Morphometric and allozyme variation in the African catfishes Clarias gariepinus and C. anguillaris

This study investigated morphological characters and electrophoretic polymorphism at 25 protein loci in nine wild populations of the African clariid catfish Clarias gariepinus and seven wild populations of C. anguillaris. Two other clariid species, Clarias albopunctatus and Heterobranchus longifilis , were used as outgroups in the allozyme study. Morphometric and allozyme data are congruent for the Nilo-Sudanian populations of C. gariepinus and C. anguillaris. Both approaches also distinguished two groups amongst the C. gariepinus populations, one containing Nilo-Sudanian populations and the other including Lake Victoria and southern African populations. However, allozyme data suggest that C. gariepinus is not a monophyletic group and show that C. albopunctatus is more divergent from C. gariepinus and C. anguillaris than it is from H. longifilis , stressing the need for a revision of clariid systematics. The variation observed in C. gariepinus is discussed in terms of palaeogeographical events and its use in aquaculture.     

Immunological study of muscle parvalbumin isotypes in three African catfish during development

Eleven parvalbumin isotypes expressed during the development of clariids Heterobranchus longifilis and Clarias gariepinus and claroteid Chrysichthys auratus were purified and electrophoresed on sodium-dodecyl-sulfate polyacrylamide gels. Immunochemical cross-reactions among these proteins were investigated by immunoblotting, using purified antibodies raised against three isotypes chosen at different stages of fish development. Antibodies raised against H. longifilis PA I (larval-juvenile isotype) and against C. gariepinus PA IIIa (juvenile-adult isotype) cross-reacted to a rather similar extent despite a weaker cross-reaction of anti-PA IIIa with larval-juvenile isotypes. On the other hand, antibodies raised against H. longifilis PA IV (an exclusively adult isotype) recognized markedly only H. longifilis PA IV and C. gariepinus PA IIIb. These two adult isotypes most likely belong to the alpha lineage, and all the others to the beta lineage. These results show that parvalbumin isotypes synthesized at different stages of fish growth differ structurally, and that the most marked difference is between larval-juvenile and adult clariid isotypes.     

A karyological analysis of the artificial hybridization between Clarias gariepinus (Burchell, 1822) and Heterobranchus longifilis Valenciennes, 1840 (Pisces; Clariidae)

A karyological analysis of an artificial hybridization (reciprocal crosses) between two African clariid catfish, Clarias gariepinus (Burchell, 1822) and Heterobranchus longifilis Valenciennes, 1840, was performed. C. gariepinus has a standard karyotype of 2 n = 56, while H. longifilis has 2 n = 52. The hybrids revealed an intermediate karyotype (2 n = 54), and it appears as if they have totalized the haploid chromosome number of both parental species, excluding gynogenesis or androgenesis. The hybrid karyotype is considered as aneuploid, although the hybrids proved to be fertile. No variation was found in the hybrids karyotypes.     

Muscle parvalbumin isoforms of Clarias gariepinus, Heterobranchus longifilis and Chrysichthys auratus: isolation, characterization and expression during development

The white-muscle parvalbumin isoforms of Clarias gariepinus, Heterobranchus longifilis and Chrysichthys auratus were purified and their physicochemical parameters determined. The three catfish isoforms are distinct but those of C. gariepinus and H. longifilis are more similar. In the course of development, the successive appearance of larval and adult parvalbumins was observed. Larval isoforms (PA I, PA IIa, PA IIb) displayed a lower isoelectric point (pI) and molecular mass than adult ones (PA IIc, PA IIIa, PA IIIb, PA III, PA IV). The PA IIa isoform appeared as an omnipresent typical larval isoform. PA IIb appeared mostly larval, being insignificant in adult specimens; its physicochemical features were the same in the three catfish species. In Chrysichthys auratus , there were three PA II isoforms, one appearing as an adult isoform (PA IIc). The fact that the two types of parvalbumin isoforms appear at different times should reflect specific physiological needs (mobility, feeding) of different developmental stages.     

