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Ecology and Potential for Fishery of the Small Barbs (Cyprinidae, Teleostei) of Lake Tana, Ethiopia

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Ecology and Potential for Fishery of the Small Barbs (Cyprinidae, Teleostei) of Lake Tana, Ethiopia Promotoren: prof. dr. Jan W.M. Osse, hoogleraar in de algemene dierkunde, co-promotor: dr. Ferdinand A. Sibbing, universitair hoofddocent bij de leerstoelgroep Experimentele Zo¨ologie, dr. Jacobus Vijverberg, senior onderzoeker bij het NIOO-KNAW Centrum voor Limnologie, Nieuwersluis, overige leden promotiecommissie: prof. dr. Paul Skelton, JLB Smith Institute of Ichthyology, South Africa, prof. dr. Marten Scheffer, Wageningen Universiteit, dr. Leo A.J. Nagelkerke, Wageningen Universiteit, dr. Seyoum Mengistu, Addis Ababa University, Ethiopia, dr. Jan H. Wanink, Universiteit Leiden. ECOLOGY AND POTENTIAL FOR FISHERY OF THE SMALL BARBS (CYPRINIDAE, TELEOSTEI) OF LAKE TANA, ETHIOPIA Eshete Dejen Dresilign Proefschrift ter verkrijging van de graad van doctor op gezag van de rector magnificus van Wageningen Universiteit, prof. dr. ir. L. Speelman, in het openbaar te verdedigen op dinsdag 2 september 2003 des namiddags te vier uur in de Aula. Dejen Dresilign, E. (2003). The small barbs of Lake Tana. Doctoral thesis, Experimental Zoology Group, Wageningen University, P.O. Box 338, 6700 AH Wageningen, The Netherlands. Subject headings: small Barbus / taxonomy / ecology / zooplankton / zooplanktivorous / fishery / Lake Tana, Ethiopia ISBN: 90-5808-869-3 To Neter Alemu (my mother) and Dejen Dresilign (my father) vi Contents 1 Introduction 1 1.1 General introduction .................................... 1 1.1.1 Importance of small pelagics for fisheries ..................... 1 1.1.2 Zooplankton and zooplanktivorous fish in the food web of tropical waterbodies 2 1.1.3 Lake Tana and its environment .......................... 3 1.1.4 Lake Tana and its fish community ........................ 4 1.1.5 Initiation and objectives of the research ..................... 6 1.1.6 Sampling Programme ............................... 8 1.2 Introduction to the various chapters ........................... 9 1.2.1 Chapter 2; Taxonomy of small barbs ....................... 10 1.2.2 Chapter 3; Barbus tanapelagius, a new small barb species from Lake Tana . 10 1.2.3 Chapter 4; Zooplankton composition and distribution in relation to environ- mental factors ................................... 11 1.2.4 Chapter 5; Resource partitioning ......................... 12 1.2.5 Chapter 6; Reproductive strategies ........................ 13 1.2.6 Chapter 7; Parasitism and its effect on small barbs ............... 14 1.2.7 Chapter 8; Estimation of biomass and production of the small barbs and potential for fishery ................................ 15 2 The ‘small barbs’ Barbus humilis and B. trispilopleura of Lake Tana (Ethiopia): are they ecotypes of the same species? 17 2.1 Introduction ......................................... 18 2.2 Materials and methods ................................... 20 2.2.1 Sampling sites and collection ........................... 20 2.2.2 Identification of species .............................. 20 2.2.3 Measurements and observations .......................... 22 2.2.4 Cluster analysis ................................... 24 2.3 Results: Identification and spatial distribution of B. trispilopleura and B. humilis ....... 25 2.3.1 Comparison with Boulenger’s description .................... 25 2.3.2 B. trispilopleura .................................. 30 2.3.3 B. humilis ...................................... 30 2.3.4 Spots and body colour ............................... 31 2.3.5 Cluster analysis ................................... 32 2.3.6 Ecology and reproduction ............................. 34 2.4 Discussion .......................................... 35 3 Barbus tanapelagius, a new species from Lake Tana (Ethiopia): its morphology and ecology 39 3.1 Introduction ......................................... 39 3.2 Materials and methods ................................... 42 3.2.1 Collection of type material and morphometrics ................. 42 vii 3.2.2 Spatial distribution sampling program ...................... 42 3.2.3 Analysis of gut content .............................. 43 3.3 Results ............................................ 43 3.3.1 Morphology ..................................... 43 3.3.2 Spatial distribution ................................. 44 3.3.3 Diet composition .................................. 47 3.4 Discussion .......................................... 47 4 Temporal and spatial distribution of microcrustacean zooplankton in relation to turbidity and other environmental factors in a large tropical lake (L. Tana, Ethiopia) 51 4.1 Introduction ......................................... 51 4.2 Materials and methods ................................... 53 4.2.1 Study site ...................................... 