64: 3244 (2009) Japanese Journal of Benthology 1994 2005 Changes in the Macrobenthic Fauna of the Shigetomi-higata Tidal Flats in Kagoshima Bay: Comparison between 1994 and 2005 1), 1) 2) 2) 1) 8900056 45020 2) 8908580 12135 Tomoko YAMAMOTO 1),, Ai MASUYA 1),, Koji MATSUSHITA 2) and Masanori SATO 2) 1) Faculty of Fisheries, Kagoshima University. 45020 Shimoarata, Kagoshima 8900056, Japan 2) Faculty of Science, Kagoshima University. 12135 Korimoto, Kagoshima 8908580, Japan Abstract: Long-term macrobenthic faunal change was investigated in the Shigetomi-higata tidal flats of northern Kagoshima Bay by means of surveys conducted in spring and autumn of 1994 and 2005. Density and biomass of all species of macrobenthic animals and sediment characteristics were surveyed along four transect lines covering the entire intertidal area of the flats. In total, 37 and 38 species were collected from this area in 1994 and 2005 respectively. Density and biomass of the various benthic animals have changed di#erently among the transects, but gastropods remarkably increased in both density and biomass during this period along most of the transects. Especially, Batillaria multiformis dominated the flats in 2005 but were not found in 1994. In contrast, the density, biomass, and species number of bivalves decreased, and decrease of Ruditapes philippinarum in both density and size of individuals was remarkable. The dominant polychaete changed from Notomastus sp. with a larger body size to Armandia lanceolata with a smaller body size. In general, epibenthic and smaller species increased whereas endobenthic and larger species and larger individuals decreased during this period. We could not identify the environmental factors responsible for causing these faunal changes, but topographic changes in the tidal flats may have played a role in the biomass change of R. philippinarum. Key Words: Armandia, Batillaria, macrobenthos, faunal change, Notomastus, Ruditapes, tidal flats. 1993; 1994 (1994) 19891992 50 1945 6 1977 200 ha 2003 60 ha 1990 1995; 2009 20 km, 80 km, 1,130 Received 19 May 2008 Accepted 30 June 2009 Corresponding author km 2 1978 2000 1.5 km 197 2007 1993 1995 1997 1977 2003 26 60 ha 30 ha 2009 32
20042005 1617 1995 1976; 1981; 1984; 1985; 1988; 1993; 1994; 1996 1997; 1999 1974; 2005; 2007 2001 1990 1994 2005 11 11 (Fig. 1). 900 m 400 m 500 m 1997 Fig. 1. Maps showing the location of the study site (Shigetomi-higata tidal flats) in Kyushu, Japan (right), and the locations of four transect lines on the tidal flats (left). The limits of the area exposed at the lowest tide in October, 1994, are indicated by dashed lines. 1994 3 28 4 10 27 6 12 9 16 20 2005 4 7 11 10 14 16 4 14 (Fig. 1) 39 2005 1 4 1 4 3 1994 1 4 15 cm15 cm, 2 3 30 cm30 cm 40 cm40 cm 2005 25 cm25 cm 1994 1.19 m 2 0.61 m 2 2005 1.69 m 2 0.56 m 2 30 cm 1mm 10 70 80 (nmds) (Bray-Curtis Index) 33
