Phenetic relationships and probable ways of morphological diversification of the African large barbs (<Emphasis Type="Italic">Barbus intermedius</Emphasis> complex) in Lake Tana (Ethiopia)

29 quantitative and 18 qualitative skull characters were studied in the African large barbs (Barbus intermedius complex) from Lake Tana (Ethiopia). Based on results of principle components analysis phenetic relationships of 12 morphotypes were assessed. External features diagnostic of the morphotypes were found in individuals substantially different in the skull characters. The obtained data along with the data on morphological diversity of large barbs from Ethiopia beyond the Lake Tana basin suggest origination of the Lake Tana complex of forms from a complex of forms that existed in the upper reaches of the Blue Nile before the lake came into being, and the extant morphotypes of the Lake Tana barbs originated from three or four forms that composed the ancestral riverine local complex.     

Polymorphism of skull proportions in large African barbs <Emphasis Type="Italic">Barbus intermedius</Emphasis> sensu Banister (Cyprinidae) from Lakes Awasa and Langano (Rift Valley,Ethiopia)

After performing principal component analysis (PCA) of 13 characters (indices) of skulls of large African barbs (Barbus intermedius complex sensu Banister, 1973) from Lakes Awasa and Langano (Ethiopia) distributions of individuals in coordinates BL and PC1 were studied where BL is the basal skull length and PC1 is the first principal component. It is shown that in both the lakes there are groups of barbs whose individual ontogenetic trajectories are situated within separated ontogenetic channels. Separation of the channels is by far less distinct than that of the channels of generalized and specialized forms of barbs from Lake Tana and not correlated with differences in external traits. It is suggested that the revealed polymorphism is connected with food resource partitioning and might serve as a prerequisite of further divergence of sympatric forms of large African barbs.     

Assessment of the role of chemoreception in the mate choice in barbs of the Barbus intermedius complex from Lake Tana, Ethiopia

The role of chemoreception in mate choice was studied in barbs of the complex Barbus intermedius from Lake Tana. It was found that chemical communication may be used in reproductive interactions of the barbs, and that males are responsible for the choice. Mature males tend to prefer ripe females and avoid both immature females and mature males, or are indifferent to them. It was shown that selectivity in mate choice by certain barb morphotypes could reduce the probability of interbreeding between different morphotypes, but not provide for reproductive isolation between them. It was hypothesized that the mate groups are finally formed as a result of direct interactions between potential partners, mediated by tactile reception or specific behavioral patterns.     

Phenetic diversity of the large barbs (Barbus intermedius complex sensu Banister) from the coastal zone of the Bahar Dar Gulf (Lake Tana, Ethiopia)

Phenetic structure of a sample of large African barbs Barbus intermedius complex) taken in November 2010 from the inshore part of the Bahar Dar Gulf (Lake Tana) was studied. Principal components analysis was performed on 28 indices characterizing skull proportions and shape of bones. The individuals under study formed distinct clusters in the principal component space. Composition of these clusters was considered. Many morphotypes described in the 1990s were absent in the sample taken in 2010, and there were individuals which did not comply with the diagnoses of the previously described morphotypes. It was suggested that some of these fish represented phenetic groups which did not occur in the offshore part of the gulf, some were, most probably, hybrids of the known morphotypes or originated from homonomic crossings but did not had diagnostic characters of the parental morphotype.     

Changes of Skull Proportions in Ontogeny of Barbus canis from Lake Kinneret and of the Lake Tana large-mouthed barbs, Barbus intermedius Complex

Skull proportions of Barbus canis individuals drastically change during their life. Yearlings have skull proportions characteristic for barbs of the NEC phenetic group (sensu Mina et al. 2001 Environ. Biol. Fish. 61: 242–252), but with age become similar to those of the Lake Tana large-mouthed morphotypes which acquire their specific features at 3–7 years old while at earlier ages they have skull proportions of a NEC form identified as 'intermedius' sensu Nagelkerke et al. (1994 Environ. Biol. Fish. 39: 1–21). The large-mouthed Lake Tana barbs originated from an ancestral NEC form through peramorphosis, but in relation to B. canis they are paedomorphic in skull proportions. B. canis supposedly originated from a NEC form as a consequence of an adaptive radiation that took place in western Asia.     

