African Journal of Marine Science 2010, 32(1): 85–93 Printed in South Africa — All rights reserved

Copyright © NISC (Pty) Ltd

ISSN 1814–232X EISSN 1814–2338 doi: 10.2989/18142321003714575

Distribution patterns of striped mullet Mugil cephalus in mangrove creeks, Zanzibar, Tanzania
AW Mwandya1, YD Mgaya2, MC Öhman3, I Bryceson4 and M Gullström3,5* Department of Animal Sciences and Production, Sokoine University of Agriculture, PO Box 3004, Morogoro, Tanzania Department of Aquatic Sciences and Technology, University of Dar es Salaam, PO Box 60091, Dar es Salaam, Tanzania 3 Department of Zoology, Stockholm University, SE-106 91 Stockholm, Sweden 4 Norwegian University of Life Sciences, PO Box 5003, NO-1432 Aas, Norway 5 Department of Systems Ecology, Stockholm University, SE-106 91 Stockholm, Sweden * Corresponding author, e-mail:
1 2

Manuscript received August 2009; accepted January 2010

Spatial and seasonal variations in density of striped mullet Mugil cephalus were investigated in four mangrove creeks in Zanzibar, Tanzania, during a one-year cycle. Fish were collected monthly in the lower, intermediate and upper reaches of each creek using a beach-seine net. All fish collected were juveniles between 2 and 16 cm standard length. The density of juvenile mullet differed significantly among the creeks, but the spatial patterns within them were consistent with higher densities upstream in three of the creeks. Generally, small mullet (2–10 cm) were more abundant in the upper reaches compared to the lower and intermediate reaches. Seasonal patterns were weak, although mullet densities were high during the period of heavy rains (March–May). Principal component analysis showed that a muddy substrate with microphytobenthos was positively correlated with high mullet densities, although site-specific variables such as shallow water depth and water clarity were also significantly correlated. Our findings suggest that the densities of juvenile striped mullet vary among sites and creeks in response to refuge availability from turbid, shallow water and the accessibility of food from benthic microalgae. Keywords: East Africa, marine embayment, Mugil cephalus, population density, spatial and seasonal variation

Introduction Mangroves are variable in terms of structural features and environmental conditions, which may uniquely influence spatial and seasonal patterns of fish populations (Nagelkerken et al. 2008). Different habitats are recognised within mangrove areas, including tidal creeks in which fish proliferate in high abundances (Tongnunui et al. 2002, Singkran and Sudara 2005). Juvenile fish are commonly found in tidal creeks because many adults use them as nursery areas, before migrating to other habitats such as seagrass beds or coral reefs (Robertson and Duke 1990, Cocheret de la Morinière et al. 2004, Dorenbosch et al. 2006). The striped mullet Mugil cephalus is a circumtropical species found in nearshore coastal waters. It predominately occupies the mouths of rivers and streams or estuaries and freshwater environments, but spawning takes place at sea. The species has a bipartite life cycle because it undergoes an ontogenetic shift in terms of both habitat and food preferences (De Silva 1980). In the tropics, mangrove creeks seem to play an important role as nursery area for juvenile mullet (Barletta et al. 2003, Koutrakis 2004). This has implications for the management of mangrove resources because of the commercial importance of striped mullet (Mahmoudi 2000, Jiddawi and Öhman 2002). Being tolerant to environmental changes, M. cephalus has good potential for aquaculture (Oren 1981, Pillay and Kutty 2005). For this reason, it is a priority species for future development of marine aquaculture in areas where such activities are in their infancy, such as in East Africa (Mmochi and Mwandya 2003). The success of such development is largely dependent on the availability of juveniles, for which mangrove creeks may play an important role. However, information is lacking on the distribution and habitat preferences of M. cephalus in mangrove creeks, particularly in the Western Indian Ocean region (but see Kimani et al. 1996, Lugendo et al. 2005, 2007, Mwandya et al. 2009). The aim of this study was to investigate spatial and seasonal variation in the density and size distribution of M. cephalus within and among mangrove creeks, and to explore how patterns may relate to habitat characteristics and environmental conditions. Such information has implications in defining productive habitats for this economically important species.

