Cytological and morphological ontogenesis and involution of the thymus in cichlid fishes (Cichlidae,Teleostei)

The ontogenesis and involution of thymus in cichlid fishes was studied with the aim of comparing development in the bottom-spawning species Tilapia zillii and T. tholloni, and in the mouth-brooding species Oreochromis auratus, O. niloticus, O. mossambicus, and Sarotherodon galilaeus. For comparison, data are also given on bottom-spawning Cichlasoma spp. from America and mouth-brooding Pseudotropheus auratus and Aulonocara nyassae from Africa. Developmental changes were followed histologically by means of light and electron microscopy of sections, beginning with embryos 24 h after fertilization, up to 14-year-old specimens of O. auratus. In all these fish, the anlagen of the thymus glands begins from the third and fourth gill pouches, and their development shows a high correlation with the pace of general organogenesis, which differs greatly in the bottom-spawning and mouth-brooding cichlids. In juveniles of bottom-spawners of 20–40 mm total length and in mouth-brooders of 40–60 mm total length, three cell types are present in the thymus: thymocytes, with large, dense nuclei; epitheliocytes, with long cell extensions containing bundles of tonofibrils; and reticulocytes, with short, granulated cell extensions. Hassall's corpuscles start to develop in larvae of T. zillii at 20–35 mm total length, and in specimens of 40 mm and more total length the corpuscles are typical, formed by inner and outer rings of epitheliocytes. At 30–45 mm total length, cell debris starts to accumulate in the interior of the corpuscles as an early sign of regression. As involution continues, macrophages accummulate within and around the Hassall's corpuscles. The epitheliocyte rings are eventually completely broken down. Isles of thymocytes persist in tilapias from the age of 1–14 years, but most of the thymus volume is occupied by blood lacunae and pigmented macrophage aggregations. The morphology is similar in the mouth-brooding species Pseudotropheus beginning at 1.5 years of age. © 1995 Wiley-Liss, Inc.     

Molecular phylogeny and biogeography of the Neotropical cichlid fish tribe Cichlasomatini (Teleostei: Cichlidae: Cichlasomatinae)

We have conducted the first comprehensive molecular phylogeny of the tribe Cichlasomatini including all valid genera as well as important species of questionable generic status. To recover the relationships among cichlasomatine genera and to test their monophyly we analyzed sequences from two mitochondrial (16S rRNA, cytochrome b) and one nuclear marker (first intron of S7 ribosomal gene) totalling 2236 bp. Our data suggest that all genera except Aequidens are monophyletic, but we found important disagreements between the traditional morphological relationships and the phylogeny based on our molecular data. Our analyses support the following conclusions: (a) Aequidens sensu stricto is paraphyletic, including also Cichlasoma (CA clade); (b) Krobia is not closely related to Bujurquina and includes also the Guyanan Aequidens species A. potaroensis and probably A. paloemeuensis (KA clade). (c) Bujurquina and Tahuantinsuyoa are sister groups, closely related to an undescribed genus formed by the 'Aequidens'pulcher-'Aequidens'rivulatus groups (BTA clade). (d) Nannacara (plus Ivanacara) and Cleithracara are found as sister groups (NIC clade). Acaronia is most probably the sister group of the BTA clade, and Laetacara may be the sister group of this clade. Estimation of divergence times suggests that the divergence of Cichlasomatini started around 44Mya with the vicariance between coastal rivers of the Guyanas (KA and NIC clades) and remaining cis-andean South America, followed by evolution of the Acaronia-Laetacara-BTA clade in Western Amazon, and the CA clade in the Eastern Amazon. Vicariant divergence has played importantly in evolution of cichlasomatine genera, with dispersal limited to later range extension of species within genera.     

