journal 6.indb
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journal 6.indb
POLISH JOURNAL OF ECOLOGY (Pol. J. Ecol.) 54 2 231–242 2006 Regular research paper Elżbieta DUMNICKA Institute of Nature Conservation, Polish Academy of Sciences, al. Mickiewicza 33, 31-120 Kraków, Poland, e-mail: [email protected] COMPOSITION AND ABUNDANCE OF OLIGOCHAETES (ANNELIDA: OLIGOCHAETA) IN SPRINGS OF KRAKÓWCZĘSTOCHOWA UPLAND (SOUTHERN POLAND): EFFECT OF SPRING ENCASING AND ENVIRONMENTAL FACTORS ABSTRACT: Studies were performed on 25 karstic springs located in Southern Poland (50º10’ – 50º46’ N, 19º17’ – 19º54’E). They are natural or encased, differing also in discharge rate (0.01– 15 l s–1 in the southern part of KrakówCzęstochowa Upland while 20 – 1440 l s–1 in its northern part) and kind of bottom sediments (fine or coarse). 27 species and 1 genus representing 5 families were determined, among them Enchytraeidae were represented by the highest number of species whereas Tubificidae (mainly juvenile forms) were the most abundant. Among oligochaetes no crenobionts were found while crenophiles: Rhyacodrilus falciformis Bretscher and Stylodrilus heringianus Claparède were present in a half of the studied springs. In the majority of the studied springs the density of oligochaetes was significantly higher in fine sediments (ranges 200 – 13 200 ind. m–2) than in coarse ones (ranges 22–7900 ind. m–2) (P <0.05). The species diversity (H’) was also higher in fine sediments, especially in springs of the southern part of Upland. Oligochaeta were more abundant and diversified in encased springs than in natural ones and they inhabited preferably fine sediments. The discharge and springs localization (in southern or northern part of the studied area) were found to be not of importance for oligochaete composition and abundance. KEY WORDS: karstic springs, Oligochaeta, Poland 1. INTRODUCTION Although ecological and zoological studies on springs have long tradition (D emel 1922, Thienemann 1924, Pa x and Masch ke 1936, Miche j d a 1954, O du m 1957, Thor up and L i nde ga ard 1977) taxonomic composition of many invertebrate groups living in this environment and their relationships with various environmental factors (hydrogeology, discharge, temperature, bottom character, etc.) is still little known. Therefore in last years intensive studies on benthic fauna were conducted in Europe as well as in North America (B otos ane anu 1998, Hof fsten and Ma l mq v ist 2000, Smit h et al. 2001, 2003). Oligochaetes were rarely determined during such studies, especially in these made recently in Europe (Smit h and Wo o d 2002, Smit h et al. 2003) or only a few large and easily recognized taxa (Eiseniella tetraedra, Stylodrilus sp.) were stated (L i nde ga ard et al. 1998). The knowledge of benthic fauna living in Polish springs is also incomplete (Biesi adka and C z achorowsk i 1999) while oligo- 232 Elżbieta Dumnicka chaetes fauna is almost unknown. Detailed faunistic studies were performed only in big limnocrene springs (“Niebieskie Źródła” springs situated in Central Poland) (Woj as 1972, Pie cho ck i 2000). In the first Polish paper concerning winter fauna of springs situated near Wigry Lake (North-East Poland) only Eiseniella tetraedra was mentioned (D emel 1922) and in materials collected from 12 springs situated in Śnieżnik Kłodzki Massif (Sudety Mts, S-W Poland) (Pax and Mas ch ke 1936) only 3 taxa were determined in a single spring (Eiseniella tetraedra, Bimastos tenuis (Eisen) and Cognettia sp). Up to now oligochaetes from big limnocrene karstic springs situated in Central Poland (“Niebieskie Źródła”) were elaborated in detail and results published in a separate paper (Ka h l 2000). Moreover some data concerning the occurrence of oligochaete species in springs could be found in faunistic papers (Kasprza k 1979a, b) and in ecological ones dealing with oligochaete fauna of streams and rivers, in which their springs were also elaborated (Kasprza k 1976, Kasprza k and Sz cz ęsny 1976, D umnicka 2000). The knowledge of species composition of this group living in various kind of lowland springs (karstic, moraine and others) situated in Eastern Europe (D embick ij 1972, Timm 1987) is fairly good. Recently the studies on spring fauna (including oligochaetes) have