African Journal of Marine Science is co-published by NISC (Pty) Ltd and Taylor & Francis

as well as water clarity. and Hypnea sp. Patches of Halimeda spp. were the dominant macroalgal species in Kiongwe Creek. half of the sediment was placed in a crucible and burnt in a furnace at 450 °C to constant weight (for a period of ~24 h). forming small patches at the intermediate and upper sites. the intermediate and lower reaches) were deeper. the downstream regions of the creeks (i.5 cm). were measured twice every sampling trip. Fish were divided into three size classes based on maximum lengths of M. through which several tidal creeks and channels intertwine. it was not possible to sample over the whole tidal cycle at each site. Rainfall data were obtained from the Tanzania Meteorological Agency. subadults one. the second haul was taken 10 minutes after the first. 39°24. with a mean depth at low tide of 1. especially in Kiongwe and Kinani creeks where there were thick layers of microphytobenthos on the muddy bottom.5–3 m (Table 1). the heavy rains from March to May and the moderate rains from October to December. macroalgae. Data analyses The variations in mullet densities and environmental variables were analysed using one-way analysis of variance and the t-test. cephalus (Froese and Pauly 2007). The Games-Howell post hoc test (Field 2000) was used to . in Makoba Bay it is 4 m and 2 m respectively (AWM pers. Thalassia hemprichii and Cymodocea two-thirds of SL. To assess sediment organic matter content and grain size.5 m). Hypnea spp. Both bays are fringed by dense mangrove forests. Sampling procedures In order to assess the variation in mullet densities within each of the four creeks. and to some extent rivers. Levene’s (1960) test was applied to check if the assumptions of homogeneity of variances were met. Tanzania (Figure 1). three replicate sediment samples were taken from each site to a depth of 5 cm. Microphytobenthos species were identified using a light microscope and the relative percentage frequency of individual taxa was determined. However. obs.13′ S. Salinity.0 m. some sites were characterised by macrophyte bottom vegetation (i. The area usually experiences two rainy seasons.. whereas water depth was measured only once. intertidal waterbodies. and with relatively high water clarity. characterised the lower part of Kinani Creek. Fish samples were collected at low tide each month from January to December 2005 (except September).8–1. seagrass and macroalgae). The catches of each haul were placed in separately labelled plastic bags and transported in an icebox to the laboratory. the second dry season was extended and occurred from June to October. 39°11.). non-parametric tests (Kruskal-Wallis or Mann-Whitney U) were applied. Padina sp. The standard length (SL) of each individual was measured. intermediate and upper reaches (Figure 1). biases due to tidal variation were alleviated to some degree. Water temperature and dissolved oxygen were measured using an oxygen meter.86 Mwandya. The samples were homogenised and oven-dried at 65 °C to constant weight. The upstream parts of the four creeks were shallow (mean depth at low tide = 0.17′ E) and Chwaka Bay (6°6. lower. 1995). Because of logistical constraints. Seagrass areas in both bays were composed mainly of Enhalus acoroides.58′ S.e. Mgaya. Environmental variables were recorded during daytime and at low tide. When the assumptions of homogeneity of variance were not met. Life stage categories recorded for each fish followed Nagelkerken and van der Velde (2002). water temperature and dissolved oxygen (in the upper 0. Makoba Bay and Chwaka Bay are both shallow. The four areas selected for this study were Kiwani and Kiongwe creeks in Makoba Bay and Mapopwe and Kinani creeks in Chwaka Bay (Figure 1). The fish were further divided into size class intervals of 2 cm and grouped for each site (i. microphytobenthos. Kiwani and Mapopwe are the largest creeks of the respective bays and comprise the main water exchange routes through the forests. Bryceson and Gullström Material and methods Study sites The study was conducted in the mangrove creek systems of Makoba Bay (5°43. Two replicate hauls were taken at every site during each sampling trip. The average depth in Chwaka Bay is 3.9 cm. Kinani Creek is relatively small. each influenced by an asymmetric semi-diurnal tidal cycle. Fish were caught using a 17 m long and 2 m wide beach-seine net with a stretched mesh size of 1. Benthic microalgae were sampled by scooping triplicate subsamples of sediment (about 20 g each) from the upper layer of the bottom sediment (to a depth of 0.31′ E). may temporarily alter the salinity gradient of these embayment systems (Johnstone and Mohammed 1995). Prior to each analysis. and adults more than two-thirds SLmax. located at Zanzibar Island (Unguja).e. Prior to the establishment of permanent sampling sites.e. Halimeda spp. as well as two dry seasons. To minimise the effect of consecutive sampling. Salinity was estimated using a hand refractometer and water clarity was taken by means of a Secchi disk. Their abundance decreased farther upstream.5 m at low spring tide (Cederlöf et al. Table 1). from January to February and July to September (McClanahan 1988). three sampling sites were established — at the lower. and Sargassum spp. In contrast. To estimate organic matter content. Öhman. during this study. channel-like tributary to Kiwani Creek. rains. and Kiongwe Creek is even smaller but is an important. which covered an area of about 170 m2 in each haul. The tidal dynamics. Table 1) with relatively low water clarity and a high percentage of microalgal coverage.2 m at high spring tide and 2. Gracilaria salicornia. dominated the macroalgae assemblages at the lower site in Mapopwe. The other half of the sediment was used for grain size analysis according to Folk and Ward (1957). determination of coverage of the major bottom substrate types (seagrass. even after transformations (square root or log10(x+1)). Juveniles represented fish of less than one-third of the maximum SL (SLmax). intermediate and upper reaches) to assess spatial distribution patterns in size frequency. by always sampling at low tide. However. sand and/ or mud) was conducted by snorkelling with a 2 m2 square frame that was randomly placed within each site (n = 75. and Dictyota spp. A posteriori multiple mean comparisons were performed using Tukey’s test for data that showed homogeneity of variances.