Bray-Curtis Index i x Ai x Bi (1) X A X B x Ai, x Bi A, B i X A, X B SPSS MDS 1994 6 22 10 15 16 2005 4 12 2 cm 1994 9 2005 4 10 1cm 29 mm 1994 2005 2, 3 120 12 63 mm 63 mm 1994 2005 MRK-RETAC 10 2008 10 4 7 29 mm 1cm 3 120 12 2 1994 12 14 2005 4 10 1994 13 1 1 50 m 3 2005 1 120 12 650 2 1994 9 2005 4 29 mm 1cm (IS/Mill-E iuchi) 11 Wilcoxon Kendall 1. 1994 37 14 10 9 1 3 2005 38 19 10 4 3 2 (Table 1). 4 1994 27 12 7 6 1 1 2005 35 18 9 3 3 2 1 4 1994 15 6 3 6 2005 16 6 2 4 3 1 1994 2005 (Musculista senhousia), (Nuttallia japonica), (Psammotaea minor), (Solen strictus), (Phacosoma japonicum) 1994 2005 1 (Moerella rutila), (Clithon oualaniensis), (Batillaria multiformis), (B. multiformis) (B. cumingii) ( 2003) nmds 1994 1, 4 2005 2, 3 34
Fig. 2. Non-metric multidimensional scaling analysis of the benthic animal community in each quadrat. The distances between plots represents their similarity. Each symbol indicates quadrats along the same transect line in the same season and year. Closed symbols represent the data from spring of 1994, and open ones the spring of 2005; grey symbols show the data from autumn in both years. Positions (transect line number and distance from land) are shown beside the plots of quadrats isolated from others in the same transect line during the same season and year. MDS (Fig. 2). 1 4 1994 180 m 270 m 2005 180 m 2005 180 m 4 2 2005 180 m 90 m 0.239 0.1 2. 1 (Ruditapes philippinarum) (Fig. 3a, Table 2). 1m 2 100 g (Fig. 3b). 1994 1m 2 1000 g 2005 10 g/m 2 170 g/m 2 1994 2005 Fig. 3. Changes in macrobenthic community composition on the Shigetomi-higata tidal flats between 1994 and 2005. Averages of (a) density and (b) biomass (wet weight) for all four transect lines were calculated. 35
Table 1. List of benthic species sampled along each transect line during Phylum Class Japanese Name Species Name Cnidaria Anthozoa 1 Actiniaria gen. sp. Platyhelminthes Turbellaria 1 Platyhelminthes gen. sp. Nemertea 1 Nemertinea gen. sp. Mollusca Gastropoda Clithon oualaniensis Batillaria multiformis Reticunassa festiba Bivalvia Musculista senhousia Mactra veneriformis Moerella rutila Macoma tokyoensis Nuttallia japonica Psammotaea minor Solen strictus Phacosoma japonicum Ruditapes philippinarum Laternula marilina Annelida Polychaeta Glycera nicobarica Hemipodia yenourensis 1 Goniada sp. Sigambra hanaokai 1 Syllidae gen. sp. 1 Hediste sp. Ceratonereis erythraeensis 1 Leonnnates sp. 1 Nephtyidae gen. sp. 1 Chrysopetalidae gen. sp. Marphysa sanguinea 1 Marphysa sp. 1 Lumbrineridae gen. sp. Pseudopolydora cf. kempi 1 Scolelepis sp. 1 Prionospio sp. Mesochaetopterus japonicus Cirriformia tentaculata Armandia lanceolata 1 Notomastus sp. 1 Capitellidae gen. sp. 1 Polychaeta gen. sp. Arthropoda Crustacea sp. 1 Amphipoda gen. sp. 1 sp. 2 Amphipoda gen. sp. 2 Neomysis japonica 1 Cumacea gen. sp. 1 Caprellidae gen. sp. Hippolytidae gen. sp. Crangon a$nis complex Callianassa japonica Upogebia yokoyai Philyra pisum Scopimera globosa Hemigrapsus penicillatus 1 Grapsidae gen. sp. Polyonyx sinensis 1 Paguridae gen. sp. Diogenes nitidimanus Echinodermata Holothuroidea Patinapta ooplax No. of species Total area of quadrats (m 2 ) ) Identified by Ken-ichi Hayashi. ) Identified by Katsuji Sakai. N) Only nest tubes were sampled. 36
surveys in the Shigetomi-higata tidal flats of Kagoshima Bay in 1994 and 2005. 1994 spring 2005 spring 1994 autumn 2005 autumn line 1 line 2 line 3 line 4 line 1 line 2 line 3 line 4 line 1 line 2 line 3 line 4 line 1 line 4 N 12 11 10 15 19 14 25 12 6 9 14 11 12 8 27 35 23 (lines 1, 4: 15) 16 0.28 0.44 0.27 0.20 0.50 0.31 0.44 0.44 0.16 0.09 0.16 0.20 0.38 0.19 1.19 1.69 0.61 (lines 1, 4: 0.36) 0.56 37