Structure and development of first-generation teeth in the cichlid Hemichromis bimaculatus (Teleostei, Cichlidae)

In order to build a reference system to assess results of ongoing in vitro experiments on the study of epithelial-mesenchymal interactions during odontogenesis in actinopterygians, we have chosen to study the first-generation teeth of the cichlid Hemichromis bimaculatus from initiation until attachment both at the light and transmission electron microscopical level. Although their development follows the general pattern of teleost tooth formation, first-generation teeth show peculiarities compared with later tooth generations, including their size, bare emergence from the epithelium, absence of dentinal tubules and of nerves and capillaries in the pulp cavity, and organization of the outer dental epithelium. Four developmental stages (a to d) prior to attachment (stage e) have been distinguished. The oral epithelium invaginates into the underlying mesenchyme (stage a) and is later folded to form a bell-shaped dental organ (stage b) without any primordial thickening, or any other morphological indication of imminent invagination. Then, the collagenous enameloid matrix is laid down, most probably by the odontoblasts (early stage c), soon followed by predentine deposition and the beginning of enameloid mineralization (late stage c). With ongoing dentinogenesis, the enameloid matrix matures (stage d), i.e. the organic constituents are removed and the matrix further mineralizes. Finally (stage e), an annular collar of attachment bone is deposited to fix the tooth onto the underlying bone.     

Ontogenesis and functional metamorphosis of the head-kidney in bottomspawner and mouthbrooder cichlid fishes (Cichlidae,Teleostei)

Histological and electron-microscopical studies of the head-kidney ontogenesis in a group of bottomspawner and mouthbrooder cichlid fishes reveal that this organ undergoes three functional and structural stages. At the initial stage, 2–12 days after fertilization, the head-kidney is a filtering organ of two glomeruli and adjacent renal tubules; the second stage, beginning 12–30 days after fertilization, is a functionally mixed one, with the nephron and developing hemopoietic and endocrine, chromaffin, and interrenal cells; during the third stage, which begins 30–60 days after fertilization, the nephron degenerates, and the head-kidney starts functioning solely as a lympho-adrenal organ. With age the interrenal tissue expands, forming fields, especially along the blood vessels. The number of melano-macrophage centers progressively increases within the hemopoietic parts. Data are given on the various cell types, the reticuloendothelial stroma, and the renal part of this organ in various species. The earlier ontogenesis and function of the head-kidney, as thymus and other organs, in bottomspawner larvae compared to the more protected mouthbrooders provide support for earlier free functioning in a dangerous environment. © 1996 Wiley-Liss, Inc.     

Comparative ecology of Serranochromis species (Teleostei: Cichlidae) in the Upper Zambezi River floodplain

The Upper Zambezi River drainage of Zambia contains six Serrunochromis species belonging to the largemouth, piscivorous subgenus Serranochromis and three smallmouth, invertebrate-feeding species of the subgenus Sargochromis. A survey of the Barotse floodplain and associateddrainages revealed that two species, S. (S.) longimanus (Boulenger) and S. (S.) thumbergi (Castelnau), were uncommon on the floodplain. S. thumbergi appeared to be more abundant in the Kafue, Chobe, and Kabompo rivers than the Upper Zambezi proper. Adults of the three largest species (i.e. S. (S) robustus (Günther), S. (S.) altus (Winemiller & Kelso-Winemiller), and S. (Sar.) giurdi (Pellegrin)) inhabit the main channel of the Zambezi River during low water, whereas most adults of the other four species (i.e., S. (S.) macrocephalus (Boulenger), S. (S.)ungusticeps (Boulenger), S. (Sar.) codringtoni (Boulenger), and S. (Sar.) carlottae (Boulenger)) were collected from lagoons, canals and backwaters. Comparisons of population size structure and gonad condition during the falling water (May-August) and low water (September- December) seasons indicated that the seven common Serranochromis initiate spawning near the end of the low water period. Examination of gonad condition during the pre-spawning period indicated that Serrunochromis attain maturity at the following sizes: S. robustus, males (275-300 mm s.l .), females (250-275 mm); S. macrocephalus, both sexes (150-200 mm); S. altus, males (275-325 mm), females (250-300 mm);S. angusticeps, males (250-275 mm), females (175-225 mm); S. giardi, males (275-300 mm), females (225-250 mm); S. codringtoni, males (150 mm), females (125-150 mm); S. carlottue, males (150-175 mm), females (100-125 mm). Age was estimated from scale annuli comprised of closely spaced circuli that corresponded to the period of rising water (December- March). S. robustus, S. altus, S. angusticeps, and S. giurdi matured at approximately 3 years, and S. macrocephalus, S. codringtoni, and S. curlottae matured at approximately 2 years in the Barotse floodplain. The average numbers of mature oocytes per female were as follows: S. robustus- 1165; S. macrocephalus-810; S. altus-920; S. angusticeps-627; S. codringtoni-580; S. carlottue-362. Fecundity exhibited a weak positive relationship with female size. Analysis of stomach contents showed that interspecific diet overlap was low among both immature and mature size classes of Serrunochromis. Adult size classes of S. robustus fed primarily on small Synodontis catfishes, S. mucrocephalus fed mostly on mormyrid and cichlid fishes, S. altus specialized on mormyrids, and S. angusticeps fed primarily on small cyprinid and characid fishes. The three members of the subgenus Sargochromis consumed mostly invertebrates. Molluscs were the most important item in the diet of S. giardi, and aquatic insects and fish scales were predominant in the diet of S. carlottae. Larger size classes of S. codringtoni consumed large amounts of seeds (Graminaceae). If habitat and diet are considered together, niche separation among Upper Zambezi Serrunochromis species appears to be nearly complete.     