the sediment organic matter content increased from the lower to upper reaches in all creeks .African Journal of Marine Science 2010. p < 0.0 0.5 <2 60 30 <1 U 1. with low values (15 ± 7) occurring during the rainy season and high (27 ± 6) during the dry season (Mann-Whitney U-test. Grain size and organic matter contents were not included in PCA analysis on account of the small sample sizes. Significant seasonal variations were only recorded for salinity in the upper reaches of Kiongwe.0 28 12 57 <5 U 0.05). with water temperature and dissolved oxygen being relatively stable across all sites. Kiongwe Creek (4–6). as well as the lower site of Mapopwe.7 ± 0. Thereafter.0 59 27 12 <2 L 3. water temperature. Water clarity at the intermediate site of Kiwani was higher during the dry season (0.5 <5 80 5 10 Variable Water depth (m) Macrophyte cover (%) Mud cover (%) Sand cover (%) Microphytobenthos cover (%) L 2. the intermediate and upper sites at Mapopwe.4 ± 0.5 km 0 0 5 10 km 1 2 km Mangrove forest Mangrove forest Figure 1: Map of Zanzibar Island. water clarity and dissolved oxygen were found during the period of heavy rains. varied from very fine sand to silt (Figure 2). 32(1): 85–93 87 (a) (b) Kiongwe Creek AFRICA Tanzania ●4 ●1 Kiwane Creek Zanzibar TANZANIA 5● Chwaka Bay 39°30′ E ●6 ●3 ● 2 Makoba Bay ●7 8 ● Mapopwe Creek (a) 6° S ZANZIBAR ●10 Kinani Creek INDIAN OCEAN ● 9 ● 11 12 ● (b) 0 0.0 <2 60 35 <2 L 3. as well as the sampling sites in each creek within the bays.0 25 25 5 <2 68 70 <2 5 Kinani I 1. Results The environment Low values of salinity.0 65 26 5 <5 L 3. Tanzania. Salinity was lower at the upper site compared to the lower and intermediate sites of Kiongwe. being highest at the lower sites and lowest at the upper sites (Table 2). and the lower sites of Kiongwe and Kinani were characterised by mediumsized grain sand (Figure 2).8 4 40 <2 55 identify differences among sites. intermediate and upper sites are shown for Kiwani Creek (1–3). and at the upper site relative to the lower site of Kinani (Table 2). The sediment grain size at the intermediate and upper sites at Kiongwe and Kinani. showing (a) Makoba Bay and (b) Chwaka Bay. the significance levels were adjusted using the Bonferroni method. Water clarity showed consistent and significant gradients within all creeks. In general.1 m) (t-test. Principal component analysis (PCA) was used to minimise the dimensionality of the abiotic data (Manly 1994). Significant spatial patterns were found only in salinity and water clarity.2 m) than the rainy season (0.8 25 – 25 40 45 5 <2 50 Mapopwe I U 1. For all multiple comparison tests. There was no significant difference in water clarity between the intermediate and upper sites of Kiwani (Table 2). Mapopwe Creek (7–9) and Kinani Creek (10–12) Table 1: Habitat characteristics of the the lower (L).5 1.5 <2 55 40 <2 Kiongwe I U 2. especially in April and May.05). Sediment characteristics The bottom sediment at all three sites of Kiwani. p < 0. Data are mean values Kiwani I 1. The lower. intermediate (I) and upper (U) sites within each mangrove creek. This approach overcomes problems related to the colinearity of abiotic factors in which the principal component scores are used as explanatory variables. multiple regression was applied to investigate the relationship between (ln+1)-transformed mullet densities and the PCA.