been made in Southern Europe (Me zzanotte and S ambugar 2004). Springs form specific life environment for aquatic fauna due to low, almost stable temperature, usually small area (what facilitates contacts with the surrounding soil) and low food resources – especially in reo- and helocrenes (St ar mach et al. 1976). The bottom may have very diversified character: silty, sandy or stony, large springs being partly covered by plants. The springs are from a long time under anthropogenic impact that causes not only changes in water chemistry (increase of nutrients and chloride content) but mainly changes in springs morphology due to various kinds of structures made for the facilitation of water taking. Such changes (dredgening, encasement or pipe intake) to various degree and in various ways modified life conditions of spring fauna and their effects are still little known. The aim of this study was to elaborate the composition and abundance of oligochaete fauna in karstic springs of KrakówCzęstochowa Upland in Southern Poland and to find its relationships with environmental parameters. An attempt to evaluate the impact of various kinds of spring encasing on the composition of oligochaete taxocens and species distribution was also undertaken. 2. STUDY AREA The samples were taken from 25 springs (Fig. 1) situated in the southern (springs no 1–15) and northern (springs no 16–25) parts of the Kraków-Częstochowa Upland (50º10’ – 50º46’ N, 19º17’ – 19º54’E, with mean elevation in its southern part 400–450 m and 250–300 m in the north). This Upland is formed by Jurassic limestone, covered mainly by brown soils of loess origin in the southern part and sandy or sandy-loamy soils in the northern (C he ł mick i 2001). The description of hydrogeological situation as well as characteristic of the studied springs concerning localization of each spring, its local name, water chemistry, organic matter content and granulometric composition of sediments were published by G a l as (2005). The southern springs, characterized by small discharge (0.01–15 l s–1) (D y nowska 1983, C he ł mick i 2001) are situated in the drainage area of Prądnik stream and its effluent – Sąspówka stream (drainage basin of Vistula river), whereas northern springs have distinctly higher discharge (20 – 1440 l s–1) (D y nowska 1983, C he ł mick i 2001) and are situated in the drainage area of various rivers: nos 17, 18, 19 – Pilica River; no. 25 – Przemsza River – both situated in the drainage area of Vistula River and nos 16, 20–24 – Warta River – the drainage area of Oder River. Permanent springs were chosen for the studies, but due to small amount of precipitation and very low groundwater level spring no 4 dried up in summer and spring no 23 in autumn 2003. Mainly natural springs of rheo-limnocrene or rheocrene type were studied and some modified by man to various degree and ways were also chosen: in springs nos 9, 10, 11, 12 and (partially) no 20 concrete well-heads were placed on the water outflow, springs nos 1 and 2 were enclosed Oligochaeta in karstic springs in small house–like concrete boxes and in spring no 14 water was taken in the iron pipe and outflowed through this pipe to a small basin situated 1 m below. Samples of macrobenthos were taken from this basin. 3. MATERIAL AND METHODS In 2003 samples were collected seasonally (in January/February, May, August and September/October) from 25 springs. Samples of benthic fauna were taken using two methods 233 but due to small dimensions of the majority of the studied springs the number of samples collected each time was small. Fine sediments were collected using polyethylene corer (diameter 4 cm) – each time two samples were taken. For coarse sediments hand scraper (15 × 15 cm) with mesh size 0.2 mm was used – only one sample was collected from each spring at each season. Both devices were pushed 5–6 cm into sediments. In cases when only one type of sediment was present in the spring (sand or silt in springs nos 3, 16, 18 and 23) samples were taken from the bottom of the same kind using two above mentioned methods. Oligochaetes were sorted out under a stereoscopic microscope, fixed in 4% formaldehyde and preserved in 45% ethyl alcohol. Slides in Canada balsam were prepared. The density per square meter and percentage share of each taxon were calculated for both habitats. Basing on oligochaete density cladograms showing similarity of fauna were calculated using Ward’s method. The significance of differences was calculated using non-parametric test (Mann-Whitney test). All calculations were done using STATISTICA 6.0 software package (STAT SOFT). Organic matter content was determined in fine sediments only. Beside sediment’s samples, each time the water was taken for physico-chemical analysis and the results were published by G a l as (2005). 