The Barbs of Lake Tana, Ethiopia: Morphological Diversity and Its Implications for Taxonomy, Trophic Resource Partitioning, and Fisheries Promotor: Prof
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The barbs of Lake Tana, Ethiopia: morphological diversity and its implications for taxonomy, trophic resource partitioning, and fisheries Promotor: Prof. Dr. J.W.M. Osse Hoogleraar Algemene Dierkunde CO-promotor: Dr. F.A. Sibbing Universitair Hoofddocent Functionele Diermorfologie aan de Ieerstoelgroep Experimentele Dierkunde, departement Dienvetenschappen Leo Nagelkerke The barbs of Lake Tana, Ethiopia: morphological diversity and its implications for taxonomy, trophic resource partitioning, and fisheries Proefschrift ter verkrijging van de graad van doctor op gezag van de rector magnificus van de Landbouwuniversiteit te Wageningen, dr. C.M. Karssen, in het openbaar te verdedigen op dinsdag 7 oktober 1997 des namiddags te half twee in de aula Abstract Nagelkerke, L.A.J. 1997. The barbs of Lake Tana, Ethiopia: morphological diversity and its implications for taxonomy, trophic resource partitioning, and fisheries. Doctoral thesis. Esperimental Animal Morphology and Cell Biology, Wageningen Agricultural University, P.O. Box 338, 6700 AH Wageningen. The Netherlands ISBN: 90-5485-755-2 Subject headings: Barbus 1 biodiversity 1 cyprinids 1 ecomorphology 1 feeding 1 food partitioning 1 lakes l species flock I sustainable fisheries The rediscovery of a unique species flock of cyprinid fish, its taxonomy and its feeding-biology are described. Fourteen species of barbs (Barbus spp, Cyprinidae, Teleostei) were found in highland (1800 m) Lake Tana, in northwestern Ethiopia. Lake Tana is an isolated fresh-water system, because its only outflowing river, the Blue Nile, of which it is the source, drops over 40 m high waterfalls only 35 km from the lake. The carplike barbs in the lake are unique and show a wide variety of discrete, and consistent types, widely differing in 1) size and shape, 2) feeding behaviour, and 3) spatial distribution. Together with 4) the early morphological divergence of juvenile barbs, 5) the reproductive segregation of the adults, and 6) part of their genetic characters, this diversity was decisive in our description of fourteen endemic species. seven of them new to science. Lake Tana was formed by volcanic blocking of the Blue Nile and subsequent flooding of the Lake Tana basin. The Barbus species probably evolved within the lake itself from one common ancestor, very similar to the present-day, riverine B. intermedius, making it a unique species flock. The driving force of their evolution most likely involved radiation into the new lacustrine trophic niches that became available when Lake Tana filled up. The presence of this group of very closely related fishes provides a unique opportunity to study the evolution of adaptation, and the relation of morphology and ecology: it forms a 'natural laboratory'. By using an ecomorphological approach, based on properties of fish foods, and functional morphological studies of the relations between fish parameters and their significance in dealing with food properties. predictions on the diets and food partitioning of the barbs have been made. These predictions were based 011 a large set of characters (33, which made the method robust and allowed for accurate resolving power Testing of the predictions with field data revealed that individual diets, but especially food partitioning can bc well predicted. This new method provides insights into the dynamic trophic interactions among species without the need for an extensive ecological sampling programme, and could be instrumental in predicting shifts in fish fauna composition, due to environmental impact, such as overfishing or thc introduction of ncw species. The understanding and prediction of such shifts will help in developing a strategy towards sustainable fisheries and the protection of biodiversity. This research was supported by: NWO-WOTRO, project number W88-176 Interchurcl~Foundation EthiopiaIEritrea (ISE), Urk Interchurch Coordination Commission for Develop~nent(ICCO), Zeist, Fisheries Resources Development Department of the Ministry of Agriculture, Addis Ababa Bureau of Agriculture of theA~nharaNational Regional State, Bahar Dar Ethiopian Orthodox Church (EOCIDICAD), development department Agricultural University Wageningen Cover: Lake Tana with Great white pelicans (Pelecanus onocuotalus) and Black kitcs (Mi1vzi.s migrans) at Bahar Dar in the rainy season: photograph by G.J.M. van Snik. Cover design: Wim Valen. Caput Nili quaerere ("De bronnen van de Nijl zoeken": een zeer lastig kanvei beginnen) +fifl+CI h9MA flh% +&ss\ ("Langzaam maar zeker zal een ei pootjes krijgen en gaan lopen"; Arnhaars spreekwoord) Vlum en nzijn vuren lach drinkt uii onizaggelijke schnlen van luchi en aarde den opalen dag (H. Marsman, Verzamelde Gedichten, 1938, uitgeverij Querido) Chapter 1 General Introduction 7 Chapter 2 The barbs (Barbus spp.) of Lake Tana: a forgottten species flock? 