05 ** = p < 0.5 28.15 0.8 27. as well as at the intermediate site of Kinani (Mann-Whitney U-test. Microcoleus spp. no noticeable cover of these microorganisms was recorded at the lower and intermediate sites in any creek (Table 1). Nodularia sp.6 2.5 6.1 0.0 0.5 0.3 28.8 5. Kinani: p < 0.6 28.8 2.0 0.7 32.2 5.1 0. more taxa were found in the upper sites of Kiongwe and Kinani — although in different proportions — compared to the upper sites of Mapopwe and Kiwani.9 29.1 0.3 28.1 0.2 0.3 33.2 7.1 0.2 32.8 Mean 28.3 2. .9 1.4 21.4 2.4 0. p < 0.9 28.6 0.9 1.3 0.7 5.9 30.4 33. Others ORGANIC MATTER (%) 8 6 4 2 0. Mgaya. The highest values of organic matter were at the intermediate and upper sites of Mapopwe.88 Mwandya.5 0.9 Post hoc comparisons L vs I L vs U I vs U ns ns ns ns * ** ns ns ns ns * ns ns ns ns ns ** * ** ** ns ns ns ns ns ns ns ns ** ** ** ** ns ns ns ns ns ns ns ns ns ** ** * ns ns ns ns Variable Salinity Creek Kiwani Kiongwe Mapopwe Kinani Kiwani Kiongwe Mapopwe Kinani Kiwani Kiongwe Mapopwe Kinani Kiwani Kiongwe Mapopwe Kinani Water temperature (°C) Water clarity (m) Dissolved oxygen (mg l–1) * = p < 0.3 I SD 8.3 0.5 5.7 27.1 0.8 2. Kiongwe and Kinani. Spirulina spp.1 0.7 3.5 24. p < 0.001 ns = not significant (p > 0.2 5.6 5. There was spatial variation in mullet densities within all four creeks.6 SD 9.5 0.20 0.2 0.5 5.1 8.6 1.9 0. However.4 5. distribution patterns and size Variation within creeks Seasonal patterns in mullet density were generally obscured by the high monthly variability within each season (Figure 4).2 5.4 2.01. Results of post hoc tests (Tukey’s and Games-Howell tests) are shown L Mean 35. intermediate (I) and upper (U) sites Mapopwe SITE Kinani Figure 3: Relative frequency of occurrence of the most dominant taxa of microphytobenthos taken from the upper site in each creek (Figure 2).2 5. intermediate (I) and upper (U) sites of each creek.5 32. p < 0. df = 18.4 2.6 29. (Figure 3).01) respectively.3 6.7 2.3 5.7 1. Bryceson and Gullström Table 2: Spatial variations in mean (±SD) of environmental variables (n = 11) at the lower (L).6 0.5 5.0 28. there were significantly higher densities during the rainy season at the intermediate (t-test.10 0.8 0.05 L I U Kiwani L I U L I U L I U Kiongwe Mapopwe Kinani SITE GRAIN SIZE (mm) 40 30 20 10 Kiwani Kiongwe Figure 2: Sediment organic matter contents and sediment grain size measured in each site for lower (L). For microphytobenthos. In contrast.2 0.6 5.5 U SD 12.1 0.05).05) and intermediate sites (p < 0.7 29.9 27.5 7. Kiongwe: p < 0. Spirulina spp.6 0.05) Organic matter Grain size Oscillatoria spp.7 2.3 0.7 2.5 0.9 Mean 26.0 0. df = 18.2 0. whereas Mapopwe had highest densities at the lower site (Figure 5).05) and upper sites (t-test. Densities were significantly higher in the upper sites in both Kiongwe and Kinani than the lower (Games-Howell tests. whereas the least was at the lowest site of Kiwani (Figure 2). RELATIVE FREQUENCY 0.1 0.1 29.001) at Kiongwe. Mullet densities.25 Lyngbya sp.3 0.7 11. the highest mean densities were at the upper sites. and Microcoleus spp.3 5.1 0. In Kiwani. Öhman.4 4..7 2.2 0. The dominant taxa in the upper sites were Oscillatoria spp.

