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A Seasonal Survey of the Fish of the Kavango River Floodplain Along The

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A Seasonal Survey of the Fish of the Kavango River Floodplain Along The A Decade of Namibian Fisheries Science Payne, A. I. L., Pillar, S. C. and R. J. M. Crawford (Eds). S. Afr. J. mar. Sci. 23: 449–464 2001 449 FISH RESPONSE TO THE ANNUAL FLOODING REGIME IN THE KAVANGO RIVER ALONG THE ANGOLA/NAMIBIA BORDER C. H. HOCUTT* and P. N. JOHNSON† The results of the first seasonal survey of the fish of the Kavango River floodplain along the Angola/Namibia border are reported. The river experiences peak flooding from February through June, with the 375-km long flood- plain extending up to 5 km across. The floodplain was sampled five times in 1992 by seine, fish traps and rotenone. The data indicated a pronounced structural and functional response of the fish community in relation to the alternating flood and drought conditions in the river. Catch per unit effort and diversity were highest during months of peak flooding (May and June), and lowest during the month of least flow (November). The reproductive strategies of K-selected piscivorous cichlids and tigerfish were in advance of flooding. Many r-selected invertivores, especially cyprinids, were in relative synchrony with flooding and the stimulation of littoral zone plant growth, while other invertivores lagged the cyprinids. Herbivores had lowest relative abundance during peak flooding; this seemingly inverse relationship with the invertivores should not be interpreted as replacement, but rather the swamping of the system with young-of-the-year r-selected invertivores. The data support the Flood Pulse Concept, which hypothesizes that flooding is the major “driver” of productivity in lowland or floodplain rivers. Key words: behaviour, floodplain fishery, migration, Okavango Delta, r-/K-selection, subsistence fishery A seasonal survey of the fish of the Kavango River long-term shifts in the structural and functional com- floodplain along the Angola/Namibia border was ponents of the fish community. However, a funda- conducted during 1992 and 1993, as a cooperative ef- mental requirement of utilizing any measure of envi- fort between the University of Namibia, the Namibian ronmental alteration of an aquatic system is the need Ministry of Fisheries and Marine Resources (MFMR) to understand natural variability, both spatially and and the University System of Maryland. The Kavango temporally (Bruton and Jackson 1983). This manuscript (also known as the Cubango) flows south from its addresses that need, at least in part. origin in the highlands of central Angola, and then Knowledge of the seasonal structure and functioning turns east to form the 415-km long border between of the floodplain fishery is largely anecdotal, despite Namibia and Angola (Fig. 1); approximately 375 km laudable efforts made by the MFMR. As the MFMR of this reach has the lowest gradient in the drainage, seeks solutions to manage the fishery in light of in- with extensive floodplains (Hocutt et al. 1994), and creasing demand, it is critical to understand the rela- is the study area for this manuscript (Fig. 2). The tionship between seasonal flooding and productivity, river then flows southeast through Namibia for an- particularly given the current regional plans to change other 35 km before it reaches Botswana and terminates or interrupt the natural processes of this important flood- in the 15 000 km2 Okavango Delta. plain river system (Breen et al. 1984). The Kavango River remains a pristine catchment The survey described here differs from historical in the context of the definition used in the western efforts of the MFMR, which for a variety of reasons world, with minor human impact along the Namibia/ has been restricted to surveying the Kavango once Angola border other than through organic enrichment only per year, thus not including a cycle that can be and riparian zone mismanagement. Development in correlated to the annual flooding regime. The database the basin has to date been retarded; however, with here is taken from Hocutt et al. (1994), whose data political normalization of the region, various regional hinted (their Fig. 6) that a seasonal trend might be schemes for water resource utilization are being con- discernible if interpreted further. sidered for the catchment (Baldwin 1991). For this Jubb’s (1967) and Skelton’s (1993) works on the reason, Hocutt et al. (1994) and Hay (1995) developed freshwater fish of southern Africa and Poll’s (1967) the rationale for a routine monitoring protocol based tome on the fish of Angola were hallmark contributions on the Index of Biotic Integrity (IBI) as a measure of to regional ichthyology. Castelnau (1861) described * Formerly [email protected]; BENEFIT Secretariat, P.O. Box 912, Swakopmund, Namibia. E-mail: [email protected] † MEVATEC Corporation, Fisheries Engineering Team, P.O. Box 40, N. Bonneville, WA 98639, USA Manuscript received November 2000; accepted June 2001 A Decade of Namibian Fisheries Science 450 South African Journal of Marine Science 