Morphometric and genetic characterization of sympatric populations of Clarias gariepinus and C. anguillaris from Senegal

A sample of African Clarias catfishes from the Senegal River was studied using morphometry, allozyme variation, microsatellites and RFLPs of mitochondrial DNA. They all confirmed the presence of two species , C. gariepinus and C. anguillaris . The two species were closely related genetically and no diagnostic loci were found in allozymes and microsatellites studies. Two of the 11 haplotypes of mtDNA observed were shared by both species. Three of the four assays (morphometry, allozymes and microsatellites) allowed a precise characterization of both. One specimen occupied an intermediate position in the analysis of the data; it was considered an F_l hybrid whose possible origin is discussed.     

A comparative study on morphology, growth rate and reproduction of Clarias gariepinus (Burchell, 1822), Heterobranchus longifilis Valenciennes, 1840, and their reciprocal hybrids (Pisces, Clariidae)

A detailed study on reproduction indicated that female C. gariepinus mature earlier (5–7 months) than female H. longifilis (12–14 months). In the reciprocal hybrids, female first sexual maturity was attained at 20–21 months. Both reciprocal hybrids and parental species displayed an equilibrated sex ratio. Macroscopical and microscopical observations revealed numerous abnormalities in gonadal development of the hybrids. GSI and fecundity in female hybrids were considerably lower than those found in the parental species. Moreover, intra-ovarian tumors occurred in 20% of the hybrids. In hybrid males, GSIs were generally higher than those found in the parental species, but the spermatozoa concentration in the semen was about 100 times less. Despite these abnormalities, viable larvae, resulting from F2 and backcross fertilizations, were obtained.     

Redescription of Proteocephalus glanduligerus (Cestoda: Proteocephalidea), a parasite of clariid catfishes in Africa with a unique glandular apical organ

The tapeworm Proteocephalus glanduligerus (Janicki, 1928) Fuhrmann, 1933 (Cestoda: Proteocephalidea), a parasite of clariid catfishes (Clarias spp.) in Africa, is redescribed on the basis of an evaluation of voucher material and newly collected specimens from Clarias gariepinus and C. cf. anguillaris from the Sudan, Ethiopia, and South Africa. The most typical characteristic of P. glanduligerus (previously misspelled as P. glanduligera or P. glanduliger) is the presence of an extremely large glandular apical organ, the size of which is 1.6-3.5 times larger than that of the suckers. In addition, other morphological characteristics unreported in previous accounts are provided, such as an unusual position of osmoregulatory canals (situated close to each other, with the dorsal canal latero-ventral to testes), the presence of a vaginal sphincter, several (usually 3-5) uterine pores, and eggs with paired lateral auricular swellings (extensions) of the outer envelope.     

Influence of temperature on egg hatching, growth and survival of larvae of Heterobranchus longifilis Val. 1840 (Teleostei: Clariidae)

Eggs of Heterobranchus longifilis Val. 1840 were artificially fertilized and incubated at a range of temperatures (20, 23, 25, 27, 29 and 32°C). The time from fertilization to hatching decreased with increasing temperature. No eggs survived to hatch at 20 and 32°C incubation temperatures, while at 23 and 29°C hatching was only minimal. Optimum hatching was obtained at 25 and 27°C, which corresponds to the ambient temperature range during the breeding season. Larvae of H. longifilis were reared for 11 days post-hatching at 20, 25, 27, 29 and 32°C. Growth increased with temperature (P < 0.05), whereas survival depicted an inverse relationship. Growth was minimal at 20°C and larvae rarely survived to the end of the experiment. Optimum temperature for the primary nursing of H. longifilis larvae was within the 25–27°C temperature range.     