4. RESULTS Fig. 1. Map of Kraków-Częstochowa Upland showing the distribution of the studied springs. a – springs (1–25), b – borders of the Ojców National Park, c – watershed between Vistula and Oder drainage areas. Oligochaetes have been found in 24 among 25 investigated springs. They were absent only in completely encased spring (no 1), where samples were taken from inside of deep concrete “house” from its walls. Moreover they were not found in fine sediments in springs nos 16 and 23 and in coarse sediment in springs nos 3 and 5 (Fig. 2). Animals from this group were quite abundant in almost half of the studied springs – they formed more than 20% of the benthic invertebrate community in fine sediments of 5 springs (nos 4, 5, 9, 11 and 21) and in coarse sediments of 4 springs (nos 9, 10, 11 and 14). Additionally, in fine sediments of 4 springs (nos 10, 14, 15 and 24) and in coarse sediments of 1 spring (no 24) they formed more than 10% of the benthic fauna. 234 Elżbieta Dumnicka Fig. 2. Proportional participation (in total numbers) of oligochaete families in studied springs (1 – 25) (see Fig. 1). ¤ – springs with well-heads (blue) # – strongly modified springs (red) The studied springs can be divided in a few groups depending on families dominating in the collected material: Tubificidae prevailed in samples taken with both methods from seven springs situated mainly in the southern part of the Upland (Fig. 2), moreover this family prevailed in samples taken from coarse sediments in spring no 25. They dominated in a few encased springs with high oligochaete density (nos 11, 15, 24) as well as in springs where oligochaete fauna was extremely poor (nos 7 and 23). In this last mentioned case, the calculation of the percentage share does not give good results (e.g. in corer samples taken in spring no. 7 juvenile specimens of Tubificidae were found exclusively). In the same kind of samples in spring no. 4 Eiseniella tetraedra was stated only. Family Enchytraeidae was more abundant in springs from the northern part of the Upland, whereas family Lumbriculidae prevailed in the southern part. Only in one spring (no 18), situated near the clean river with sandy bottom Propappidae (represented by Propappus volki) were the most numerous taxon in samples of both kinds. Naididae prevailed mainly in encased springs (no 2, 9, 10 and 14) moreover in scraper samples from springs 20 and 22. Results of cluster analysis (Fig 3A, 3B) showing the similarity of oligochaete taxocens in two kinds of sediments were a little different: in coarse sediments two groups of springs could be distinguished (Fig 3A) – the first one includes almost all the studied encased springs, where Naididae prevailed. In the second group, gathering mainly natural springs, one sub-group with very poor oligochaete fauna could be distinguished, the other being springs with fairly diversified oligochaete fauna. The clustering of samples taken from fine sediments divided studied springs into a few groups (Fig. 3B): the most similar were springs with very poor oligochaete fauna with the domination of Eiseniella tetraedra (nos 4 and 22) or domination of Tubificidae (nos 8, 19 and 25). Similarity of oligochaete taxocens living in coarse sediments was higher than in those from fine sediments. Basing on the composition of oligochaete fauna the springs from the northern and southern parts of the Upland were not separated. Oligochaeta in karstic springs 235 A 16 23 25 3 5 7 8 4 22 19 6 21 17 20 15 9 12 13 24 10 2 14 11 18 0 500 1000 1500 2000 2500 Linkage Distance B 16 23 3 7 22 4 21 17 19 8 25 5 18 10 6 13 14 11 12 9 15 20 24 2 0 1000 2000 3000 4000 5000 6000 7000 8000 Linkage Distance Fig. 3. Similarity of oligochaete fauna in 1 – 25 springs (see Fig. 1) inhabiting coarse (A) and fine (B) sediments. In the studied material 27 oligochaete species and 1 genus have been found (Tables 1, 2). The majority of species belonged to the family Enchytraeidae (10 species and 1 genus), families Tubificidae and Naididae were represented by 6 species each, moreover 3 species of Lumbriculidae and 1 species of Propappidae and Lumbricidae were stated. Stylodrilus heringianus and Rhyacodrilus falciformis were found in the highest number of springs (13 and 12 springs, respectively), Nais elinguis, N. communis and Marionina argentea occurred in 8 – 9 springs, Trichodrilus cernosvitovi, Tubifex tubifex and Cernosvitoviella atrata – in six springs, whereas the majority of species (19) was found in 1–5 studied 0.9 2.8 1.3 1.5 25 25 2.0 9 9 27 18 15 3 5 F 9 33 25 25 4 C 1 67 3 F 3 1 3 59 36 C 9 2¤ 3 10 70 7 7 F 1.1 12 12 75 6 F 0.8 75 25 7 C 1.9 9 9 18 54 9 F 8 0.7 20 80 C 2.8 3 3 21 21 6 21 F 9 6 9 9¤ 2.7 3 24 3 9 3 3 32 9 3 C 12 8 1 1 1 46 1 1 1 21 7 5 3 C 4 2.5 10¤ 1.7 10 40 10 40 F 1.0 2 >1 85 >1 2 2 2 5 1.1 1 >1 3 83 >1 >1 1 >1 9 >1 11¤ F C >1 >1 9 9 9 9 64 C 1.7 12¤ 2.9 10 15 5 5 5 25 5 10 F 20 The spring’s habitats without oligochaetes and in which only one taxon was found were excluded from the table (see Fig. 2). spring number (Fig. 1) Taxa Stylodrilus sp. juv. Stylodrilus heringianus Clap. Trichodrilus sp. juv. Trichodrilus cernosvitovi Hrabe Lumbriculus variegatus O.F. Műll. Lumbriculidae gen. spp. juv. Nais elinguis O.F. Műll. Nais communis Piguet Nais bretscheri Mich. Nais variabilis Piguet Nais sp. Specaria josinae (Vejd.) Pristina idrensis Sperber Rhyacodrilus falciformis Bretscher Limnodrilus hoffmeisteri Clap. Limnodrilus udekemianus Clap. Limnodrilus claparedeanus Ratzel Tubifex ignotus Štolc Tubifex tubifex (O.F. Műll.) Tubificidae gen. spp. juv. Cernosvitoviella atrata (Bretscher) Cernosvitoviella spp. juv. Buchholzia appendiculata (Buch.) Marionina argentea (Mich.) Marionina riparia Bretscher Cognettia sp. juv. Achaeta sp. juv. Henlea perpusilla Friend Enchytraeus buchholzi Vejd. Enchytraeidae gen. spp. juv. Eiseniella tetraedra (Sav.) Shannon-Wiener index H’ Table 1. Percentage share of particular taxa in total numbers of oligochaetes in fine (F) and coarse (C) sediments in springs nos 2–15 located in the southern part of the Upland (see Fig. 1). ¤ – encased springs 2.1 8 8 8 46 8 23 F 13 1.7 45 35 3 3 C 15 3 7 4 1 3 3 13 18 48 C 2.3 14¤ 2.1 37 6 13 25 19 F 1.7 1 1 1 1 66 3 17 1 1 F 8 15 2.1 5 2 3 58 3 3 2 C 13 11 236 Elżbieta Dumnicka 13 2.1 1.5 1.4 29 0.5 12 18 0.9 6 1 1 2 87 1 2 C F 1.9 20 40 20 57 88 F 13 13 50 14 C 20 17 6 F 6 25 50 25 16 C 19 1.0 50 C 50 1.9 20 20 F 40 20 20¤ 2.0 4 7 49 7 C 26 7 8 8 2.2 42 8 17 F 17 21 2.5 4 4 10 4 10 50 7 4 4 4 C The spring’s habitats without oligochaetes and in which only one taxon was found were excluded from the table (see Fig. 2). spring number (Fig. 1) taxa Stylodrilus sp. juv. Stylodrilus heringianus Clap. Lumbriculidae gen. spp. juv. Nais elinguis O.F. Műll. Nais communis Piguet Nais bretscheri Mich. Nais variabilis Piguet Propappus volki Mich. Rhyacodrilus falciformis Bretscher Limnodrilus hoffmeisteri Clap. Tubifex tubifex (O.F. Műll.) Tubificidae gen. spp. juv. Cernosvitoviella atrata (Bretscher) Cernosvitoviella tatrensis (Kowal.) Cernosvitoviella spp juv. Marionina argentea (Mich.) Marionina riparia Bretscher Cognettia sphagnetorum (Vejd.) Cognettia spp. juv. Enchytraeus buchholzi Vejd. Fridericia bulbosa (Rosa) Fridericia perrieri (Vejd.) Fridericia spp. juv. Enchytraeidae gen. spp. juv. Eiseniella tetraedra (Sav.) Shannon-Wiener index H’ 50 1.0 F 50 22 1.0 50 C 50 1.5 1 44 12 43 F 24¤ 1.1 >1 3 69 27 C 1 1.0 50 50 F 25 0.6 14 86 C Table 2. Percentage share of particular taxa in total numbers of oligochaetes in fine (F) and coarse (C) sediments in springs (Nos 16 – 25) located in the northern part of the Upland (see Fig. 1). ¤ – encased springs Oligochaeta in karstic springs 237 238 Elżbieta Dumnicka Fig. 4. Comparison of density (ind. m–2) of oligochaete taxa in natural and modified springs. A – in southern part of the Upland, B – in northern part of the Upland (see Fig. 1). Statistically significant differences are marked by asterisks: *P <0.05, ** P <0.01, *** P <0.005. Oligochaeta in karstic springs 239 