Genetic divergence among morphotypes of Lake Tana (Ethiopia) barbs

The genus Barbus is a paraphyletic assemblage that includes three different ploidy levels: diploid, tetraploid and hexaploid. 'B.' intermedins , which inhabits Lake Tana (Ethiopia), is a hexaploid taxon that forms a 'species flock' consisting of at least 14 morphotypes. Thirty-one presumptive allozymic loci were analysed in a large sample of 'B.' intermedius. No diagnostic loci were observed between the morphotypes, but substantial differences in allele frequencies were found. One morphotype, Acute, differs significantly from the others. The presence of this genetic differentiation within the lake strongly suggests that the morphotypes are diverging lineages. Because of some parallel characteristics with cichlid species flocks, we put forward the hypothesis that the morphotypes of barbs in Lake Tana represent a species flock that originated via sympatric speciation with more rapid morphological than molecular differentiation. If the differentiation and the reproductive isolation are real, it is probable that the morphotypes already are or will soon reach the status of species.     

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.     

Spawning migrations of the endemic Labeobarbus (Cyprinidae, Teleostei) species of Lake Tana, Ethiopia: status and threats

The reproductive biology of the only known intact species flock of large cyprinids, the 16 Labeobarbus species of Lake Tana (Ethiopia), has been extensively studied for the past two decades. Seven species of Labeobarbus are known to migrate >50 km upstream into tributary rivers for spawning during the rainy season (July to October), whereas eight other species are absent from these rivers and probably developed a new strategy of lacustrine spawning (macro-spatial segregation). One species (L. intermedius) probably spawns in the lake as well as in the rivers. Between the early 1990s and 2000s, the riverine spawners showed a decline of 75% in both biomass and number in both fishery independent surveys and in commercial catches. Reproductive migration makes fishes vulnerable to fisheries and other threats like habitat modifications. Lacustrine spawners are probably more resilient as they are not known to form spawning aggregations that can easily be exploited by fishermen. In addition, upstream rivers and catchments around Lake Tana are highly degraded by erosion and recently subjected to intensive habitat modification for irrigation and hydroelectric power generation. This article reviews results of field studies on the Labeobarbus spawning migration from Lake Tana to spawning rivers, giving emphasis on segregation and homing. It also summarizes existing and emerging threats which form potential causes for the decline of the migratory Labeobarbus species. Knowledge gaps on the reproductive biology are identified for further investigation.     

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.     

Seasonal variability of secondary production of cladocerans and rotifers,and their trophic role in Lake Tana,Ethiopia, a large,turbid, tropical highland lake

Daily and annual production rates of eight cladoceran and two rotifer species, and their seasonal variation and trophic role in the large, turbid, tropical Lake Tana, Ethiopia, were assessed in 2003–2005. Laboratory cultures were used to infer cladoceran development times, and secondary production was estimated using the growth increment summation and recruitment methods. Production for both taxa was highest in October–November, after the rainy season, and lowest in January–April during the dry season. Cladocerans and rotifers comprised 24% of the metazoan zooplankton biomass of 45.1 mg DW m^?3, but comprised 53% of its production. Daily production for cladocerans and rotifers, respectively, was 1.23 and 0.94 mg DW m^?3 d^?1, and annual production was 447.9 and 353.5 mg DW m^?3 y^?1. Energy transfer efficiency from producers to zooplankton was 1.3% and 4.4% from zooplankton to planktivores. Herbivores consumed 3.4% of primary production and planktivores 36% of zooplankton production. High biomass turnover rates of cladocerans and rotifers sustain planktivores and, after a month's delay, decomposed Microcystis provides their main food source during the pre- and post-rainy months in Lake Tana.     

金花小檗与滇小檗的名实问题

Léveillé在1911年发表了小檗科滇小檗(Berberis bodinieri Lévl.)(新拟),但在原始文献中未说明其模式标本收藏处,并且在之后的一个世纪里,世界小檗属主要分类学文献都未收载该种。2014年,作者在英国皇家爱丁堡植物园标本馆(E)找到了滇小檗模式标本。之后,通过比较滇小檗和古宗金花小檗的模式标本及其原始文献,发现在枝条具棱,叶片为匙形或倒卵状匙形,花序为总状花序或近伞形花序,果实近球形或扁球形、红色、顶端具明显宿存花柱等主要分类学特征上,滇小檗与古宗金花小檗基本一致。因此,将滇小檗归并入古宗金花小檗。     

Metacercariae of new trematode species of the genus Diplostomum (Trematoda,Diplostomidae) from fishes of Lake Tana,Ethiopia

Four new species of trematodes of the genus Diplostomum are described from metacercariae found in fishes from Lake Tana in Ethiopia: Diplostomum garrae sp. n. from the lens of Garra dembecha, D. longicollis sp. n. from the lens of Barbus humilis, D. montanum sp. n. from the lens of Barbus humilis, and D. tilapiae sp. n. from the vitreous body of Oreochromis niloticus. The metacercariae of the new species considerably differ in morphology and size from those of all known Holarctic species. It is the first African record of metacercariae of the genus Diplostomum in fish.     