Effects of vitamin C supplementation on the growth of Heterobranchus longifilis fingerlings

A 12-week growth experiment was conducted to establish the necessity of vitamin C in the nutrition of Heterobranchus longifilis fingerlings. Vitamin C was supplemented at levels of 0 and 50 mg x kg(-1) to a basal diet (42.5% CP), which was fed to triplicate groups of H. longifilis fingerlings. Fish receiving the vitamin C-supplemented diet had significantly improved weight gain (20.7 vs. 16.7 g per fish), feed efficiency ratio (1.03 vs. 1.42), specific growth rate (3.00 vs. 2.74%), protein efficiency ratio (2.26 vs. 1.64), and survival rate (90% vs. 50%). There was a significant decrease in haematocrit and haemoglobin levels in the blood of fish fed no supplemental vitamin C. Furthermore, this group exhibited retarded growth and pathological changes such as vertebral curvature, condensation associated with fragility of the spinal bones. Supplementation of 50 mg vitamin C per kg diet was adequate to prevent the occurrence of vitamin C deficiency in H. longifilis and it was concluded that vitamin C is essential in the nutrition of these fishes.     

The growth and feeding habits of Clarias anguillaris during their first season in the floodplain pools of the Sokoto-Rima river basin, Nigeria

The growth and feeding habits of juvenile Clarias anguillaris (Linné 1762) during their 5-month residence in the floodplain pools of the Sokoto-Rima river basin are described. A mean growth increment of 202 mm and 69.7 g attained in the October to March season compares favourably with first-year growth recorded for other Clarias species. The diet of C. anguillaris is relatively unspecialized and includes a wide variety of items ranging from algae, higher plant material and detritus through larval dipterans, rotifers and copepods, to fish. There is variation in the diet according to month of capture and capture location, and a reduction in diet diversity with fish size. The findings are discussed in the context of work carried out on other Clarias species. It is proposed that C. anguillaris is suitable for locally operated fish culture programmes.     

African sharptooth catfish Clarias gariepinus does not detoxify ammonia to urea or amino acids but actively excretes ammonia during exposure to environmental ammonia

The African sharptooth catfish Clarias gariepinus lives in freshwater, is an obligatory air breather, and exhibits high tolerance of environmental ammonia. This study aimed at elucidating the strategies adopted by C. gariepinus to defend against ammonia toxicity during ammonia exposure. No carbamoyl phosphate synthetase (CPS) I or III activities were detected in the liver or muscle of the adult C. gariepinus. In addition, activities of other ornithine-urea cycle (OUC) enzymes, especially ornithine transcarbamylase, were low in the liver, indicating that adult C. gariepinus does not have a "functional" hepatic OUC. After being exposed to 50 or 100 mM NH4Cl for 5 d, there was no induction of hepatic OUC enzymes and no accumulation of urea in tissues of the experimental animals. In addition, the rate of urea excretion remained low and unchanged. Hence, ammonia exposure did not induce ureogenesis or ureotely in C. gariepinus as suggested elsewhere for another obligatory air-breathing catfish of the same genus, Clarias batrachus, from India. Surprisingly, the local C. batrachus did not possess any detectable CPS I or III activities in the liver or muscle as had been reported for the Indian counterpart. There were no changes in levels of alanine in the muscle, liver, and plasma of C. gariepinus exposed to 50 or 100 mM NH4Cl for 5 d; neither were there any changes in the glutamine levels in these tissues. Yet even after being exposed to 100 mM NH4Cl for 5 d, there was no significant increase in the level of ammonia in the muscle, which constitutes the bulk of the specimen. In addition, the level of ammonia accumulated in the plasma was relatively low compared to other tropical air-breathing fishes. More importantly, for all NH4Cl concentrations tested (10, 50, or 100 mM), the plasma ammonia level was maintained relatively constant (2.2-2.4 mM). These results suggest that C. gariepinus was able to excrete endogenous ammonia and infiltrated exogenous ammonia against a very steep ammonia gradient. When exposed to freshwater (pH 7.0) with or without 10 mM NH4Cl, C. gariepinus was able to excrete ammonia continuously to the external medium for at least 72 h. This was achieved while the plasma NH4+ and NH3 concentrations were significantly lower than those of the external medium. Diffusion trapping of NH3 through boundary layer acidification can be eliminated as the pH of the external medium became more alkaline instead. These results represent the first report on a freshwater fish (C. gariepinus) adopting active excretion of ammonia (probably NH4+) as a major strategy to defend against ammonia toxicity when exposed to environmental ammonia.     