Foraging on patchily distributed prey by a cichlid fish (Teleostei,Cichlidae): A test of the ideal free distribution theory

Cichlid fish (Aequidens curviceps) distributed themselves and allocated their foraging time between two drift food patches in close approximation to the patch profitability ratio, as predicted by the ideal free distribution theory. The fish thereby achieved similar average feeding rates in the two patches, in two of three patch profitability ratio experiments. However, one major assumption of the ideal free model was violated, since individual fish differed in their competitive abilities for limited food resources, which resulted in unequal payoffs among individuals within each patch. Individual variation in feeding rates, and thus in competitive ability, was not related to despotism, but perhaps rather to individual differences in perceptual ability and in the ability to learn which patch was currently the more profitable. The strategy used by the fish to assess patch profitability included sampling available patches. However, individual fish switched (sampled) patches with varying frequency. Sampling had an associated cost, since high-frequency switchers had lower feeding rates on average than low-frequency switchers. Differences in foraging strategy among the fish therefore contributed to the observed in-equality in individual payoffs within patches.     

Breeding behavioural pattern of the lake fish Tilapia discolour Gunther (Teleostei, Cichlidae) in captivity

Close observation of the breeding activities of specimens of Tilapia discolour obtained from Lake Bosomtwe and held captive in glass aquaria, a net cage and a pond showed that this species practises substrate-brooding; the eggs were incubated and hatched in prepared nests. However, brief egg-carrying tendencies were observed; both the male and female brood fish carried the eggs in their mouths from the sites of spawning to new sites. Both sexes also made and guarded the nests, incubated the eggs and provided protection to the young fry.     

Phylogeny of the Neotropical cichlid fish tribe Cichlasomatini (Teleostei: Cichlidae) based on morphological and molecular data, with the description of a new genus

Phylogenetic relationships among cichlasomatine cichlids were studied using an extensive taxon sampling and both morphological and molecular data sets. A new genus, Andinoacara n. gen. with six species ( A. pulcher-rivulatus group of previous authors) from trans-andean South America and NW cis-andean South America, is described based on results of phylogenetic and diagnosability analyses and tests of alternative topologies Our results demonstrate that cichlasomatine cichlid diversity is divided into five principal lineages composed of eleven genera and three suprageneric clades: the [( Bujurquina , Tahuantinsuyoa ), ( Andinoacara ) (BAT) clade; the ( Cleithracara , ( Nannacara , Ivanacara )] clade (NIC) plus Laetacara and 'Aequidens' hoehnei ; and the ( Aequidens , Cichlasoma ) clade, where Aequidens is paraphyletic to Cichlasoma . Two former Aequidens species are additionally transferred into Krobia ( K. potaroensis , K. paloemeuensis ). 'Aequidens' hoehnei probably represents a unique evolutionary lineage and would thus qualify for a separate generic status. Molecular data are yet not available for this species and its generic status requires further study. Relationships between the three suprageneric clades and between Acaronia and Krobia could not be convincingly resolved with our data set of two mitochondrial (16S and cyt b ) and two nuclear (S7 and RAG1) molecular markers and 96 morphological characters.     