23 2001 Fig. 1: Map of the Kavango (Cubango) River basin (after Skelton et al. 1995) the first specimens from the Okavango Swamp region 1965, Bell-Cross 1968, 1972, 1976, Hay et al. 1999), of Lake Ngami, based on specimens collected by although the historical relationships remain unre- Daviaud (Jubb and Gaigher 1971). The first account solved (Gaigher and Pott 1973, Greenwood 1983). of the upper Cubango River fish fauna in Angola was Bell-Cross (1968, 1972, 1976), Bowmaker et al. given by Pellegrin (1936), and three years later the (1978) and Bell-Cross and Minshull (1988) reviewed first collection of fish from the Kavango River in the fish of the upper Zambezi River system, and their Namibia (then called South West Africa) was obtained checklists of fish included the Okavango drainage. by Eedes (Jubb and Gaigher 1971). Explanation of current distributions may be the pres- The ichthyofauna of the Namibia and Botswana ence of obstacles or barriers during periods of former portion of the Kavango drainage is well documented drainage connections (Gaigher and Pott 1973), such (Jubb and Gaigher 1971, Skelton et al. 1983, 1985, barriers being physical (Crass 1962), ecological Skelton and Merron 1984, 1985, 1987, Minshull 1985, (Jackson 1962), climatic (Stuckenburg 1969, Gaigher van der Waal 1987, 1991, Merron 1991, 1993, van Zyl and Pott 1973), behavioural (Bell-Cross and Minshull 1992, Hay et al. 1999), but the Angolan portion of the 1988), or the result of competitive interactions. Gaigher catchment is only modestly known, primarily through and Pott (1973) list other influences on current distri- the works of Poll (1967). Merron and Bruton (1985, bution patterns: extermination by predation, retreat 1989, 1995) considered the fishery of the Delta in due to competition, and catastrophic destruction of Botswana. The latter work, particularly, parallels the fish life in intermediate areas. research reported here in that seasonal shifts in abun- The river continuum concept (Vannote et al. 1980) dance and diversity of the Delta’s fish community are is a widely accepted model of longitudinal structure discussed. and functioning of lotic systems, although it is chal- From a zoogeographic perspective, the ichthyofauna lenged in some circles (e.g. Statzner and Higler of the Kavango has affinities with the adjacent Cuveili, 1985, Davies et al. 1993). Junk et al. (1989) outlined Cuando (Mashi) and upper Zambezi drainages (Jubb the flood pulse concept, and hypothesized that flooding 1964, 1967, Gabie 1965, Jubb and Farquharson is the major “driver” of lowland or floodplain rivers. 2001 Hocutt & Johnson: Seasonal Response of Kavango River’s Floodplain Fish 451 Fig. 2: Zones of the lower Kavango River and the study area (Zone 2) along the Namibia/Angola border (after Hocutt et al. 1994) The concept implies that the frequency, magnitude plain, the flats may have standing trees, or appear as and rates of rise and fall of flooding will be reflected a savannah, depending upon location. Smith (1976) in the ecological character of the associated biota and Bethune (1991) discussed the vegetation along (Davies et al. 1993). Junk et al. (1989) defined the the Kavango floodplain. floodplain as an aquatic-terrestrial transition zone Hocutt et al. (1994) subdivided the Kavango channel that excludes permanent lotic and lentic habitats within into four latitudinal zones along the Namibia/Angola the zone of inundation, so emphasizing the floodplain border, based on the occurrence of extended flood- as an area of alternation of wetting and drying phases plain and associated habitats (Fig. 2): Zone 1, from (Davies et al. 1993). Katwitwi to Kasivi, has little floodplain development The Kavango River floods seasonally, normally and mostly shallow-water environments with sand from December to July, with a general maximum 6 m and/or rock substrata. Small rapids are scattered above low flow conditions and an average discharge throughout, with prevalent submerged vegetation and of 10 500 × 106 m3 per year at the Botswana border aquatic weed beds in a rather well defined river bed. (Baldwin 1991). Flooding peaks from February In Zone 2 from Kasivi to Mbambi, the primary study through June, i.e. some 3–4 months later than peak area discussed here, the floodplain is well developed, rainfall in Angola. The floodplain can extend more extending at times to >5 km in breadth (Sandlund than 2 km away from the stream bed during flooding. and Tvedten 1992). The river valley is wider, with Local rainfall is usually ineffectual in the flooding the main channel seasonally broadening into innu- pattern itself, other than perhaps assisting the saturation merable inferior back channels, bays and oxbow of the water table. ponds forming over submerged pasture-land during According to the definition of Welcomme (1979a), flooding, and connected by faster-flowing anasto- the Kavango River floodplain is a fringing floodplain mosed channels; these features usually dry out as the lying between river valley walls (v. internal, e.g. the river retreats. The substratum in the main river is Okavango Delta).
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