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Multiple repeated patterns of adaptive radiation were revealed in cyprinid fish inhabiting the compact geographic region of the ethiopian Highlands. We found four independently evolved radiations in the evolutionary hexaploid (2n = 150) Labeobarbus lineage based on matrilineal relationships of >800 individuals. Each radiation displayed similar patterns of mouth phenotype diversification, and included ecomorphs/species of the generalized, lipped, scraping (one or two), and large-mouthed (one to three) types. All radiations were detected in geographically isolated rivers, and originated from different ancestral populations. This is the first documented case in which numerous parallel radiations of fishes occurred-via different ways-in a riverine environment. Some radiations are very recent and monophyletic, while others are older and include ecomorphs that originated in separate mini flocks and later combined into one. The diversification bursts among Ethiopian Labeobarbus were detected in the mid-upper reaches of rivers (1050-1550 m above sea level), which likely offer ecological opportunities that include diverse habitats yet poor fauna (i.e. lower competition and relaxed selection). this promising example of parallel evolution of adaptive radiation warrants further investigation.In the pioneering work on South African cichlids and catfishes, Joyce et al. 21 and Day et al. 19 suggested that the riverine flocks are remnants of radiations that emerged in a paleolake. This suggestion is corroborated by the finding of the Lake Tanganyika endemics beyond the lake basin 33 . While some riverine radiations clearly have lacustrine origins, it is likely that other riverine radiations originate from sources other than lakes. Hence, we aimed to test whether the same environmental factors that trigger lacustrine radiation can also act in rivers.In general, the main prerequisites for adaptive radiation are: (1) environmental stability for evolutionarily significant periods (starting from thousands of years ago); and (2) physical isolation from the river drainages harboring diverse fish faunas 15,20,31 . The latter serves a dual purpose: first, it provides the ecological opportunities for niche divergence at the initial stages of radiation, and later it prevents the extinction of diverged forms caused by competition with highly specialized species from other systematic groups. These processes can operate in specific and rather rare parts of the riverine network. For example, the segment of the Genale River that harbours the radiating assemblage of distinct forms of the Labeobarbus is situated in the apparently old canyon, isolated by the Baratieri Falls from the lower reaches of the river system and characterized by the depauperate fish fauna 20 .If the fish radiations in rivers are similar to the evolutionary phenomena experienced in lakes, they should display similar evolutionary patterns. It is well known that adaptive radiations are often repeatable in different organisms as exemplified from spiders 34 , fish 35 , and liz...

Large African barbs of the genus Labeobarbus constitute a hexaploid lineage (2n = 150). This group is widely distributed in African freshwaters, and exhibits profound phenotypic variation that could be a prerequisite for adaptive radiation. Using morphological, molecular, and stable isotope analyses, we investigated whether an adaptive radiation has occurred in a riverine assemblage of the Labeobarbus gananensis complex. This complex is composed of six phenotypically distinct sympatric forms inhabiting the Genale River (Ethiopian highlands, East Africa) in the Juba and Wabe–Shebelle drainages (Indian Ocean basin). Of the six forms, five were divergent in their mouth morphology, corresponding to generalised, lipped, scraping (two forms), and large‐mouthed phenotypes. One form had no mouth specialisation, but differed from the others in body shape (short and deep body; short form). Stable isotope analysis revealed differences in 15N among these forms, representing different foraging strategies (omnivorous, scraping, and piscivorous). Phylogenetic analysis of two mitochondrial DNA markers confirmed the monophyly of L. gananensis, suggesting an intra‐riverine radiation. However, the Genale assemblage appears to have originated through a combination of allopatric and sympatric events. Some of the specialised forms within the Juba and Wabe–Shebelle drainage originated independently from the local generalised forms in three different river regions. Each of these cases could be considered as a small species flock composed of two or three sympatric forms. Our study shows that adaptive radiation in rivers can be enhanced by a combination of sympatric speciation and temporal geographic isolation, leading to local sympatric speciation followed by migration.

Chromosome sets of nine species of Barbus and a species of Varicorhinus from Ethiopia were investigated. All large Barbus (B. bynni, B. intermedius, B. ethiopicus) and V. beso are shown to have 2N =150, whereas all small Barbus (B. anema, B. kerstenii, B. paludinosus, and three unidentified (probably undescribed) species have 2N = 50. Based on the karyological data and on the published morphological data, an independent origin of small and large Barbus of Africa is suggested. The authors believe that large Barbus of Africa constitute a monophyletic group and that this group has no sister‐group relationships with any of the small Barbus of Africa.

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During the rainy season (September-November 1994), in tributaries of Lake Tana (Ethiopia), the Gumara, Gelda and Negashu, barbs of different morphotypes were ready to spawn, running and spent. The most numerous were barbs similar to Barbus intermedius common in many rivers of the Ethiopian Highlands. They were also found spawning in the Gumara and Gelda Rivers in the dry season (February 1996(February , 1997. There were no evident differences in time and place of spawning of barbs representing different morphotypes. Only zurki, bigmouth mini-eye and bighead were not found in the rivers. Spatial and temporal segregation may contribute to reproductive isolation of morphotypes spawning in rivers, but if reproductive isolation exists the main role must be played by differences in mating behaviour. Morphotypes of the Lake Tana barbs have different migration patterns. Barbs similar to the riverine B. intermedius may spend their lives in rivers or in the lake, moving to and fro. Barbs of acute, bigmouth big-eye and, perhaps, other morphotypes migrate downstream as alevins or fry, stay in the lake for several years and migrate back into rivers for spawning after reaching maturity. 1999 The Fisheries Society of the British Isles

In the Lake Tana barbs the PCA of skull measurements reveals "ontogenetic channels" within which individual ontogenetic trajectories are located. Some individuals seem to acquire states of characters specific for a certain morphotype quite late when they are 20-25 cm SL and 3-5 years old. Morphologically different sympatric forms of large barbs are also found in the Didessa (Blue Nile basin), Genale (Wabi Shebeli basin), and Gibe (Omo basin) rivers. In all the three rivers barbs resembling certain morphotypes of the Lake Tana barbs occur. Most probably, allopatric bigmouthed forms originated independently from sympatric forms which have no conspicuous specific traits and that their similarity results from homoplasies.