Phylogeographic structure and gene flow in the scale-eating cichlid Perissodus microlepis (Teleostei, Perciformes, Cichlidae) in southern Lake Tanganyika

One of the most fragmented habitats in freshwater lakes is the rocky littoral zone, where the already richly structured habitat is frequently interspersed with more pronounced barriers such as sandy bays, river estuaries and deep slopes. Although habitat fragmentation generally constrains the dispersal of specialized rock-dwelling species, patterns of population structure vary in sympatric taxa due to species-specific traits. In the present study, we examine the phylogeographic and population genetic structure of Perissodus microlepis , a presumptively highly mobile scale-eating cichlid fish endemic to Lake Tanganyika with a lake-wide distribution in the rocky littoral zone and no obvious geographical colour variation. Analysis of the mitochondrial DNA of six populations in the southern end of the lake suggests isolation by distance along rocky shoreline. Across a large muddy bay, a phylogeographic break indicates that environmental barriers restrict gene flow even in this highly mobile species. Restricted dispersal across the bay is not necessarily a consequence of an intrinsic propensity to avoid sand, but may be connected with the association between P. microlepis and other rock-dwelling fish, which the scale-eaters mimic and intermingle in order to be able to approach other fish to rip off scales from their bodies.     

First description of monogenean parasites in Lake Tanganyika: the cichlid Simochromis diagramma (Teleostei, Cichlidae) harbours a high diversity of Gyrodactylus species (Platyhelminthes, Monogenea)

Lake Tanganyika harbours the most diverse endemic cichlid fish assemblage of Africa, but its monogenean fish parasites have not been investigated. Here we report, for the first time, on the Gyrodactylus parasites in this hotspot of fish biodiversity. Haptor morphometrics and nuclear ribosomal DNA sequences revealed 3 new species on Zambian Simochromis diagramma: Gyrodactylus sturmbaueri n. sp., G. thysi n. sp. and G. zimbae n. sp. Their distinct morphology and strong genetic differentiation suggest that they belong to distant lineages within the genus Gyrodactylus, and phylogenetic reconstructions suggest affinities with other genera of gyrodactylids. Additional U-shaped haptoral plates in G. thysi n. sp. and a second large spine-like structure in the male copulatory organ of G. zimbae seem to represent new features for the genus. Such large diversity on a single host species can probably be explained by host-switching events during the course of evolution, in agreement with the generally accepted concept that ecological transfer is an important aspect of gyrodactylid speciation. Additional parasitological surveys on other host species, covering a broader phylogenetic and geographical range, should clarify the evolutionary history of Gyrodactylidae on cichlids in the African Great Lake and other parts of Africa.     

A new Apistogramma species (Teleostei, Cichlidae) from the Rio Negro in Brazil and Venezuela

Apistogramma diplotaenia sp.n. occurs in the upper and middle Rio Negro. The largest among 186 specimens studied is a 29.2 mm SL male, the largest female is 22.9 mm SL. Distinguishing characters include a unique colour pattern, consisting of two dark bands along the side which converge anteriorly and posteriorly. A comparative osteological study, including other species of Apistogramma and the genera Taeniacara and Apistogrammoides , indicates a progressive forwards loss of infraorbital ossicles and foramina among dwarf cichlids. The long, rod-like interarcual cartilage of Apistogramma, Apistogrammoides and Satanoperca possibly represents a plesiomorphic condition otherwise unknown among cichlids.     