The African sharptooth catfish Clarias gariepinus can tolerate high levels of ammonia in its tissues and organs during four days of aerial exposure

The African sharptooth catfish Clarias gariepinus lives in freshwater, is an obligatory air breather, and can survive on land during drought. The objective of this study was to elucidate how C. gariepinus defends against ammonia toxicity when exposed to terrestrial conditions. During 4 d of aerial exposure, there was no accumulation of urea in its tissues, and the rate of urea excretion remained low. Thus, exposure to terrestrial conditions for 4 d did not induce ureogenesis or ureotely in C. gariepinus. Volatilization of NH(3) was not involved in excreting ammonia during aerial exposure. In addition, there were no changes in levels of alanine in the muscle, liver, and plasma of C. gariepinus; nor were there any changes in the glutamine levels in these tissues. However, there were extraordinarily high levels of ammonia in the muscle (14 micromol g(-1)), liver (18 micromol g(-1)), and brain (11 micromol g(-1)) of fish exposed to terrestrial conditions for 4 d. This is the first report on a fish adopting high tolerance of ammonia in cells and tissues as the single major strategy to defend against ammonia toxicity during aerial exposure. At present, it is uncertain how C. gariepinus tolerates such high levels of ammonia, especially in its brain, but it can be concluded that, contrary to previous reports on two air-breathing catfishes (Clarias batrachus and Heteropneustes fossilis) from India, C. gariepinus does not detoxify ammonia to urea or free amino acids on land.     

Temperature-induced changes of early development and yolk utilization in the African catfish Clarias gariepinus

Artificially fertilized eggs and yolk-sac larvae of a freshwater tropical/subtropical fish Clarias gariepinus receiving no external food were incubated at 22, 25 and 28° C until full yolk resorption. Developmental time, size and matter composition (CHNS-O Analyzer and ashing) were assessed at egg fertilization, hatching and yolk resorption; respiration was measured every 4–5 h. The course of acceleration of C. gariepinus embryonic developmental rate with temperature ( Q_{10 dev}) was compared over the temperature range to those of Cyprinus carpio and Oncorhynchus mykiss ; they differed greatly, but were similar when compared on the basis of effective temperatures specific to each fish. Specific growth rates for energy (88, 150 and 183% per day at 22,25 and 28° C, respectively) as well as the conversion efficiencies of egg energy (64, 71 and 68%, respectively) and protein (71, 78 and 76%, respectively) in C. gariepinus larval tissues were higher than those known for the endogenous feeding period of coldwater and temperate fish species. In C. gariepinus at the end of yolk resorption, the carbon percentage and caloric values of dry weight, size (in terms of dry matter, minerals, protein and energy per larva) and transformation efficiencies were lowest at 22° C, highest at 25° C and had slightly decreased at 28° C. A tentative mechanism which leads to the positive or negative response of body size to temperature over the viable temperature range is defined.     

The Bathyclarias-Clarias species flock. A new model to understand rapid speciation in African great lakes

Phylogenetic relationships between seven species of the catfish species flock from Lake Malawi (genus Bathyclarias) and other Clariid catfish have been investigated using cytochrome b partial sequences. Here we demonstrate that this species flock originated from a widespread, generalist species, Clarias gariepinus, still occurring in the lake. Bathyclarias species and their ancestor C. gariepinus form a simple model that can be used to understand the mechanisms of adaptation and rapid speciation in African Great lakes.     