Table 2. Mean density (no. of individuals per 1 m 2 ) of macrobenthic species along each transect line in the Shigetomi-higata tidal flats no data indicates that the sample was too small or in poor condition. line 1 spring Phylum Class Japanese Name Species Name 1994 2005 Cnidaria Anthozoa 1 Actiniaria gen. sp. 4.0 (0.15) Platyhelminthes Turbellaria 1 Platyhelminthes gen. sp. Nemertea 1 Nemertinea gen. sp. 4.0 no data Mollusca Gastropoda Clithon oualaniensis Batillaria multiformis 96.0 (0.85) Reticunassa festiba 6.0 Bivalvia Musculista senhousia Mactra veneriformis Moerella rutila 22.0 (0.17) Macoma tokyoensis Nuttallia japonica Psammotaea minor 5.7 Solen strictus Phacosoma japonicum Ruditapes philippinarum 129.3 16.0 (0.49) (0.42) Laternula marilina 3.7 Annelida Polychaeta Glycera nicobarica 11.7 Hemipodia yenourensis 22.0 1 Goniada sp. 2.0 Sigambra hanaokai 1 Syllidae gen. sp. 2.0 1 Hediste sp. Ceratonereis erythraeensis 26.0 2.0 1 Leonnnates sp. 1 Nephtyidae gen. sp. 1 Chrysopetalidae gen. sp. Marphysa sanguinea 1 Marphysa sp. 2.0 1 Lumbrineridae gen. sp. 11.0 10.0 Pseudopolydora cf. kempi 7.3 1 Scolelepis sp. 1 Prionospio sp. Mesochaetopterus japonicus 2.0 Cirriformia tentaculata Armandia lanceolata 11.7 10.0 (0.01) (0.01) 1 Notomastus sp. 7.3 2.0 (0.35) no data 1 Capitellidae gen. sp. 2.0 1 Polychaeta gen. sp. Arthropoda Crustacea 1 (sp. 1) Amphipoda gen. sp. 1 4.3 (0.002) 1 (sp. 2) Amphipoda gen. sp. 2 Neomysis japonica 1 Cumacea gen. sp. 16.0 1 Caprellidae gen. sp. 1 Hippolytidae gen. sp. Crangon a$nis complex Callianassa japonica 11.7 8.0 (0.06) (0.96) Upogebia yokoyai Philyra pisum Scopimera globosa Hemigrapsus penicillatus 3.7 1 Grapsidae gen. sp. Polyonyx sinensis 1 Paguridae gen. sp. 2.0 Diogenes nitidimanus 38
of Kagoshima Bay ( indicates absence). Wet weight (g) per individual is shown in parentheses for the dominant species of each class; line 2 spring line 3 spring line 4 spring line 1 autumn line 4 autumn 1994 2005 1994 2005 1994 2005 1994 2005 1994 2005 2.3 no data 4.9 (0.01) 2.3 5.3 no data no data 6.4 10.7 153.6 216.0 26.7 (0.42) (0.76) (0.35) 2.0 16.0 9.1 13.3 2.7 7.3 9.9 18.7 2.3 4.9 5.3 22.4 34.3 11.4 96.0 16.0 (0.15) (0.06) (0.05) (0.06) (0.03) 6.3 4.9 39.6 12.7 4.9 4.3 3.7 2.0 0.4 33.3 11.4 404.9 11.4 285.6 13.3 19.7 10.7 (3.23) (5.06) (0.53) (0.39) (0.24) (0.29) (2.69) (0.40) (1.42) (0.32) 24.7 5.3 3.7 9.1 4.9 13.7 24.7 9.1 2.0 3.2 3.7 12.6 6.9 4.6 2.7 4.9 5.3 148.0 2.0 9.8 2.7 24.7 4.9 4.9 6.9 2.3 3.7 4.0 4.6 2.3 2.7 10.3 28.8 18.3 32.0 69.1 22.9 56.9 10.7 34.4 4.9 2.7 3.2 4.6 2.3 2.3 4.6 307.2 326.9 2.3 (0.004) (0.01) (0.001) 50.7 36.7 202.4 49.3 5.3 (0.12) (0.23) (0.03) (0.02) (0.003) 2.3 18.3 4.6 (0.001) (0.006) 43.4 (0.001) 8.3 9.6 6.9 2.3 12.8 2.3 7.3 4.3 6.4 11.4 2.3 (1.68) (0.02) (0.09) (0.24) 9.8 2.3 4.9 5.3 4.9 6.3 4.9 3.2 2.3 16.0 6.4 39