intermediate and upper sites. Variables negatively correlated with 0 500 Kinani 400 300 200 100 0 Jan Feb Mar Apr May Jun Jul Aug Oct Nov Dec Dry Rainy Dry MONTH AND SEASON Rainy Figure 4: Seasonal (dry and rainy seasons) variations in mean density of M.5 ± 2. In Makoba Bay.001). Multiple regression showed a significant relationship between mullet densities and the explanatory variables. water depth and macrophytic coverage than the intermediate and upper sites (Figure 7). and positively correlated with microphytobenthos and mud cover.05).3) (p < 0. According to the partial regression coefficients and the correlations among the principal components and the original abiotic variables. Error bars denote +1SE. mainly at the upstream sites (Figure 6). water depth. whereas in Chwaka Bay.2) compared to Kiwani (6. Generally. Note the different scales on the y-axis Variation between creeks Within each bay. Axis 3 was negatively correlated with salinity.01) and in Kinani compared to Kiwani (p < 0.4%) of the total variance (Table 3). mullet densities were positively correlated with microphytobenthos and mud cover (which were positively correlated with PC1). The first principal component separated the lower.8 ± 31. the mean size of mullet increased from upstream to downstream in most creeks. only the partial regression and coefficients for the first and second PC axes were significantly different from zero (Table 4).3) than in Mapopwe (1.African Journal of Marine Science 2010. water clarity and macrophyte coverage. intermediate and upper sites of the four creeks. whereas larger fish (8–16 cm) formed a greater proportion of the fish at the lower site of Mapopwe.05). cephalus in each of the four creeks. with the lower sites having higher values for salinity. and positively correlated with sand cover. The upper sites were characterised by microphytobenthos and mud cover. Axis 2 was negatively correlated with dissolved oxygen. water clarity. A relatively large number of intermediate-sized mullet (6–8 cm) were found in the upper site of Kinani. The first axis was negatively correlated with salinity. Kiongwe had a higher mean density (30.6 ± 71. temperature and sand cover. 32(1): 85–93 89 Kiwani 40 MEAN DENSITY (100 m−2) Lower Intermediate Upper 100 Kiwani Kiongwe Mapopwe Kinani 20 50 0 120 100 80 MEAN MULLET DENSITY (100 m−2) 60 40 20 0 10 8 6 4 2 Kiongwe Lower Intermediate SITE Upper Figure 5: Spatial variations in mean density of M. temperature and mud cover. Size distribution Small mullet (4–6 cm SL) dominated the catches in Kiwani. mullet density differed significantly between creeks. p < 0. water clarity. whereas the intermediate site was influenced by high cover of mud and sand (Figure 7). Error bars denote +1SE Mapopwe However. the mean density was higher in Kinani (43. and positively correlated with macrophyte cover (Table 3). There were higher densities of mullet in Kiongwe compared to Mapopwe (p < 0.5 ± 9. However. . cephalus in the lower. Relationships between mullet density and environmental variables The first three axes of a PCA of abiotic factors contributed to a significant proportion (67. Kiongwe and Kinani. mullet densities did not differ between Kiwani and Mapopwe or between Kiongwe and Kinani.2) (Mann-Whitney U-test.