Role of thyroid hormone in the ontogeny and morphological diversification of <Emphasis Type="Italic">Barbus intermedius sensu</Emphasis> Banister, 1973 of Lake Tana in Ethiopia

The role of thyroid hormone in the ontogeny of the Large African barb Barbus intermedius sensu Banister, 1973 (Teleostei; Cyprinidae)—the presumed ancestral species for the species flock of the large African Barbs of Lake Tana (Ethiopia)—has been studied. It has been shown that the thyroid hormone influences the timing and rate of many morphogenetic processes and a change in its level causes heterochronies, affecting the definitive morphology of fish. Thus, it can be assumed that even a slight change in activity of the thyroid axis during ontogeny can be the cause of wide morphological variability in B. intermedius and may underlie explosive diversification of the species flock of Large African Barbs of Lake Tana.     

Individual ontogenetic trajectories and ontogenetic channels of three forms of large African barbs (<Emphasis Type="Italic">Barbus intermedius</Emphasis> complex). Analysis of experimental data

Changes of external morphological characters in three forms (morphotypes) of African barbs from Lake Tana (Ethiopia) belonging to Barbus intermedius complex were traced using electronic tags. It was shown that within age interval from two to four years individual ontogenetic allometric curves often cannot be described with a single power function equation. Individual ontogenetic trajectories in the principal components space within this interval strongly differ in shape meandering within ontogenetic channels outlined on the basis of cross-sectional data. Morphological differences between siblings are found that do not allow to classify some of them with the morphotype of their parents.     

Mitochondrial DNA phylogeography of the Labeobarbus intermedius complex (Pisces,Cyprinidae) from Ethiopia

Mitochondrial DNA phylogeography of populations of the Labeobarbus intermedius complex (hexaploid barb) was investigated using 88 complete and 71 partial cytochrome b (cytb) sequences originating from 21 localities in five major drainages in Ethiopia and two localities in northern Kenya. The samples included 14 of the 15 Labeobarbus species described from Lake Tana. Discrete phylogeographic analyses of 159 cytb sequences employing Bayesian Markov Chain Monte Carlo (MCMC) simulations using Bayesian evolutionary analysis by sampling trees (BEAST) supported the monophyly of the L. intermedius complex, including the Lake Tana species. This analysis, in combination with statistical parsimony analysis, identified two mitochondrial DNA lineages within the complex. Divergence dating employing coalescent simulations suggested that the geographic split in the L. intermedius complex that led to the formation of these lineages occurred during the Pleistocene (c. 0·5 M b.p. ), consistent with the timing of volcano‐tectonic events postulated to have shaped the current landscape of East Africa.     

Morphological diversity and age dependent transformations in progeny of the large barbs (Barbus intermedius complex sensu Banister) of several morphotypes from Lake Tana (Ethiopia). Results of a long-time experiment

Results of a long-time (2005?C2011) experiment in crossing and rearing the large African barbs from Lake Tana (Ethiopia) are presented. One may conclude that progeny from homonomic crossings of large-mouthed barbs acquire diagnostic traits of their parental morphotype at different age. Some males originated from different crossings reached sexual maturity before acquiring diagnostic traits of a certain morphotype. Siblings from both homonomic and heteronomic crossings living in the same conditions considerably differed both in external and in skull characters. Differences both in external and skull characters between some siblings originated from a homonomic crossing that lived in a pond and those that lived in aquaria were equivalent to the differences between moiphotypes.     

Scale structure variability of “forms flock” of Barbus (=Labeobarbus) intermedius and Varicorhinus jubae from the Genale River (Eastern Ethiopia)