1 0.4 g 2.7 g 0.06 g 0.17 g (Table 2). 1 2.7 g 0.4 g (Table 2). 2 3 2: 100 /m 2 600 /m 2 3: 130 /m 2 450 /m 2 (Fig. 3a). 2 1m 2 100 g (Fig. 3b, Table 2), 3 50 g/m 2 20 g/m 2 1994 (Fig. 3b). 1 0.5 g 0.4 g (Table 2). 2, 3 Notomastus sp. (Armandia lanceolata) 1 0.1 g 0.004 g (Table 2). 4 (Fig. 3a). 1000 /m 2 70 /m 2 10 1 170 /m 2 80 /m 2 180 g/m 2 10 g/m 2 40 g/m 2 10 g/m 2 (Fig. 3b). (Table 2). 1994 Hediste diadroma H. atoka 2005 Notomastus sp. (Fig. 4) 1 1994 180 m 493 /m 2 ; 1466 /m 2 (Fig. 4a, b), 2005 3090 m (Fig. 4a, b), 656 /m 2 ; 912 /m 2 60 m 30 m 2, 3 1994 2005 2: 30 m 75 m 3: 10 m 210 m (Fig. 4c, d). 2005 2 (Fig. 4c), 3 (Fig. 4d). 4 1994 240270 m 2005 90 m 180 m 4 30 m, 60 m, 150 m, 210 m (Fig. 4e). 1994 270 m 2005 (Fig. 4f). 3. Fig. 5 Table 3 1994 2005 Fig. 5ad 4 4 1994 250 m 300 m 2005 200 m 3 50 cm Fig. 5ad 2005 4 1: 0 12 2: 211 3: 14 4: 1 5 1994 9 1: 171 2: 11 19 3: 310 4: 461 Wilcoxon n20, p0.01 1 4 120150 m 1 4 2005 9 1: 0.2 12 4 016 1994 9 Wilcoxon n10, p0.05 2005 4 9 Wilcoxon n10, p0.05 1994 2005 1994 2005 Wilcoxon n7, p0.05 (SD) 2.21 ( 40
Fig. 4. Changes in vertical distribution of macrobenthic animals on the Shigetomi-higata tidal flats between 1994 and 2005. The number of individuals per 1 m 2 was calculated for each quadrat. Figures of zero ( 0 ) in (e) indicate quadrats from which no macrobenthic animals were recovered. Symbol explanations in (f) pertain to all graphs. 0.61) (SD) 0.43 (0.43) 2005 4 9 14 Fig. 5ad 1994 4 1 50 m 24 1994 Kendall n33, p0.05 (Table 3), 1994 9 2005 4 1994 9 1: 30 32 2: 31 3: 2832 4: 30 33 2005 4 1: 2225 2: 17 32 3: 932 4: 2331 2 75175 m 3 90 m 4 30 m 1994 2005 11 37 1.80 m 2 38 2.25 m 2 1 2003 MDS 1994 2005 (Fig. 2). 1 4 11 1994 2005 41
Fig. 5. Environmental characteristics of each transect line (a-d: lines 14, respectively). Upper graphs show the profiles of the tidal flats along the transects. Middle graphs indicate the mud contents at each sampling point along the transects. Lower graphs show the proportion of ignition loss at each sampling point along the transects. MHWS: mean high water level in spring tides; MTL: mean tidal level; MLWS: mean low water level in spring tides. Symbol explanations in (d) pertain to all graphs. Sato (1994) 1991 1992 76 11 2005 (Table 2). 2005 1994 1 1994 Notomastus sp. 2005 1994 1 1 2005 1994 2005 11 Notomastus sp. (1996) 2005 11 1994 2005 42
Table 3. Salinity of the interstitial water in each quadrat along the four transect lines in the Shigetomi-higata tidal flats of Kagoshima Bay. Water samples were not taken in some quadrats, indicated by. Transect Distance from land September 1994 () April 2005 () Line 1 30 m 31.5 23.5 60 m 31.0 23.5 90 m 30.0 21.5 120 m 31.0 23.0 150 m 31.5 25.0 180 m 31.0 25.2 210 m 30.0 MeanSD 30.90.6 23.61.4 Line 2 30 m 31.0 31.5 75 m 31.0 17.2 135 m 18.5 175 m 18.2 MeanSD 31.0 21.46.8 Line 3 30 m 32.0 31.6 60 m 28.0 31.0 90 m 30.0 8.5 120 m 31.0 26.8 150 m 29.0 25.9 180 m 31.9 210 m 18.8 MeanSD 30.01.6 24.98.6 Line 4 30 m 30.0 22.9 60 m 31.0 29.7 90 m 32.0 27.7 120 m 32.0 31.3 150 m 33.0 31.2 180 m 33.0 31.2 210 m 33.0 240 m 32.0 270 m 32.0 MeanSD 32.01.0 29.03.3 2002 5 2007 4 1994 2005 240 m 1994 4 200 m 1994 4 2000; 2005 4 1994 2005 2007 2005 2002 1990 2005 Notomastus sp. 2006 1994 2005 2005 1cm 1994 1 4 60 2005 1994 2005 1994 2005 11 43
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