Model of jaw depression during feeding in Astatotilapia elegans (Teleostei: Cichlidae): Mechanisms for energy storage and triggering

Suction feeding in Astatotilapia elegans occurs by a series of rapid, coupled movements of various head parts. The lower jaw rotates with respect to the neurocranium through an angle of 62° in less than 15 ms. The power requirements for jaw depression are calculated from a mathematical model and may reach a peak of ±4 watt in a 12-cm-long specimen. Data from the literature on mechanical output of fish muscle suggest that a muscular volume equal to 44% of the volume of the fish (= 45 cm³) should be required, if it is premised that movement and muscle shortening are directly coupled. Therefore, we assumed that storage and release of strain energy must be considered. The work demands for depression equal 0.014 J. The amount of energy storable in only three pairs of head ligaments can be estimated between 0.008 J and 0.05 J. The use of strain energy implies initial blocking and subsequent triggering of the movement. The position of the hyalomandibular connection, dorsal to the mandibular-suspensorial articulation, appears to be of crucial importance in balancing the forces of sternohyoideus and the body muscles. Triggering at the onset of jaw depression occurs by the contraction of the levator operculi. The line of action of the hyalomandibular connection is lowered beneath the jaw suspension, which raises the equilibrium of forces. Elastic recoil can occur thereafter.     

Variability of the fast suction feeding process in Astatotilapia elegans (Teleostei: Cichlidae): a hypothesis of peripheral feedback control

Movement analysis of the 'volume suction' feeding type in Astarotilapia elegans suggests the existence of an inhibiting peripheral feedback control on the fast movements of the head parts, apparently triggered by the food items entering through the mouth aperture. As soon as the prey passes the mouth, rostral expansion of the buccopharyngeal cavity stops. On the basis of a mathematical model and physiological evidence, respectively, visual and chemical perception must probably be excluded as the initial stimulus of the feedback control. The simulation of the hydrodynamic characteristics of the suction flow at the level of the gape reveals sudden changes in the pressure and acceleration waves coupled to the moment of prey uptake. These fluctuations are premised to generate the triggering signal. The possibility of modulation entails re-evaluation of the neuro-motoric preprogamming concept.     

The intramandibular linkage in Astatotilapia elegans (Teleostei: Cichlidae): appearance and function of the meckelian cartilage

During expansion of the bucco-pharyngeal cavity in Astatotilapia elegans , the lower jaws depress and follow the movements of the suspensoria (abduction). Moreover, the angulo-articular part of each ramus rotates around its longitudinal axis in such a manner that the retro-articular process moves laterally (= lateral inclination). As a result, a part of the meckelian cartilage undergoes a torsion.
Elastic moduli of the meckelian cartilage are calculated. It is supposed that during adduction of the head parts, the gained strain energy (due to torsion of the cartilage rod) may be sufficient to move the retro-articular process medially (= medial inclination) by means of elastic resilience. The processes are then held in their resting position, closely to the ventral side of the head. Undoubtedly, the meckelian cartilage acts as a spring, preventing passive lateral movements of the retro-articular processes. In this way, the streamlining of the fish-head is improved when the head parts are in an adducted position.     

Observations on tooth development and implantation in the upper pharyngeal jaws in Astatotilapia elegans (Teleostei,Cichlidae)

Prominent stages in the development of teeth, associated with the upper pharyngeal jaws in early postembryonic stages of the mouth brooding cichlid A statotilapia elegans were studied on semithin sections in relation to changes in the underlying endoskeletal parts and to the formation of the dentigerous bone. Because the pattern of tooth implantation on infrapharyngobranchial III-IV is constant, at least in early postembryonic stages, it is possible to trace the life history of a given tooth by tracing its homologue throughout the ontogenetic series. A probable causal relationship exists between tooth development and erosion of the underlying cartilage. Fully developed, though unerupted teeth, differentiate annular bone of attachment, which, depending on its position, is formed either outside the cartilage or within the previously induced erosion cavities. Attachment bone of adjacent teeth fuses to build up the dentigerous bone, which, as a result, may be situated within the area previously occupied by cartilage. As soon as the tooth has built up its bone of attachment, it may erupt. The collagenous matrix between tooth and attachment bone persists and gives rise to the movable connection between both. Differentiation of teeth on infrapharyngobranchial III-IV, together with enlargement of the dentigerous bone, proceeds from the lateral and the rostral border, where new germs constantly form. The appearance of new germs on infrapharyngobranchial II is more unpredictable.     

Tilapia (Teleostei: Cichlidae) status in Nicaraguan natural waters

We surveyed the watersheds covering more than 80% of the surface area of Nicaragua, and review the history of deliberate introductions and unintentional invasion of tilapias, Oreochromis spp., into the freshwater of Nicaragua. The species have become widely established, with a range of negative consequences for the rich natural fish fauna of this Central American country. Tilapias compete directly with native cichlids in a number of ways, and have also supplanted native species as food fish in local markets. We suggest that introduced tilapias may have been responsible for the outbreak of blindness in native cichlids. We make recommendations on the management of these exotic species and on further introductions. An erratum to this article can be found at     

Initial development of Oreochromis niloticus (Teleostei: Cichlidae) otolith

Otoliths of Oreochromis niloricus initiate as separate protein granules, surrounded by numerous dark particles. These granules merge to form the sub-primordium, after a layer of calcium and a layer of protein has been added to each one. Large amounts of protein are accreted on to the subprimordium to form the primordium. Calcification takes place shortly afterwards. In the otocyst lumen filaments and vesicles are abundant and associated. They seem to contribute to the proteins of the primordium and the calcified otolith. Mitochondria, Golgi complexes and intracellular granules are abundant in the epithelium, from which vesicles extrude.     