Fig. 5. Density of oligochaetes in fine and coarse sediments from the studied springs (1 – 25, see Fig. 1) versus organic matter content (%) in the sediments. ¤ – encased springs springs only (Tables 1, 2). Six from 7 species occurring only in one spring were found in encased springs – most often in spring no 11 situated close to the main stream bed and overflooded sporadically by stream water. In springs situated in the southern part of the Upland higher number of taxa was found (22 species and 1 genus) than in the northern (18 species); only 13 species were common for springs of both parts of the Upland. The diversity of oligochaete fauna was also higher in southern springs and usually higher values were stated in corer samples than in those collected with a scraper (Tables 1, 2). Species found in the studied springs represent various ecological groups: among them there are crenophilic ones (Stylodrilus heringianus and Rhyacodrilus falciformis), characteristic of clean, running waters (Nais bretscheri, Pristina idrensis, Propappus volki), semi-aquatic or terrestrial (some enchytraeids and Eiseniella tetraedra), eurytopic (Tubifex tubifex, Limnodrilus hoffmeisteri, Nais elinguis) and only one stygobiont (Trichodrilus cernosvitovi). For the majority of the above mentioned species statistically significant differences of their density between encased and natural springs were stated (Fig. 4A, B). Juvenile specimens from families Tubificidae and Enchytraeidae as well as almost all abundant species, including stygobiontic and crenophilic ones were more abundant in encased springs. Only genus Cernosvitoviella preferred natural springs (Fig. 4A), whereas juvenile specimens of Fridericia and Tubifex ignotus were found exclusively in springs of this kind (Fig. 4B). The number of species found in a single spring varied from one (springs no 16 and 25) to twelve species (spring no 11). Higher number of species (6–12) was found in springs situated along Prądnik stream (nos 9–15) than in springs situated along Sąspówka stream (nos 1 – 8), where the number of species varied from zero to four (Table 1). In springs situated in the northern part of the Upland the number of determined species was usually small and did not exceed five (Table 2). The diversity was low (H’ < 2.0) in the majority of the studied springs situated in the northern part of Upland, whereas in some springs from its southern part these values were higher, especially for fine sediments (Tables 1, 2). In the majority of the studied springs the density of oligochaetes was significantly higher in fine sediments (Fig. 5) collected by a corer than in samples taken by a scraper from coarse sediments, what was confirmed by statistical analyses (P <0.05). Only in springs with homogenous kind of the bottom (nos 18 and 23) oligochaetes were a little more abundant in samples taken by the scraper but these differences were not statistically significant. No correlation be- 240 Elżbieta Dumnicka tween organic matter content and oligochaete density was found. Both in natural northern springs, where the amount of organic matter was usually less than 1%, and in the southern ones (situated along Sąspówka stream) where the content of organic matter was distinctly higher (2.5–4.5%) the density of oligochaetes was very low (Fig. 5). The highest densities were stated in fine sediments from encased springs (nos 2, 9, 11 and 24). 5. DISCUSSION Only a few data concerning oligochaete species from springs of Kraków-Częstochowa Upland were published so far and they dealt with the southern part of the Upland. In the study area, in one spring “Pieskowa Skała” (Prądnik Stream) Tubifex tubifex and Limnodrilus hoffmeisteri were stated exclusively (Kasprza k 1976) and in three springs of Sąspówka stream 11 species were determined (D umnicka 2000), but Propappus volki, Mesenchytraeus armatus (Levinsen), Cernosvitoviella carpatica Niel. et Christ. and Nais pseudobtusa Piguet were not found during recent studies. The abundance of oligochaetes as well as the number of species found in springs from various geographic regions and various site types change in a wide range: from less than 1% of the whole benthic fauna and a few species (2–3) (Hof fs en and Ma lmquist 2000, Lindegaard et al. 1998) to 39 species