53 4.2.2 Environmental conditions ............................. 53 4.2.3 Zooplankton sampling and analysis ........................ 54 4.2.4 Zooplankton taxonomy .............................. 55 4.2.5 Data analysis .................................... 56 4.3 Results ............................................ 57 4.3.1 Environmental factors and seasonality ...................... 57 4.3.2 Spatial and temporal patterns in the zooplankton community ......... 60 4.3.3 Structure of the zooplankton community in relation to environmental variables 62 4.4 Discussion .......................................... 64 4.4.1 Species composition ................................ 64 4.4.2 Zooplankton distribution ............................. 65 4.4.3 Conclusions ..................................... 67 5 Resource partitioning in a tropical fish assemblage of obligate and facultative zooplanktivores (Lake Tana, Ethiopia) 69 5.1 Introduction ......................................... 70 5.2 Materials and Methods ................................... 72 5.2.1 Study area and sampling ............................. 72 5.2.2 Diet analysis .................................... 74 5.2.3 Measures of diet and habitat overlap and niche breadth ............ 74 5.2.4 Morphometry of feeding structures ........................ 75 5.3 Results ............................................ 75 5.3.1 Morphometry of feeding structures ........................ 75 5.3.2 Spatial distribution ................................. 80 5.3.3 Diet composition .................................. 82 5.3.4 Habitat and diet overlap .............................. 87 5.3.5 Niche breadth .................................... 89 5.4 Discussion .......................................... 89 5.4.1 Potential and actual food niches ......................... 90 5.4.2 Diet and habitat overlap .............................. 93 5.4.3 Predation by large piscivores ........................... 94 5.4.4 Role of zooplanktivory in tropical food webs ................... 95 5.4.5 Fisheries management aspects ........................... 96 6 Reproductive strategies of two sympatric ‘small barbus’ (Barbus humilis and B. tanapelagius) in Lake Tana, Ethiopia 97 6.1 Introduction ......................................... 97 6.2 Materials and Methods ................................... 99 6.2.1 Sampling of fish .................................. 99 6.2.2 Measurements and data analysis ......................... 100 viii 6.3 Results ............................................ 101 6.3.1 Environmental conditions ............................. 101 6.3.2 Zooplankton densities ............................... 102 6.3.3 Abundance and distribution of small barbs ................... 102 6.3.4 Length-weight relationship ............................ 103 6.3.5 Tapeworm infection ................................ 103 6.3.6 Sex ratio ...................................... 104 6.3.7 Breeding season and area ............................. 104 6.3.8 Ova diameter frequency .............................. 104 6.3.9 Fecundity ...................................... 104 6.3.10 Mean size at maturity ............................... 108 6.4 Discussion .......................................... 109 6.4.1 Timing of spawning ................................ 109 6.4.2 Frequency of spawning ............................... 111 6.4.3 Fecundity, egg size and maturity ......................... 111 6.4.4 Biased sex ratio ................................... 113 6.4.5 Adaptation to lacustrine conditions ........................ 113 7 Dynamics of cestode infection and its effects on small pelagic barbs (Barbus humilis and B. tanapelagius) in Lake Tana, Ethiopia 115 7.1 Introduction ......................................... 116 7.2 Materials and methods ................................... 117 7.3 Results ............................................ 120 7.3.1 Environmental conditions ............................. 120 7.3.2 Cestode infection among ‘small’ cyprinids .................... 120 7.3.3 Infection prevalence across habitats and season ................. 121 7.3.4 The effect of parasite on reproductive strategies ................. 123 7.4 Discussion .......................................... 124 7.4.1 Fish species and size class sensitivity ....................... 124 7.4.2 Habitat variability ................................. 125 7.4.3 Seasonal effects on parasite prevalence ...................... 126 7.4.4 Parasite induced reaction norms on life-history strategies ........... 127 8 Biomass and production of pelagic small barbs (Barbus humilis and B. tanapelag- ius 129 8.1 Introduction ......................................... 129 8.2 Materials and methods ................................... 131 8.2.1 Study area ..................................... 131 8.2.2 Sampling sites and collection ........................... 131 8.2.3 Data analysis ...................................
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