Scales sculpture of 6 morphotypes of large African barbs Barbus (=Labeobarbus) intermedius and Varicorhinus jubae from the Genale River were described using light and scanning electron microscopy (SEM). It was found that in spite of several trends in difference macrostructural scale parameters (shape and lateral line scales number), cannot be used for identification of most forms of barbs and V. jubae of the Genale River. SEM analysis of scale microstructure of large barbs, V. jubae and V. beso has showed that the relief of the caudal field of the scales is not uniform and four main zones may be distinguished. The differences between forms and species in the relief details are weakly expressed however the studied species may be arranged in succession according to the development of specific; characters in transitional Zone II: B. intermedius (5 forms: GF, LMP, SH, LMS, LIP) > ??smallmouth scraper?? form??probable hybrid B. intermedius × V. jubae (SMS) > V. jubae > V. beso. By the development of granulation zone III the following series may be suggested: B. intermedius (5 forms: GF, LMP, SH, LMS, LIP), V. jubae > ??smallmouth scraper?? formprobable hybrid B. intermedius × V. jubae (SMS) > V. beso. By the degree of founded ??trabecular?? structure development the following series may be arranged according to the character reduction: B. intermedius (GF, LMP, SH, LMS, LIP) > ??smallmouth scraper?? form-probable hybrid B. intermedius × V. jubae (SMS) > V. jubae > V. beso. Thus in accordance with scale microstructure Varicorhinus jubae closer to Barbus (=Labeobarbus) intermedius than to V. beso.     

Observations on reproduction of the Lake Tana barbs

During the rainy season (September–November 1994), in tributaries of Lake Tana (Ethiopia), the Gumara, Gelda and Negashu, barbs of diVerent 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, 1997). There were no evident diVerences in time and place of spawning of barbs representing diVerent 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 diVerences in mating behaviour. Morphotypes of the Lake Tana barbs have diVerent 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.     

Diet segregation between two sympatric 'small'Barbus spp: an experimental study of mechanisms

Gut contents of two co‐occurring species of 'small' diploid barbs (<10  L _F cm) in Lake Tana revealed that zooplankton is the major diet component for B. tanapelagius (75% based on volume), but less prominent in B. humilis (40%). Functional response experiments in the laboratory were conducted to elucidate the mechanisms causing this difference. The type of functional response by the two 'small' barbs under different microcrustacean zooplankton densities (10, 20, 40, 60 and 80 ind.l⁻¹) was examined. The functional response of B. tanapelagius to increasing prey densities corroborates with Holling Type II model, whereas B. humilis exhibits a Type III functional response. Predation rate is higher for B. tanapelagius at low zooplankton density (<40 ind.l⁻¹) and equals the level of B. humilis at higher densities (>40 ind.l⁻¹). This suggests that at lower zooplankton densities B. humilis is a less efficient forager on zooplankton prey items than B. tanapelagius . In Lake Tana average zooplankton density is relatively low (<35 ind.l⁻¹). Under these food conditions, B. humilis is forced to feed on other food items (e.g. benthic invertebrates), whereas B. tanapelagius primarily feeds on zooplankton. The feeding potentials of the two 'small' barbs, as deduced from their morphology explain their different performances and their segregation in space and food resources.     

Major histocompatibility genes in the Lake Tana African large barb species flock: evidence for complete partitioning of class II B, but not class I, genes among different species

The 16 African large barb fish species of Lake Tana inhabit different ecological niches, exploit different food webs and have different temporal and spatial spawning patterns within the lake. This unique fish species flock is thought to be the result of adaptive radiation within the past 5 million years. Previous analyses of major histocompatibility class II B exon 2 sequences in four Lake Tana African large barb species revealed that these sequences are indeed under selection. No sharing of class II B alleles was observed among the four Lake Tana African large barb species. In this study we analysed the class II B exon 2 sequences of seven additional Lake Tana African large barb species and African large barbs from the Blue Nile and its tributaries. In addition, the presence and variability of major histocompatibility complex class I UA exon 3 sequences in six Lake Tana and Blue Nile African large barb species was analysed. Phylogenetic lineages are maintained by purifying or neutral selection on non-peptide binding regions. Class II B intron 1 and exon 2 sequences were not shared among the different Lake Tana African large barb species or with the riverine barb species. In contrast, identical class I UA exon 3 sequences were found both in the lacustrine and riverine barb species. Our analyses demonstrate complete partitioning of class II B alleles among Lake Tana African large barb species. In contrast, class I alleles remain for the large part shared among species. These different modes of evolution probably reflect the unlinked nature of major histocompatibility genes in teleost fishes.Electronic Supplementary Material Supplementary material is available for this article at .An erratum to this article can be found at     