3 88. Although seasonality in abundance was not clear.51 0. cm) Table 3: Eigenvalues and percentage of variation of the first five principal components and loading of abiotic variables on the first three principal components (PC1.35 −0. 0 14 . 0 6. 0– 4. Note the different scales on the y-axes 4. 1– 10 .4 PC3 −0.4 78. water clarity. 0 14 .1 –1 6. The high densities of mullet in the muddy areas of the upper reaches of Kiongwe and Kinani might be attributable to the presence of thick layers of microphytobenthos on the sediment. they were most abundant in the upper reaches in the creeks of both Chwaka Bay and Makoba Bay. Predation on juveniles seems to decline with decreasing water clarity (Stunz and Minello 2001) and water depth (Ruiz et al. 0 2. 1993).27 −0.55 −0.1 –1 6.0 10 . 1– 8.10 1. which could be an important food source.33 −0.0 10. 0 12 . 1– 6.07 −0. 1– 6. 0– 4.1 –1 2.0 PC2 −0.22 0.12 Cumulative 34. 1– 8.01 −0.48 −0.01 −0.98 0. Bryceson and Gullström Kiwani 200 150 100 50 Mapopwe 1 200 1 000 800 600 400 200 Kiongwe Lower Intermediate Upper NUMBER OF INDIVIDUALS 30 25 20 15 10 5 2 500 2 000 1 500 1 000 500 Kinani Figure 6: Length frequency distribution of M. Baker and Sheaves (2006) argued that the presence of large piscivores in shallow tropical estuaries highlights their importance as predators of young fish in such an environment.26 0.42 PC1 (salinity. PC2 and PC3) Percentage variation Individual 34.37 −0. the presence of organic matter and fine sediment particles in the upstream areas may provide Component 1 2 3 4 5 Variable Salinity Dissolved oxygen Water clarity Water temperature Water depth Macrophytes Mud Sand Microphytobenthos Eigenvalue 3.26 0.5 18.20 0.0 10 . Öhman.43 0.4 67.5 53.17 −0. cephalus among sites in each of the study creeks. 0 2. Major (1978) reported that juvenile mullet prefer shallow waters due to enhanced food availability and increased predator avoidance capabilities provided in such habitats.90 Mwandya. on account of the high monthly variation.04 −0. Access to food may influence the spatial distribution of juvenile mullet. 0 6. mostly ranging between 2 and 10 cm. The upper reaches of the creeks under study provide suitable conditions for juvenile recruitment and survival.70 1.43 −0. water depth and macrophytes) were also negatively correlated with mullet densities.9 10. Sand cover was positively correlated with PC2 but negatively correlated with mullet densities.31 −0. 0 . 8. Mgaya.24 Discussion All mullet collected at the study sites were juveniles. 0 SIZE CLASS (SL.48 0.90 PC1 −0.1 –1 2. some of the creeks seemed to have higher abundance during the rainy seasons (especially during the heavy rains in March–May). 0 12 .1 –1 4.45 −0. However. 1– 10 .1 –1 4.41 −0. 0 8. Generally.02 −0. Furthermore. 0 4.50 0.09 −0.9 14.