Feeding habits of cichlid species (Perciformes: Cichlidae) in Caobas lake, Quintana Roo, Mexico

Feeding habits of seven cichlid species (Archocentrus octofasciatus, A. spilurus, "Cichlasoma" robertsoni, "C." synspilum, "C." urophthalmus, Petenia splendida, Thorichthys meeki) in Lake Caobas, southern Yucatan Peninsula, were studied. Samples were taken with enclosure and cast nets during the dry and rainy seasons of 1995 (day and night). The environment was characterized by measuring temperature, conductivity and pH. All individuals were below 41 mm SL (N = 281). Frequency of occurrence and prey abundance were analyzed. Main prey items were chironomids, mites, copepods, cladocerans, and ostracods. The cichlids fed mainly on zooplankton, with the partial exceptions of P. splendida (piscivore), "C." synspilum and A. spilurus (herbivores). A cluster analysis showed that the most similar trophic spectra were those of T. meeki, "C." robertsoni and "C." salvini, which were also the least diverse. "C." synspilum and A. spilurus had an intermediate distance between their diets and those of other species. The species with the most distinctive feeding composition were P. splendida (with the most diverse and equitable diet) and the omnivore A. octofasciatus (whose diet was the richest one). T. meeki showed quantitative diel, ontogenetic, and seasonal diet changes, but none between sexes. "C." robertsoni, "C." salvini and "C." synspilum differ in food habits in Caobas and in other localities, a fact that underscores the trophic adaptability of cichlids. Trophic overlap between cichlids in Caobas could imply absence of competition, perhaps because resources are abundant in the ecosystem.     

Histochemical study on the intestine goblet cells in cichlid and poecilid species (Teleostei)

Histochemical properties of intestine goblet cells in firemouth cichlid, zebra mbuna, freshwater angelfish and platyfish are described. Goblet cells occurred regularly in the epithelial cell layer throughout the entire intestine, they were strongly coloured by alcian blue at pH 2.5. This colour got gradually weaker when the pH was reduced, but still after alcian blue at pH 0.2 these cells displayed a distinct blue colour. When the goblet cells were treated with periodic acid-Schiff (PAS), they displayed a strong purple-magenta colour. The findings that a number of goblet cells displayed various colours between blue and purple-magenta when acidic alcian blue was followed by PAS, and between blue and red-brown when acidic alcian blue was followed by neutral red, may reflect different ages or stages of development and differentiation for these cells. However, such results may also suggest a true cellular heterogeneity in the present population of goblet cells, reflecting that the intestine mucus layer has a number of roles in teleosts like lubrication, protection, immunological defence, digestion and absorption.In the ferritin injected specimens of firemouth cichlid and platyfish, a number of macrophage-like cells in intestine wall displayed Prussian blue precipitations in tissue treated with acid ferrocyanide, suggesting that these cells play a cleansing role in the intestinal wall. No ferritin uptake was seen in the intestine goblet cells and eosinophilic granule cells.     

The inheritance of aggressiveness in the convict cichlid fish, Cichlasoma nigrofasciatum (Pisces: Cichlidae).

Intraspecific comparisons in aggression were made between X-irradiated (2 times 500 R) ancestors and untreated control animals. Bites delivered to sub-adult conspecifics were counted to measure aggressiveness. When compared with corresponding controls, a significant reduction in male aggressiveness was found in the post-irradiation F1 generation. A greater variation in bites delivered occurred among the F2 post-irradiation generations than among the control equivalents. Three distinct back-cross generations were obtained as follows. a single F1 female from the irradiated line was mated to (a) her high aggressive (I-A) and (b) her low aggressive (I-NA) F2 son; a F1 control female was mated to her low aggressive F2 son. In the back-cross generations, both I-A and C-NA males were more aggressive than I-NA males. No differences were found between I-A and C-NA back-cross males. The findings are discussed in terms of a two-factor model based on Mendelian laws.     