Spawning of large <Emphasis Type="Italic">Barbus</Emphasis> (<Emphasis Type="Italic">Barbus intermedins</Emphasis> Complex) in a small river of the Lake Tana basin (Ethiopia) and relationships of some putative species

Direct observations on three forms (morph types) of spawning barbs, considered by some authors as different species, were carried out in September–October 2005 at the Gumara River and its small tributary, the Ducalit, in the Lake Tana basin. The spawning barbs were sampled using cast nets. Barbs of different morphotypes composed common spawning aggregations and were caught together. It is likely that troutlike females mate extensively with intermedius males and, probably, with bigmouth small-eye males. Experiments in artificial fertilization and rearing showed that, during the embryonic period, there was no increase in mortality of progeny from crosses of barbs of different morphotypes, in comparison with homonomic crosses, thus suggesting an absence of postzygotic reproductive isolation between the morphotypes. At the same time, mortality of progeny from crossing barbs with Varicorhinus beso Rüppell, 1836 was very high during the earliest stages of development, suggesting strong, even if incomplete, reproductive isolation ensured by postzygotic mechanisms. Published in Voprosy Ikhtiologii, 2007, Vol. 47, No. 5, pp. 676–683. The article was translated by the authors.     

Toothless predators: feeding performance and techniques among the piscivores within Lake Tana's endemic barbus species flock (Pisces:Cyprinidae)

With more than 2000 fish species the Cyprinidae is the largest family of vertebrates. Lake Tana, a large lake (3050 km²) situated in the NW‐ highlands of Ethiopia, harbours, as far as we know the only remaining intact species flock of large (max. 100 cm FL) cyprinid fishes (15 Barbus spp.). One of the most intriguing aspects of this endemic Barbus species flock is the large number of piscivores (8). Cyprinid fishes seem not well designed for piscivory, they lack teeth in the oral jaw, have a small slit‐shaped pharyngeal cavity and all lack a stomach with low pH for digesting large prey. Many barbs are benthivorous species, like the ancestral barb in Lake Tana's isolated system. Why then is piscivory, which is rare among cyprinids, so common in Lake Tana Barbus? The aim of present study was to compare the performance and techniques of these piscivorous Barbus with known piscivores from other fish families. We studied prey handling times over prey size, prey capture using high‐speed movies, and assessed the effect of prey size on performance and prey selection in the field. Performances were explained by functional morphology of their feeding system. Overall, Lake Tana's piscivorous Barbus perform relatively 'poor', compared to piscivores from other fish families. For example, Lake Tana's piscivores are only able to handle prey fish smaller than 16% of their own body length. However, Lake Tana lacks potential piscivorous competitors, rendering the piscivorous Barbus by far the 'best' and apparently highly successful. They have adapted to all available macro‐habitats (littoral, offshore pelagic and offshore benthic), using different techniques (ambush, pursuit and cruising), a unique scenario for barbs.     

Development of the cranium in the large African hexaploid barb Labeobarbus (=Barbus) intermedius (Cyprinidae; Teleostei)

Developmental information on the structure and composition of the cartilaginous and bony skull in the large African barb Labeobarbus (=Barbus) intermedius (Teleostei; Cyprinidae) is provided. Sequences of cartilages and bones appearance from a large series of cleared and Alizarin red- and Alcian blue-stained laboratory-reared specimens ranging from prehatching larvae to juvenile stages are described. Comparisons of cranial development are made among cyprinids: L. intermedius, Danio rerio, Barbus barbus, and Cyprinus carpio.     