Relationship between semen characteristics and body size in Barbus barbus L. (Teleostei: Cyprinidae) and effects of ions and osmolality on sperm motility

The objectives of the present study were to determine the relationships among length and weight of males, sperm volume, spermatozoa concentration, total number of spermatozoa, ionic contents and osmolality of seminal plasma in Barbus barbus. The effect of osmolality on sperm motility parameters after activation in NaCl, KCl, or sucrose solutions was also examined. There were significant correlations between spermatozoa concentration – length (R = + 0.7) and – weight (R = + 0.8) of males. No significant correlations were observed between the total number of spermatozoa, sperm volume, and length and weight of males. Seminal plasma osmolality was higher when the total number of spermatozoa (R = + 0.6) and sperm volume (R = + 0.6) were higher. Sperm motility and velocity was positively correlated with osmolality (R = + 0.5). The correlation between sperm motility and K⁺ was negative (R = 0.5), but positively correlated with Ca²⁺ (R = 0.8), Na⁺ (R = 0.8), and Cl⁻ (R = 0.8). There was a rapid decrease (P < 0.05) in sperm motility parameters after sperm activation. Just after sperm activation, beating waves propagated along the full length of flagella. At latter stages post sperm activation, the waves appeared only in proximal part of the flagellum. The highest spermatozoa velocity and percentage of motility were observed at 215–235 mOsmol kg^− 1 in NaCl, KCl or sucrose. The tip of the flagellum became curled into a loop shape which shortened the flagellum after activation of sperm in distilled water. B. barbus sperm is very similar to that of other cyprinids in terms of ionic contents and osmolality of the seminal plasma, mechanism of sperm activation and behavior and motility of sperm during swimming period.     

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.     

Bioaccumulation of heavy metals in two morphotypes of African large barb Labeobarbus intermedius (Osteichthyes: Cyprinidae) in Lake Hawassa,Ethiopia

Water and muscle tissue samples from two morphotypes of the African large barb Labeobarbus intermedius collected from three sites in Lake Hawassa in 2012–2013 were analysed for eight heavy metals, including Cd, Co, Cr, Cu, Mn, Ni, Pb and Zn. Five metals (Cr, Cu, Mn, Ni and Zn) were detected in fish muscle samples, whereas only Cr, Cu and Ni were detected in water samples. Of the five metals detected in the muscle samples, Cu and Zn were present in higher concentrations in the golden morphotype, whereas Cr, Mn and Ni were found in higher concentrations in the silver morphotype. Bioaccumulation factor (BAF) values indicated that Cr, Cu and Ni have a tendency to accumulate in fish muscle in amounts exceeding those in water. In both morphotypes the highest concentrations of Zn and Mn were detected at the Hospital site, whereas the concentrations of Cr, Cu and Ni were highest at the Tikur Wuha site. Chromium, Cu and Ni concentrations recorded in fish muscle at all sampling sites exceeded the safe limits recommended by FAO/WHO and UNESCO, suggesting that water and fish from Lake Hawassa are contaminated with heavy metals originating from factories, a hospital and agricultural activities in proximity to the lake.     

Trematode metacercariae from the cranial cavity of African catfish Clarias gariepinus (Burchell, 1822) from Lake Tana,Ethiopia

Metacercariae of two species of trematodes from the fam. Diplostomidae from the cranial cavity of the African catfish Clarias gariepinus are described. The metacercaria Tylodelphys grandis n. sp. is described for the first time. Data are presented on the infection rate of catfishes in Lake Tana.     

Seasonal variation in primary production of a large high altitude tropical lake (Lake Tana,Ethiopia): effects of nutrient availability and water transparency

Primary production rates, chlorophyll and phytoplankton biovolume were measured monthly from April 2003 to November 2004 in Lake Tana, a large tropical lake in the highlands of Ethiopia. The lake is characterised by low nutrient concentrations, and a low water transparency due to high silt load of the inflowing rivers during the rainy seasons (May–November) and daily resuspension of sediments in the inshore zone. The mean chlorophyll-a concentrations varied seasonally and ranged from 2.6 mg m⁻³ to 8.5 mg m⁻³ (mean: 4.5 mg m⁻³) in the offshore zone. Primary production was measured using the light–dark bottles technique. We incubated only at three depths, i.e. 0.6, 1.2 and 1.8 m. Therefore, we may have missed a substantial part of the depth production profile and probably also frequently missed P _max. Gross primary production in the openwater averaged 2.43 g O₂ m⁻² d⁻¹ and ranged between 0.03 g O₂ m⁻² d⁻¹ and 10.2 g O₂ m⁻² d⁻¹; production was significantly higher in the inshore zone. The highest production rates were observed in the post-rainy season (Oct–Nov), which coincided with a bloom of Microcystis and higher chlorophyll levels. This seasonal high production is probably caused by a relatively high nutrient availability in combination with favourable light conditions. The gross primary production rates of L. Tana are among the lowest compared with other tropical lakes. This will be partly the result of our underestimation of gross primary production by often missing P _max. Another cause is the oligotrophic nature of the lake in combination with its relatively low water transparency. The gross primary production per unit chlorophyll in the openwater zone was in the same range as in 30 other tropical lakes and reservoirs. The higher primary production in the inshore zone is probably the result of the daily water column mixing (Z _mix ≥ Z _t) in this area, enhancing nutrient recycling. A large proportion of the annual primary production is realised in one of the four seasons only. This productive post-rainy season is relatively short (2 months) and therefore efficiency of transfer of matter between the first and second trophic level of the Lake ecosystem will be poor.     