31 published: L.A.J. Nagelkerke, F.A. Sibbing, J.G.M. van den Boogaart. E.M.R.R. Lammens & J.W.M. Osse (1994). Environtnental Biology ofFishes 39: 1-22. Chapter 3 Morphological divergence during growth in the large barbs 57 (Barbus spp.) of Lake Tana, Ethiopia. published: L.A.J. Nagelkerke, F.A. Sibbing & J.W.M. Osse (1995). Netherlands Journal of Zoology 45: 43 1-454. Chapter 4 Reproductive segregation among the large barbs (Barbus intermedius 79 complex) of Lake Tana, Ethiopia. An example of intralacustrine speciation? published: L.A.J. Nagelkerke & F.A. Sibbing (1996). Journal ofFish Biologv 49: 1244-1266. Chapter 5 A revision of the large barbs (Barbus spp., Cyprinidae, Teleostei) of 105 Lake Tana, Ethiopia, with a description of seven new species. accepted: L.A.J. Nagelkerke & F.A. Sibbing. Zoologische Verhandelingen Leiden. Chapter 6 Trophic segregation within the Barbus species flock of Lake Tana, 171 Ethiopia: 1. Diets predicted from food properties and fish morphology. in preparation: F.A. Sibbing & L.A.J. Nagelkerke. Chapter 7 Trophic segregation among the Barbus species flock of Lake Tana, 229 Ethiopia: 2. Test of diet predictions and food partitioning. in preparation: L.A.J. Nagelkerke & F.A. Sibbing. Chapter 8 In Lake Tana, a unique fish fauna needs protection. published: L.A.J. Nagelkerke, M.V. Mina, T. Wudneh, F.A. Sibbing h J.W.M. Osse (1995). Bioscience 45: 772-775. References Summary / Samenvatting Dankwoord en epiloog Curriculum vitae List of publications Chapter l General introduction Leo A.J. Nagelkerke' ' Agricultural University, Wageningen Institute of Animal Sciences (WIAS), Experimental Zoology Group, Marijkeweg 40, 6709 PG Wageningen, The Netherlanh In 1986, the Lake Tana Fisheries Resources Development Program (LTFRDP) was started by the Ethiopian Ministry of Agriculture, in cooperation with the Development Branch of the Ethiopian Orthodox Church (EOC). The project was supported by the lnterchurch Foundation Ethiopia (ISE) from Urk, The Netherlands. At this time, hardly anything was known about the identity and dynamics of the Lake Tana fish resources. Therefore, Tesfaye Wudneh was assigned as fisheries expert, to investigate the fish stocks. One of the species in the lake, Barh1r.s intermedirrs Riippell showed such an immense morphological variability that it was suspected that there were at last several different populations of barbs in Lake Tana and possibly several species. The presence of different species or populations, and therefore of different units of fish stock complicates rational management of the resources. To investigate the identity and the evolution of the Lake Tana Barbus biodiversity and its role in the ecosystem, a research project was initiated. The results of that study are reported in this thesis. Before I will evaluate the results of the research project, I will first introduce Lake Tana in a broad sense: its topography, climate, flora and fauna in general, and its fish in particular. Secondly I will describe the context and history of my research project, its aims, and its progress in answering the major questions. The latter is, of course, subject of the main body of this thesis and Water- falls Fig.l.1. A map of Ethiopia and Lake Tana. The southern Bahar Dar Gulf is indicated, as well as the Blue Nile and the waterfalls in it. 8 General introduction every chapter will be discussed in short. Finally there will be a short general discussion in which I will try to come to a synthesis of the research until now. This involves addressing some preliminary results that are not presented in the following chapters, and making some generalizations. This also brings me to future research that still should be performed to come to an even better understanding of Lake Tana. Short introduction to Lake Tana Topography and geology Lake Tana is situated in the north-western highlands of Ethiopia in the provinces of Gojjam and Gonder, at an altitude of approximately 1800 m (Fig. l.l). It is the source of the Blue Nile (or Abbay as it is called in Amharic, the chief language of this part of Ethiopia), and as such it has been a place that was searched for since the times of Julius Caesar and Alexander the Great. The first European to see the legendary source was the Portugese Jesuit Pedro Paez, in 16 13. The so-called Tana-rift, in which Lake Tana lies, rs a shallow trough which is not connected to the main Eastern Rift Valley as such, but it certainly has some relation to it (Baker et al. 1972, Mohr 1962). The lake probably formed through volcanic blocking of the Blue Nile in early Pleistocene times (Mohr 1962). Subsequently the Lake Tana basin filled up. It now covers an area of approximately 3 150 km2, and is Ethiopia's largest lake. Four larger, permanent rivers tribute to the lake (the largest is Gelgel Abbay; the 'small Blue Nile", which is c. 100 km), as well as many short, seasonal streams. The lake is shallow with an average depth of 8 m and a maximum depth of 14 m. Its bottom consists of volcanic basalts, usually covered with a muddy substratum with only little organic matter (1.2-2.3%: Gasse 1987). At some places volcanic peaks in the lake bottom form reefs, or even islands (Fig.