found in a big karstic spring in Poland (Ka h l 2000) and 33 species determined in ten karstic springs and springbrooks in Illinois, USA (Webb et al. 1998). In both above mentioned springs oligochaetes were one of the most abundant group among benthic invertebrates. Studied springs of Kraków-Częstochowa Upland have moderately abundant oligochaete fauna and high number of determined species (totally 30 species, including these found during an earlier study) what may result from high diversity of the studied springs considering their discharge, kind of bottom sediment, position with respect to the stream bed and other environmental factors. It seems that man-made modifications of springs morphology play also an important role in the formation of oligochaete taxocens and the abundance of these invertebrates. Various kinds of spring encasing in various ways influence oligochaete fauna. Placing well-heads on the water outflows diminishes water current and facilitates sedimentation of fine sediments what forms “a trap” for oligochaete specimens drifting with stream water (in cases when the springs are situated inside or close to the stream bed) or migrating from the surrounding soil (terrestrial or semi-aquatic species). This favors an increase in oligochaete density, number of species, as well as diversity (H’). Strong modification of spring morphology by construction of “houses” from concrete might cause the disappearance of oligochaetes – what was stated in one of the springs of Sąspówka stream (no 1) and probably in Sudetes’ springs studied by Pax and Mas ch ke (1936). In springs of such a kind, if oligochaetes are present, they are represented by ubiquitous and terrestrial species only. The influence of various natural and anthropogenic factors on spring oligochaete fauna was stated by Särk kä et al. (1997) and Webb et al. (1998) but in these studies the impact of spring encasing was not taken into consideration. Composition and density of invertebrate fauna living in springs are affected by various environmental factors (L in degaard et. al. 1998, B arquin and D e at h 2004) but sometimes it is difficult to find simple relations between them due to high diversity and complexity of these factors. Opposite to Hydracarina (Biesiad ka et al. 1990) and Trichoptera (C z achorowsk i 1990) studied in springs of Kraków-Częstochowa Upland, among oligochaetes crenobionts were not found and only a few species could be treated as crenophiles: Rhyacodrilus falciformis and Stylodrilus heringianus. Other crenophilic species – Nais communis, typical for mountain springs of southern Poland (Kaspr z a k and S z c z ęsny 1976, D u mnicka 2000) were less frequently found in the studied springs. Crenophilic species were more abundant in the wellheaded springs than in natural ones. The majority of species found in the studied springs is characteristic for stream or soil oligochaete fauna and they were more frequently found in encased springs and in springs situated inside or close to the stream bed. The oligochaete fauna of the studied springs is similar to that found in small Oligochaeta in karstic springs springs of various types (helo- reo- and limnocrenes) in Pieniny Mts (Southern Poland) (Kasprza k 1979a, b) and Sudety Mts (S-W Poland) (D umnicka 2000) – especially when fauna of natural springs was taken into consideration. Enchytraeidae were the most diversified oligochaete family in those springs and the number of species found in a single spring was usually small – from 1 to maximum 9 stated in springs of Raba River (Gorce Mts, Southern Poland) (Kaspr z a k and Sz cz ęsny 1976). In lowland springs, regardless their discharge, Naididae and Tubificidae prevailed and Enchytraeidae were less numerous (Timm 1987, Ka h l 2000), but in small springs low number of species was found (D embick ij 1972, Timm 1987), whereas in big limnocrenes high number of species and high abundances were stated (Ka h l 2000). Small springs are small patchy areas, isolated from the others, where immigration of non-flying invertebrates is difficult and competition for resources is high (Gl a z ier 1991); it explains small number of species from various groups, including oligochaetes inhabiting single spring. 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