Spatial and Temporal Variation of Cestode Infection and its Effects on Two Small Barbs (Barbus humilis and B. tanapelagius) in Lake Tana, Ethiopia

Pseudophyllidean cestodes as Ligula have a complex life cycle with cyclopoid copepods as first intermediate host, zooplanktivorous fish as second, and piscivorous birds as final host. We studied the effects of diet, season and habitat occupation on the prevalence of plerocercoid larvae of the tapeworm Ligula intestinalis in two closely related small barbs and the effects of the parasites on the barbs life histories in Lake Tana (Ethiopia) during 1 year. In all affected barbs L. intestinalis caused retardation in gonad development, maturation at reduced size and lower absolute fecundity. Infection rate, averaged over all habitats was significantly higher in B. tanapelagius (10%) than in B. humilis (6%). Below a threshold of 48 mm the infection rate was zero for both barbs, this coincided with a very low proportion of copepods in their diets, increasing up to 90 and 55%, respectively, for their largest size class (81–90 mm). The relatively high infection rate in B. tanapelagius is explained by its obligatory zooplanktivorous feeding behaviour, ingesting a relatively high proportion of infected cyclopoid copepods. This is in contrast with B. humilis, which is a polyphagous species, feeding both on zooplankton and benthic invertebrates. Significant seasonal effects in infection rates were observed. In both barb species infection rates were lower during the breeding season. Only for B. tanapelagius a significant negative correlation was observed between rain fall and infection rate, probably caused by an increased turbidity that decreases feeding efficiency on zooplankton. Habitat type had also a significant effect on infection rate. Barbus humilis showed a much higher infection rate in shallow clear water (10%) than in shallow turbid water (3%), whereas B. tanapelagius showed much higher infection rates in the shallow sublittoral (13%) than in the deeper pelagic (7%). Most likely, birds predate more efficiently on barbs in shallow clear waters than in shallow turbid and deep waters.     

Lake Tana large barbs: phenetics, growth and diversification

Among the Lake Tana large barbs of about 10 cm SL only representatives of the' acute'morphotype can be distinguished, in the size range 10–20 cm SL' bigmouth big eye' can be identified also. As for the rest, very few individuals can be confidently affiliated with a particular morphotype most of them looking like' intermedius'. Even within the range of 20–30 cm SL some individuals are still difficult to identify. Principal component analysis of cranial characters revealed discrete groups of morphotypes. Differences in both external and cranial characters of the morphotypes result from divergence which is most pronounced when fish are 4–5 years old and 20–25 cm SL. This divergence cannot be related exclusively with differences in the growth rates of individuals representing different morphotypes. Differences in food composition between the morphotypes probably increase in parallel with their morphological divergence. Differences between the morphotypes in the lateral line (11) and the gill rakers (Sb) counts were revealed using ANOVA. Comparison of the Lake Tana Barbus complex of forms with that previously known from Lake Lanao (Philippines) suggests that in both lakes the different forms arose sympatrically but that sympatric speciation in Lake Lanao has advanced further than in Lake Tana.     

Speciation of endemic Lake Tana barbs (Cyprinidae, Ethiopia) driven by trophic resource partitioning; a molecular and ecomorphological approach

A unique species flock of large barbs (Barbus spp.) from Lake Tana is presented, from the level of fish stocks to molecules. Evidence is given for the species status of 14 morphotypes of large barbs. They distinctly differ in: (1) head and body morphometrics, (2) food preferences, (3) distribution patterns, (4) maximal body size, (5) spawning area and period, and (6) molecular genetic characters. Most types show early morphological divergence at small size. Major Histocompatibility Complex (MHC) genes, encoding cell membrane proteins involved in defence against pathogens, were found to be diagnostic for the species' genetic identity. A strong selective pressure on particular amino acid positions in the MHC protein sequence most probably arose in response to different pathogen loads from the newly invaded ecological niches after formation of Lake Tana, ca. two million years ago.Arguments for a sympatric origin of this species flock are discussed. An evolutionary scenario suggests a riverine ancestral Barbus intermedius invading Lake Tana after its formation by volcanic blocking of the Blue Nile river and its isolation by waterfalls. Specialisation for particular food types and disruptive selection on many feeding structures are hypothesized as the evolutionary drive in speciation. The causal relationship between the diversity in feeding structures and food types is explained from experiments and models. As an example, the potential food niche of three barb species is predicted from parameter values measured from a large set of feeding structures and tested against the actual gut contents. The co-occurrence of eight piscivorous barb species is unique for cyprinid fish, which lack oral jaw teeth and a stomach. The significance of this aquatic ecosystem as a multidisciplinary evolutionary laboratory and the need for a wise balance between exploitation and conservation is stressed.