A new pelagic predatory pike cichlid (Teleostei: Cichlidae: Crenicichla) from the C. mandelburgeri species complex with parallel and reticulate evolution

The Crenicichla mandelburgeri species complex from the Middle Paraná shows parallel evolution of ecomorphs to the unrelated C. missioneira species complex from the Uruguay River. In this article, we describe a new species from the C. mandelburgeri species complex that has evolved a parallel morphology and ecology to an unrelated species from the C. missioneira species complex (C. celidochilus). The new species is a pelagic predator that feeds predominantly on fishes and together with C. celidochilus is the only known pelagic species in the large riverine genus Crenicichla. The new species is endemic solely to a small tributary (the Urugua-í) of the Middle Paraná River where it is sympatric and partly syntopic with two other closely related endemic species that, however, differ strongly in their ecomorphologies (one is a generalistic invertivore and the other a specialized molluscivore). Mitochondrial DNA phylogeny finds the new species nested within the widespread C. mandelburgeri. Reduced genome-representation ddRAD analyses, however, demonstrate that this new species is of a hybrid origin and shares ancestry with C. ypo, one of the two studied sympatric species.

     

Description of a new Acaronia species (Teleostei, Cichlidae) from the Rio Orinoco and Rio Negro drainages

Acaronia vultuosa sp.n. occurs in the Rio Orinoco and upper Rio Negro drainages. In the Rio Negro it overlaps slightly the range of its only congener, A. nassa (Heckel), from which it can be distinguished by (1) the lack of microbranchiospines on the first gill arch, (2) thc separate instead of coalesced anterior gill rakers on the first gill arch, and (3) the colour pattern, which includes a much larger midlateral blotch and a different pattern of dark markings on the head.     

A new Aequidens species from the Rio Trombetas, Brasil, and redescription of Aequidens pallidus (Teleostei, Cichlidae)

Aequidens pallidus (Heckel) reaches c. 140 mm standard length and occurs in the drainages of the blackwater rivers Negro, Puraquequara, Preto da Eva and Uatumã in Brasil. Aequidens duopunctata Haseman is a junior synonym. Aequidens tubicen sp.n., was collected in the Trombelas and Mapuera rivers a little upstream of Cachoeira Porteira. The largest specimen is 116 mm standard length. It differs from all other Aequidens species in the possession of a dark spot at the inner angle of the preopercle and adjacent cheek. Aequidens pallidus and A. tubicen form a group diagnosed by a high number of vertebrae (14 + 13 = 27) and three colour pattern synapomorphies: lateral band positioned more dorsally and the midlateral spot positioned more posteriorly than in other Aequidens species, and caudal spot preceded by a large light spot at the dorsal margin of the caudal peduncle.     

Spermiogenesis and sperm ultrastructure in Cichla intermedia with some considerations about Labroidei spermatozoa (Teleostei, Perciformes, Cichlidae)

Spermiogenesis and spermatozoal structure were studied in Cichla intermedia, a primitive species of Neotropical Cichlids. The analysis shows that spermiogenesis is characterized by chromatin compaction, flagellum development, nuclear rotation, nuclear fossa formation and residual cytoplasm elimination. In the spermatozoa, the head is round, the nucleus contains highly condensed filamentous clusters of chromatin and an acrosome is absent. The nuclear fossa is slightly eccentric and shows a projection that penetrates into the nuclear outline. The proximal centriole is located in the initial segment of the nuclear fossa. The midpiece and the cytoplasmic channel are long. The mitochondria, about 10 in number, are round or slightly elongated, disposed in two layers around the initial segment of the flagellum. The flagellum has a classical 9+2 axoneme and two lateral fins. The data available show that no characteristics of spermiogenesis or spermatozoa are exclusively found in members of the suborder Labroidei. However, three characteristics seem to be exclusively observed in Cichlidae: (1) compact filamentous clusters of chromatin; (2) slightly eccentric nuclear fossa; and, (3) number of mitochondria.