Bite performance in clariid fishes with hypertrophied jaw adductors as deduced by bite modeling

Within clariid fishes several cranial morphologies can be discerned. Especially within anguilliform representatives an increase in the degree of hypertrophy of the jaw adductors occurs. The hypertrophy of the jaw adductors and skeletal modifications in the cranial elements have been linked to increased bite force. The functional significance of this supposed increase in bite force remains obscure. In this study, biomechanical modeling of the cranial apparatus in four clariid representatives showing a gradual increase in the hypertrophy of the jaw adductors (Clarias gariepinus, Clariallabes melas, Channallabes apus, and Gymnallabes typus) is used to investigate whether bite force actually increased. Static bite modeling shows that the apparent hypertrophy results in an increase in bite force. For a given head size, the largest bite forces are predicted for C. apus, the lowest ones for C. gariepinus, and intermediate values are calculated for the other species. In addition, also in absolute measures differences in bite force remain, with C. apus biting distinctly harder than C. gariepinus despite its smaller head size. This indicates that the hypertrophy of the jaw adductors is more than just a correlated response to the decrease in absolute head size. Further studies investigating the ecological relevance of this performance difference are needed.     

The ontogeny of the chondrocranium in Clarias gariepinus: trends in siluroids

The ontogeny of the chondrocranium of 31 different stages of the African catfish Clarias gariepinus (Siluroidei: Clariidae) was studied, both from cleared and stained, and sectioned material. The fish ranged from 4.1 (1 day post-hatching) to 127.0 mm SL (100 days post-hatching). The chondrocranium of C. gariepinus seemed to correspond to the general adaptive trends in siluroids, especially in relation to the reduction of eye size and the dorso-ventral flattening of the skull. The platybasic neurocranium involved several modifications related to the trabecular bars, the hypophyseal fenestra, the ethmoid region and even the olfactory nerves. Certain reductions were present, which have been observed in all siluroids (e.g. absence of the pila lateralis, the commissura lateralis, the myodomes) or are part of a variable trend within siluroids (e.g. reduction of the taenia marginalis anterior and the tectum synoticum). Compared with some other siluroid species, the neurocranium of C. gariepinus is well developed, for example in the otic region. The same was observed in the splanchnocranium where some general siluroid trends persist (e.g. isolation of palatine from pterygoquadrate, presence of 'hyo-symplectic-pterygoquadrate' plate). Some trends, as observed in other siluroids, were present also (e.g. interhyal continuous with suspensorium and ceratohyal, Meckel's cartilage initially continuous with the suspensorium). The branchial basket is well developed as all expected elements are present (basibranchials I-IV, hypobranchials I-IV, ceratobranchials I-V, epibranchials I-IV). Based on the observed ontogeny of C. gariepinus and data from the literature, a hypothesis was formulated which indicated the presence of a general reductional trend within siluroids. In C. gariepinus , all four (I-IV) infrapharyngobranchials develop, although the anterior two are much reduced and fused with each other.     

Ultrastructure of spermiogenesis and spermatozoa of Gymnotus cf. anguillaris and Brachyhypopomus cf. pinnicaudatus (Teleostei: Gymnotiformes)

The ultrastructure of spermiogenic stages and spermatozoa of representatives of two gymnotiform families, Gymnotus cf. anguillaris (Gymnotidae) and Brachyhypopomus cf. pinnicaudatus (Hypopomidae) were studied. Spermiogenesis of both species is characterized by lateral development of the flagellum and formation of a nuclear fossa. Some differences were found between these species, such as whether (B. cf. pinnicaudatus) or not (G. cf. anguillaris) nuclear rotation occurs, permanence of the cytoplasmic channel, and type and localization of the nuclear fossa. In the G. cf. anguillaris spermatozoon the nucleus is spherical with highly condensed chromatin. The nuclear fossa is shallow and lateral and is associated with the centriolar complex through stabilizing fibrils. The midpiece is short, with many vesicles, a cytoplasmic channel, and elongate mitochondria. In the B. cf. pinnicaudatus spermatozoon the ovoid nucleus is elongated lateral and posterior to the centriolar complex, and has highly condensed chromatin. The eccentric nuclear fossa is of the moderate type, and contains the entire centriolar complex. The midpiece is long, with numerous vesicles, elongate mitochondria, and no cytoplasmic channel. In both species the flagella are laterally disposed in relation to the nucleus and comprise of the classical 9+2 axoneme. Most of the characteristics found in the spermatozoa of these two species of Gymnotiformes are shared with species of Characiformes, whereas only a few are also found in Siluriformes. This suggests that Gymnotiformes and Characiformes may be more closely related than previously proposed.     