241 0. the finer sediment grain size . 32(1): 85–93 91 4 (a) 3 2 1 PC2 0 −1 −2 −3 −4 4 3 2 1 PC3 0 −1 −2 −3 Sl Wc Wd Mp Microphytobenthos. PC2 and PC3): p-values < 0. the higher densities of mullet in the intermediate and upper Table 4: Results of multiple regression analysis on densities of M.044 SE of coefficient 0.668 −0. detritus and sediment.African Journal of Marine Science 2010. Wt = water temperature.485 PC1 PC2 PC3 sites of Kiongwe relative to Kiwani may be on account of the large cover of microalgae. Likewise. which had less microalgae and coarser sediment grain size. Sanchez-Rueda 2002) as they are capable of feeding on the bottom by sucking the upper layer that is comprised of benthic microalgae (diatoms and dinoflagellates). detritus material and diatoms are commonly found in the stomachs of juvenile mullet (Collins 1981. In Chwaka Bay. mud Lower Intermediate Upper Sand Mp −4 −4 −3 −2 −1 0 PC1 1 2 3 4 Figure 7: Score distribution of sites along the (a) PC1 and PC2 axes and (b) PC1 and PC3 axes. High proportions of sand. in Makoba Bay. mud Sl Wt Sand DO Wc Wt Mud Sl (b) Wc Wd Mp Microphytobenthos. cephalus against principal components (PC1.05 indicate significant partial regression coefficients Coefficient 0.001 <0. Abiotic variables highly correlated with the axes are indicated (see Table 3). Wd = water depth and Mp = macrophyte cover an additional food source.873 0. Wc = water clarity.001 0.055 0. Sl = salinity.041 0.740 −3. DO = dissolved oxygen.064 t 10. the large cover of microalgal mats and the fine grain-sized sediment in the intermediate and upper sites of Kinani may explain the greater numbers of mullet there compared to Mapopwe.701 p <0.