Aspects of reproductive biology of Labeo horie Heckel (Pisces: Cyprinidae) in Lake Chamo, Ethiopia

Labeo horie Heckel is increasingly becoming commercially important in Lake Chamo but its reproductive biology in the lake is not well enough to guide its management. Sex ratio, breeding season, length at first maturity and fecundity of L. horie were studied from 1197 fish samples collected between August 1998 and October 1999 in Lake Chamo, Ethiopia. The sex ratio was significantly different (χ ² = 12.12, P < 0.001). The peak‐breeding period was during the rainy months of March to May, during which time more than 90% of both sexes had ripe gonads. The size at maturity ( L _m50 ) of males was 52 cm while the L _m50 of females was 62 cm. The smallest ripe male in the catch was 46.7 cm and weighed 890 g while the smallest ripe female caught was 49.5 cm and weighed 1145 g. The weight of ripe ovaries ranged from 54.3 g to 991.8 g and contained between 68,400 and 1,320,400 eggs. Relative fecundity ranged between 60 and 290 eggs per g of body weight. The relationships between fecundity and total length and between fecundity and total weight were curvilinear, while the relationship between fecundity and ovary weight was linear. L. horie conformed to the general pattern of reproduction in a tropical environment where peak‐breeding activity occurred during the rainy season. In order to protect the spawning population, fishing pressure should be minimized during breeding time at the shallower littoral regions.     

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.     

Species composition and phytoplankton biomass in a tropical African lake (Lake Awassa,Ethiopia)

The species composition and phytoplankton biomass of Lake Awassa, Ethiopia were studied from September 1985 to July 1986 in relation to some limnological features of the lake. During the study period, three phases of thermal stratification were recognized: a period of unstable stratification and near-complete mixing was followed by a stable stratification period and another period of complete mixing. Complete mixing was associated with cooling of air temperature with an influx of cool rain and high rainfall. The underwater light penetration showed a similar pattern over the whole period with the highest in the red, and the lowest in the blue spectral region. Euphotic depth varied between 1.6 and 3.0 meters with the highest measurements corresponding to the stable stratification period. PO₄-P concentrations ranged between 23 and 45 µg l^–1 and NO₃-N concentrations varied between 7 and 14 µg l^–1 during the study period. Both nutrients showed increasing values associated with mixing periods and/or the rainy season.A total of 100 phytoplankton species were identified with 48% of the taxa represented by green algae, 30% by blue-green algae, 11% by diatoms, and the rest by chrysophytes, dinoflagellates, cryptomonads and euglenoids. The dominant phytoplankton species were Lyngbya nyassae, Botryococcus braunii and Microcystis species. Seasonal biomass variation was pronounced in the first two species but not in Mycrocystis. Phytoplankton biomass increased following the mixing period in December, and thermal destratification during May to July which was also a period with high rainfall and relatively high nutrient concentration. While the seasonal variation of the total phytoplankton community in Lake Awassa was relatively low (coefficient of variation < 20%), it was higher in some of the individual component species.     

Reproductive segregation among the Barbus intermedius complex of Lake Tana,Ethiopia. An example of intralacustrine speciation?

Reproductive segregation among the large barbs Barbus intermedius complex, Cyprinidae , of Lake Tana, Ethiopia, was investigated in the mouths and upstream reaches of rivers tributary to the lake, during the spawning seasons of 1993 and 1994. The percentage of running fish of four Barbus morphotypes [Acute (Ac), Bigmouth big-eye (Be), Intermedius (In), and Shorthead (Sh)] peaks with the highest water level of the lake. Large differences are apparent in the minimum fork length of running fish among morphotypes [c. 12 cm for male Sh and 28 cm for male Bigmouth small-eye (Se)]. Significant differences occur in temporal, but especially in spatial patterns of the relative abundance of running morphotypes, when the lake is compared with the four major river mouths and their upstream reaches. These different relative abundances are clearest among Ac, Be, In, Sh and Troutlike (Tr). Also, the absence from the rivers of some morphotypes that are usually abundant in the lake itself [Barbel (Ba), Dark (Da), Lip (Li), and Zurki (Zu)], and the abundant presence in the rivers of Tr, rare in the lake, strengthen the hypothesis that spatial and/or temporal spawning segregation is present among the morphotypes. There are indications that Bighead (Bh) and Zu spawn in the lake itself, while other morphotypes were only found running in or near rivers. Combined with the morphological, ecological, and genetic differences, these data suggest the existence of several Barbus species, forming a unique species flock, which probably evolved within the lake. The presence of several, instead of one Barbus species complicates management of sustainable fisheries and protection of the lake's biodiversity. Present results show the importance of the rivers for Barbus reproduction and stress the urgent need for careful management of fishing activities in spawning times and areas.