Scaling of suction feeding performance in the catfish Clarias gariepinus

Ontogenetic changes in the absolute dimensions of the cranial system together with changes in kinematics during prey capture can cause differences in the spatiotemporal patterns of water flow generated during suction feeding. Because the velocity of this water flow determines the force that pulls prey toward and into the mouth cavity, this can affect suction feeding performance. In this study, size-related changes in the suction-induced flow patterns are determined. To do so, a mathematical suction model is applied to video recordings of prey capturing Clarias gariepinus ranging in total length from 111 to 923 mm. Although large C. gariepinus could be expected to have increasing peak velocities of water flow compared with small individuals, the results from the hydrodynamic model show that this is not the case. Yet, when C. gariepinus becomes larger, the expansive phase is prolonged, resulting in a longer sustained flow. This flow also reaches farther in front of the mouth almost proportionally with head size. Forward dynamical simulations with spherical prey that are subjected to the calculated water flows indicate that the absolute distance from which a given prey can be sucked into the mouth as well as the maximal prey diameter increase substantially with increasing head size. Consequently, the range of potential prey that can be captured through suction feeding will become broader during growth of C. gariepinus. This appears to be reflected in the natural diet of this species, where both the size and the number of evasive prey increase with increasing predator size.     

The catfish fauna of Lake Victoria after the Nile perch upsurge

Stocks of the indigenous catfish species of Lake Victoria have decreased dramatically since the beginning of the 1980s. This decline coincided with the Nile perch boom and concomitant ecological changes in the lake. In deep water, where Nile perch densities were higher, the decline proceeded more quickly than in shallow water. In the former all catfishes eventually vanished. Of the two largest species, Clarias gariepinus and Bagrus docmak, juveniles disappeared faster than adults. This indicates that predation by Nile perch may have played an important role in their decline. Other possible impacts were the deoxygenation of deepwater areas and the decline of haplochromine cichlids which were an important food source for B. docmak, C. gariepinus and Schilbe intermedius. The various catfish species were not all affected to the same extent. The endemic Xenoclarias eupogon, which lived predominantly in deep water, may have become extinct. B. docmak currently seems to be mainly restricted to refugia in rocky habitats. Synodontis victoriae and S. afrofischeri are still present in small numbers in shallow littoral areas. Schilbe intermedius and C. gariepinus seem to be the least affected of the catfishes in littoral and sublittoral areas. This may be caused, among other reasons, by their smaller habitat overlap with Nile perch than the other species. S. intermedius is partly pelagic, and a considerable part of the C. gariepinus stock lives in bodies of water surrounding the lake. The patterns of decline of the catfishes are very similar to those observed for haplochromine cichlids in the lake. The importance of catfishes for the fisheries in the lake is currently negligible.     

Aquaculture implications of growth variation in the African catfish: Heterobranchus longifilis (Val.) reared under controlled conditions

Individual (mass) selection experiments were undertaken on different batches of Heterobranchus longifilis siblings to investigate factors such as significant size variation among larvae and juveniles that influence the darting (shooting) phenomenon. Darting was studied by subjecting treatment groups of fish larvae to (1) different feeding levels: 3% and 10% of body weight, and continuously at equal stocking rate. (2) different stocking rates: 600, 300 and 150 fish/30 L at the same feeding level. (3) effects of introduction of shelter under similar stocking and feeding levels and regime. A darting (shooting) event was registered when fish darted abruptly over a length of approximately three times the estimated total body length. The study concluded that food and density were the critical epigenetic factors responsible for size variation in juvenile H. longifilis.