2004. Okoth BK. Sheaves M. estuarine ecosystems (Rozas and Odum 1987). In the Indo-Pacific. Linnaeus) in two Florida habitats. World Wide Web electronic publication 2007. It could be that the relatively large sediment grain size at this creek. Field A (ed. Visual surveys reveal high densities of large piscivores in shallow estuarine nurseries.92 Mwandya. Ambio 24: 458–464. Australian estuaries and coastal waterways: a geoscience perspective for improved and integrated resource management. Viera 1991). van der velde G. 2006. 1995. Barletta M. Harris P. Western Australia. which all fluctuated during the heavy rains. Chubb CF. especially in developing countries where juvenile fish can be sourced from the wild. Temporal occurrence and size distribution of grey mullet juveniles (Pisces. The feeding periodicity of striped mullet (Mugil cephalus. van der Velde G. In addition. Australian Geological Survey Organisation. Bryceson and Gullström and high percentage of organic matter in the sediment at Kiongwe. . Collins MR. It is therefore unlikely that salinity alone has an effect on mullet densities. 2006. Aquaculture 19: 21–36. despite the high organic matter content at this creek. suggesting that this period could be appropriate for seasonal closure to protect recruitment. 2000. Marine Ecology Progress Series 256: 217–228. growth rates and movements of sea mullet. It is likely that a combination of factors such as turbidity. Record 2001/07. there are also seasonal aspects to consider. mainly composed of fragments of fresh calcareous algae (Halimeda spp. from oligohaline (0. London: Sage Publications. Most of the juveniles were caught from March to May. Anders Angerbjörn. Ambio 24: 482–486. Spatial and temporal distribution of Myrophis punctatus (Ophichtidae) and associated fish fauna in a north Brazilian intertidal mangrove forest. The higher densities of the small juveniles in the upper reaches and larger mullet at the lower and intermediate reaches were consistent with spatial pattern of other species in tidal. generously made their research facilities available. The prevalence of juveniles in the upper parts of all the creeks under study is consistent with other studies in tidal freshwater marshes (e.. Öhman MC. Hydrobiologia 426: 65–74. Nagelkerken I. Mgendi M. Cederlöf U. in the Swan-Avon river system. 2004. Cocheret de la Morinière E.g. Barletta-Bergan A. 1995. Iddi Khamis and Mcha Manzi who helped with field sampling. De Silva SS. Mwaipopo O. Australian Journal of Marine and Freshwater Research 32: 605–628. Saint-Paul U. Aldrichetta forsteri (Valenciennes). Special thanks are extended to Muhidin Abdallah. Zanzibar. Kenya. Marine and Freshwater Research 47: 857–868. Barletta-Bergan A. 1957. Zanzibar. 2000. Mohammed SM. We suggest that the habitat preferences in juvenile mullet in the mangrove creeks under study were determined by food availability and predator avoidance. is not a preferred food item for juvenile mullet. However. caused the apparent temporal variations in mullet density (Barletta et al. Kimani EN. Our findings could be relevant for resource managers in recognising productive coastal habitats in the Western Indian Ocean region. Hydrobiologia 563: 45–60. Wallace J. Mgaya. Rozas and Odum 1987). Spatial and temporal variation in water column nutrient concentrations in a tidally dominated mangrove creek. Mugilidae) in the estuarine systems of the Strymonikos Gulf (Greece).). Available at www. Olof Lindén. (1996) found that juveniles of striped mullet can thrive in a wide range of salinities. Grol MGG. Öhman. van der Meij H. Heap et al. 2001). In contrast to the other sites. 1981. biomass and diversity of estuarine fishes in tidal mangrove creeks of the lower Caetè Estuary (northern Brazilian coast. the dense macrophytic meadows at these creeks may not provide juvenile mullet with suitable habitat conditions. Grant CJ. Journal of Applied Ichthyology 20: 76–78. Johnstone RW. Michael Tedengren. Age structure. Chwaka Bay. Bryce S. Ekau W. Given their benthic-feeding behaviour. Journal of Sedimentary Research 27: 3–26. Tidal exchange in a warm tropical lagoon: Chwaka Bay. Liao (1981) and Ter Morshuizen et al. no mullet were caught at the lower reaches of Kiongwe and Kinani. Marine Ecology Progress Series 323: 75–82. prevailing recruitment patterns and tidal hydrodynamics may also have an influence on the mullet density (Chubb et al. Folk RL. Ambio 31: 518–527. 2002. 2007. The Institute of Marine 1981. Saint-Paul U. Schories D. Bertil Borg. Marine fisheries in Tanzania. Ward WC. References Baker R. Biology of juvenile grey mullet: a short review. From a management point of view. It was therefore unexpected to find the lowest abundance of mullet occurring at Mapopwe. What attracts coral reef fish to mangrove: habitat complexity or shade? Marine Biology 144: 139–144. 2001. Jiddawi NS. Nagelkerken I. Seasonal change in density.). 1996. and it is likely attributable to a combination of the presence of microphytobenthos on the sediment. low water clarity. Gazi. salinity and dissolved oxygen concentrations. Potter IC. and with increasing size they tend to move to lower reaches and deeper waters (Wenner and Beatty 1993). Sören Nylin and two anonymous referees are acknowledged for helpful comments on earlier versions of the manuscript. 1981. Hubold G. Our results further suggest that alteration of habitat characteristics and water conditions of mangrove ecosystems could have a profound effect on juvenile mullet populations. Gunilla Rosenqvist. Mugil cephalus L. Brazos River bar – a study in the significance of grain size parameters. Different surrounding landscapes may result in different fish assemblages in East African seagrass beds. Koutrakis ET. This concurs with the view that small fish in creeks and nearshore waters occur farther inland where salinity and water depth are generally lower. Discovering statistics using SPSS for windows. shallow depth and reduced salinity in farther upstream. Acknowledgements — This study was supported by the Sida/SAREC Bilateral Marine Science Programme between Sweden and Tanzania for research in marine biology. Pauly D. Heap A. 1980. Rydberg L. Radke L. Zanzibar. Ntiba JM. Mattias Sköld. east Amazon). Journal of Fish Biology 19: 307–315. Froese RJ. 2003. Ryand D. Dorenbosch M. Barletta M. 2000. Mullet were caught in the upper and intermediate sites of all four creeks as well as the lower sites of Mapopwe and Kiwani. Mwatha GK. and yellow-eye mullet. the results could be useful in locating suitable areas for aquaculture development. Smith R. Smith C. the lower reaches of Kiongwe and Kinani had considerable cover of submerged aquatic vegetation (seagrass and macroalgae). Fishes of a shallow tropical mangrove estuary. Lenanton RCJ. FishBase. as found during periods of heavy rains (April–May).5–5) to polyhaline waters (18–30).fishbase. Wakwabi EO. Heggie D.

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