (not always well visible in juveniles). (Sub)adults did not match any published species description. Most were further characterised by sperm funnels of a length exceeding the body width (length : width ratio of 8 : 1), three pairs of pharyngeal (septal) glands with the dorsal vessel originating behind the 3 rd pair (in VII), and spermathecae with ampullae reaching to VII or VIII ( Achaeta sp. 1 in Tab. 1). One subadult specimen (spermatheca well developed, no other sexual organs observed), with bottle-shaped glands dorsally and ventrally, was very small and otherwise resembled a newly hatched juvenile. This specimen most probably represented another species of Achaeta ( Achaeta sp. 2 in Tab. 1), differing from A. parva Nielsen & Christensen, 1961 by having three pairs of septal glands, the dorsal vessel originating behind the last one in VII. Several adult Fridericia specimens could not be identified to species, all having a spermatheca with two diverticula. Several individuals seemed to belong to one distinct The small annelids of a South Moravian floodplain forest with disturbed water regime 131

species ( Fridericia sp. 1 in Tab. 1: length ca. 10 mm; ca. 40 segments; chaetal formula: 4, 5 – 2, 3, 4 : 4, 5, 6 – 2, 3, 4, 5; few chaetae floating in coelome; origin of dorsal vessel around segment XVIII; oesophageal appendages short, unbranched or with short distal branches; spermatheca with two globular to kidney-shaped diverticula, its thick ental ducts communicating separately with the oesophagus, its ectal ducts of medium length having a small to mid-sized gland at the orifice; sperm funnel length ca. ¾ of body width, length : width ca. 1 : 2.5; large red-brown seminal vesicle in up to four segments, e.g. IX-XIV; clitellum not much elevated, gland cells arranged in transverse rows). Several Fridericia specimens were arbitrarily identified as F. cf. auritoides . The genus Enchytraeus was represented by E. buchholzi and E. cf. lacteus. E. buchholzi is listed as sensu lato as the species description probably represents a species complex (the taxonomic situation is complicated by strains reproducing by self-fertilization; see SCHMELZ et al., 1999). However the present specimens were uniform, probably belonging to a single species. The specimens identified as E. cf. lacteus differed from E. buchholzi by larger body size, large and elongate sperm funnels and large seminal vesicles. In one case the ectal glands on the ectal orifice matched better the description given for E. coronatus Nielsen & Christensen, 1959 and the situation is further complicated by the fact that some E. buchholzi strains may have enlarged sexual organs as well (SCHMELZ, pers. comm.). In case of Enchytronia some specimens were identified as E. cf. christenseni based on the presence of seminal vesicles (other characters, including the shape of the spermatheca, did not show clear differences to the description of E. parva , the other con-generic species identified). The Marionina specimens from the Oct. 2002 sampling were juveniles and subadults (i.e. probably not fully mature individuals without eggs and well developed clitellum). The chaetal bundles (with 1-2 chaetae) and further characters (e.g. “brain” not much incised caudally; no glands observed at orifice of spermatheca ectal duct) suggest that these were not immature specimens of M. communis (the other con-generic species identified, M. argentea , can not be mistaken), but data were not considered sufficient to add another species to the list.

ACKNOWLEDGEMENTS

The study was conducted under the Masaryk University’s Research Plans MSM 143100010 (work in 2001-2002) and MSM 0021622416 (subsequent work and presentation). Mgr. Klára Kobeti čová and Mgr. Adéla Šídová kindly assisted with the sampling and sample extraction.

REFERENCES

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