Cichlid Species Diversity in Naturally and Anthropogenically Turbid Habitats of Lake Victoria, East Africa
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Aquat Sci (2013) 75:169–183 DOI 10.1007/s00027-012-0265-4 Aquatic Sciences RESEARCH ARTICLE Cichlid species diversity in naturally and anthropogenically turbid habitats of Lake Victoria, East Africa Frans Witte • Ole Seehausen • Jan H. Wanink • Mary A. Kishe-Machumu • Marjolein Rensing • Tijs Goldschmidt Received: 11 October 2011 / Accepted: 10 May 2012 / Published online: 8 June 2012 Ó The Author(s) 2012. This article is published with open access at Springerlink.com Abstract During the past decades, major anthropogenic haplochromines from sub-littoral areas during the pre- environmental changes occurred in Lake Victoria, collapse, collapse and recovery periods. Water over mud including increased predation pressure due to Nile perch is murkier and poorer in oxygen than water over sand, and introduction, and decreases in water transparency and differences in haplochromine communities in these natural dissolved oxygen concentrations due to eutrophication. habitats during the pre-collapse period may predict the This resulted in a collapse of the haplochromine cichlids effects of anthropogenic eutrophication during the col- in the sub-littoral waters of the Mwanza Gulf in lapse and recovery periods. In the pre-collapse period, 1986–1990, followed by a recovery of some species in the haplochromine densities over sand and mud did not differ, 1990s and 2000s, when Nile perch densities declined. but species richness over sand was 1.6 times higher than We studied two data sets: (1) haplochromines from over mud bottoms. Orange- and white-blotched colour sand and mud bottoms in the pre-collapse period; (2) morphs were most common at the shallowest sand station. More specifically, insectivores and mollusc-shellers had higher numbers of species over sand than over mud, whereas for mollusc-crushers no difference was found. F. Witte (&) Á J. H. Wanink Á M. A. Kishe-Machumu Á Laboratory experiments revealed that mollusc shelling was M. Rensing Á T. Goldschmidt more affected by decreased light intensities than mollusc Institute of Biology Leiden, University of Leiden, crushing. During the pre-collapse period, spawning P.O. Box 9505, 2300 RA Leiden, The Netherlands e-mail: [email protected] occurred year-round in shallow areas with hard substrates and relatively clear water. In deeper areas with mud bot- F. Witte toms, spawning mainly occurred during months in which Netherlands Centre for Biodiversity, Naturalis, water clarity was high. No effects of hypoxia on spawning P.O. Box 9517, 2300 RA Leiden, The Netherlands periods were found. It follows that clearer water seems to O. Seehausen support differentiation in feeding techniques as well as Department of Aquatic Ecology and Evolution, year-round spawning, and both may facilitate species Institute of Ecology and Evolution, University of Bern, coexistence. Water clarity is also known to be important Baltzerstrasse 6, 3012 Bern, Switzerland for mate choice. These observations may explain why, O. Seehausen since the decline of Nile perch, haplochromine densities EAWAG Centre for Ecology, Evolution and Biogeochemistry, have recovered, the numbers of hybrids increased and Seestrasse 79, 6047 Kastanienbaum, Switzerland species diversity in the current eutrophic sub-littoral J. H. Wanink waters has remained 70 % lower than before the envi- Koeman en Bijkerk bv, Ecological Research and Consultancy, ronmental changes. P.O. Box 111, 9750 AC Haren, The Netherlands Keywords Dissolved oxygen concentrations Á M. A. Kishe-Machumu Tanzania Fisheries Research Institute, Dar es Salaam Center, Eutrophication Á Haplochromine cichlids Á P.O. Box 78850, Dar es Salaam, Tanzania Light conditions Á Molluscivores Á Spawning periods 123 170 F. Witte et al. Introduction anthropogenic nutrient pollution. Therefore, we analyzed haplochromine catches from littoral sand and mud bottoms In Lake Victoria, more than 500 haplochromine cichlid in Butimba Bay. These catches were made in 1979 and species, 99 % of which are endemic, made up some 80 % of 1980, before the major ecological changes. We compared the bottom dwelling fish mass until the beginning of the differences in densities and species diversity of haplo- 1980s (Greenwood 1974; Kudhongania and Cordone 1974; chromines between these stations, with the differences in Witte et al. 2007a). During the 1980s, the majority of the densities and diversity before and after the environmental haplochromine species in the sub-littoral (6–20 m deep) and changes in the sub-littoral waters of the Mwanza Gulf. offshore areas disappeared (Ogutu-Ohwayo 1990; Witte et al. 1992). This disappearance coincided with the upsurge of the introduced predator, Nile perch, Lates niloticus (L.) Methods (Goudswaard et al. 2008), and with strong eutrophication of the lake. The latter resulted in increased algal blooms and Study period and areas decreases in both water transparency and dissolved oxygen concentrations (Kaufman 1992; Hecky et al. 1994; See- In the present study, three periods are recognized: pre-col- hausen et al. 1997a; Wanink et al. 2001). The process of lapse (1973–1984), collapse (1986–1990), recovery eutrophication had already started decades earlier, with the (2001–2006) (Table 1). The division in these three periods is strong growth of human populations along the lake’s shores based on the following observations in the Mwanza Gulf: (1) (Verschuren et al. 2002; Hecky et al. 2010). Disruption of dominance by weight, and great diversity, of haplochromine the original food web and increased deforestation for Nile cichlids in bottom trawl catches in sub-littoral waters of the perch smoking, may have additionally contributed to Mwanza Gulf until 1984 (Witte et al. 1992); (2) the almost eutrophication and algal blooms. Beginning in the first half complete collapse of the haplochromine populations in sub- of the 1990s, after the decline of Nile perch probably as a littoral waters of the Mwanza Gulf (except for those in rocky result of over-fishing (Matsuishi et al. 2006; Kayanda et al. areas with clear water) in the period between 1986 and 1990 2009; Witte et al. 2009), a resurgence of some haplochro- (Witte et al. 1992; Seehausen et al. 1997a), together with mine species was observed (Seehausen et al. 1997b; Witte algal blooms, and low DO concentrations and Secchi values et al. 2007a, b). However, the initial diversity of haplo- since 1986 (Wanink et al. 2001; Witte et al. 2005); (3) chromines has not fully recovered. recovery of some haplochromine species in sub-littoral During the past years, a lively debate has been ongoing waters since the 1990s (Witte et al. 2007b). regarding impacts of Nile perch predation, over-fishing and The main study area comprised ten sampling stations (A eutrophication on the diversity and abundance of haplo- to J) across the Mwanza Gulf (Fig. 1). Stations A to D with chromine cichlid species in Lake Victoria. It appears that approximately the same length (each about 150 m) are over-fishing may have had a negative impact on haplo- located in Butimba Bay, which has a size of less than chromines locally, but this is unlikely to be a major driver 0.5 km2 (Fig. 1). Stations A and B have sand bottoms, of the lake-wide loss of cichlid diversity (Witte et al. stations C and D mud bottoms. The depth range of stations 2007b). Discriminating between the effects of Nile perch A and C is 2–4 m; that of stations B and D 4–6 m. The sub- predation and eutrophication is more difficult, as they littoral stations E to J with mud bottoms and depths ranging occurred in the same period (Seehausen et al. 1997a; from 7 to 14 m were pooled in the present study and further Verschuren et al. 2002; Witte et al. 2007a, b; Kolding et al. referred to as stations E–J. 2008; Hecky et al. 2010). Moreover, there are two major aspects of eutrophication that may affect haplochromine Environmental parameters communities: decreases in water transparency and decrea- ses in dissolved oxygen (DO) concentrations. As the original study in the pre-collapse period was not Before the environmental changes in the lake, we sam- aimed at relating species diversity with environmental pled relatively pristine sand and mud habitats that differed parameters, the latter were only collected occasionally in light conditions and in DO concentrations close to the (Table 1). To establish the distribution of sand and mud sediment. Mud bottoms are covered with a thick layer of bottoms in Butimba Bay, substrate samples were collected organic debris that easily stirs up, making the water near with an Ekman dredge (Wildlife Supply Company, model the sediment hypoxic and murky. Thus, comparison of 196). A van Dorn water sampler was used to get an densities and species richness of haplochromine cichlids in impression of the thickness of the flocculent mud layer. pristine sand and mud stations offers a unique possibility to Secchi measurements at stations A to D were made in identify potential impacts of decreased water transparency March and April 1980 (pre-collapse period) and in Feb- and DO concentrations that result from lake-wide ruary and April 2002 (recovery period). In the latter period 123 Cichlid species diversity in turbid habitats 171 Table 1 Summary of data sets and their use in the present study Area Data Month and year Period Aim Butimba Bay Comparing species diversity over Station A–D, undisturbed sand and mud 2–6 m bottoms Secchi values March; Apr 1980 Pre-collapse Differences over sand/mud Febr; Apr 2002 Recovery Differences over sand/mud Light (Lux) Febr; Apr 2002, March 2006 Recovery Differences