Phylogeny and biogeography of cichlid fishes (Teleostei: Perciformes: Cichlidae)

Family level molecular phylogenetic analyses of cichlid fishes have generally suffered from a limited number of characters and/or poor taxonomic sampling across one or more major geographic assemblage, and therefore have not provided a robust test of early intrafamilial diversification. Herein we use both nuclear and mitochondrial nucleotide characters and direct optimization to reconstruct a phylogeny for cichlid fishes. Representatives of major cichlid lineages across all geographic assemblages are included, as well as nearly twice the number of characters as any prior family‐level study. In a strict consensus of 81 equally most‐parsimonious hypotheses, based on the simultaneous analysis of 2222 aligned nucleotide characters from two mitochondrial and two nuclear genes, four major subfamilial lineages are recovered with strong support. Etroplinae, endemic to Madagascar (Paretroplus) and southern Asia (Etroplus), is recovered as the sister taxon to the remainder of Cichlidae. Although the South Asian cichlids are monophyletic, the Malagasy plus South Asian lineages are not. The remaining Malagasy lineage, Ptychochrominae, is monophyletic and is recovered as the sister group to a clade comprising the African and Neotropical cichlids. The African (Pseudocrenilabrinae) and Neotropical (Cichlinae) lineages are each monophyletic in this reconstruction. The use of multiple molecular markers, from both mitochondrial and nuclear genes, results in a phylogeny that in general exhibits strong support, notably for early diversification events within Cichlidae. Results further indicate that Labroidei is not monophyletic, and that the sister group to Cichlidae may comprise a large and diverse assemblage of percomorph lineages. This hypothesis may at least partly explain why morphological studies that have attempted to place Cichlidae within Percomorpha, or that have tested cichlid monophyly using only “labroid” lineages, have met with only limited success. © The Willi Hennig Society 2004.     

Midwater spawning inHaplochromis chrysonotus (Boulenger) (Teleostei: Cichlidae) in Lake Malawi

Synopsis Haplochromis chrysonotus, a semipelagic shoaling cichlid from Lake Malawi, has been observed spawning in open water up to ten metres above the substratum. It is suggested that open water spawning without the establishment of a substratum-based territory eliminates competition with other species for breeding space and that it may be a stage in the evolution of a totally pelagic mode of life.     

Phylogeny and species diversity of the genus Herichthys (Teleostei: Cichlidae)

We provide a review of the systematics of Herichthys by evaluating the usefulness of several mitochondrial and nuclear genetic markers together with morphological data. The nDNA next‐generation sequencing ddRAD analysis together with the mtDNA cytochrome b gene provided well‐resolved and well‐supported phylogenies of Herichthys. On the other hand, the nDNA S7 introns have limited resolution and support and the COI barcoding analysis completely failed to recover all but one species of Herichthys as monophyletic. The COI barcoding as currently implemented is thus insufficient to distinguish clearly distinct species in the genus Herichthys that are supported by other molecular markers and by morphological characters. Based on our results, Herichthys is composed of 11 species and includes two main clades (the H. labridens and H. cyanoguttatus species groups). Herichthys bartoni is in many respects the most plesiomorphic species in the genus and has a conflicting phylogenetic position between mtDNA and nDNA markers, where the robust nDNA ddRAD data place it as a rather distant basal member of the H. labridens species group. The mtDNA of H. bartoni is on the other hand only slightly divergent from the sympatric and syntopic H. labridens, and the species thus probably have hybridized in the relatively recent past. The sympatric and syntopic Herichthys steindachneri and H. pame are supported as sister species. The Herichthys cyanoguttatus species group shows two well‐separated basal species (the northernmost H. minckleyi and the southernmost H. deppii) followed by the closely related and centrally distributed species H. cyanoguttatus, H. tepehua, H. carpintis, and H. tamasopoensis whose relationships differ between analyses and show likely hybridizations between themselves and the two basal species as suggested by conflicts between DNA analyses. Several instances of introgressions/hybridizations have also been found between the two main clades of Herichthys.     

Polychromatism of the scale-eaterPerissodus microlepis (Cichlidae,Teleostei) in relation to foraging behavior

Variation in foraging behavior in a population of the scale-eaterPerissodus microlepis was studied on the northwest coast of Lake Tanganyika. Differences in body coloration were found among adult and subadult individuals, which were classified into 4 color morphs designated as Beige, Dark, Grey and Stripe. These color morphs were not strictly related to either sex or size. Each morph spent much time in specific microhabitats and had a major hunting technique that differed from other morphs. Beige morph. which predominated in number, ambushed prey at open surfaces of the substrate, whereas Dark morph used the shade of rock as an ambush site. Grey morph mixed in schools of fisheds hovering in midwater to attack school members, and Stripe morph cruised in the water column and stooped mainly at bottom-fishes. Prey preference differed among the morphs corresponding to their hunting techniques but successful attack rates were similar among them. Observations of marked individuals demonstrated adherence to particular hunting techniques and, in some cases, to particular hunting sites. Intraspecific foraging specialization is discussed in relation to the function of body color and diversity of life styles of prey fishes.     

Phylogeny,taxonomy, and evolution of Neotropical cichlids (Teleostei: Cichlidae: Cichlinae)

Despite recent progress on the higher‐level relationships of Cichlidae and its Indian, Malagasy, and Greater Antillean components, conflict and uncertainty remain within the species‐rich African, South American, and Middle American assemblages. Herein, we combine morphological and nucleotide characters from the mitochondrial large ribosomal subunit, cytochrome c oxidase subunit I, NADH dehydrogenase four, and cytochrome b genes and from the nuclear histone H3, recombination activating gene two, Tmo‐4C4, Tmo‐M27, and ribosomal S7 loci to analyse relationships within the Neotropical cichlid subfamily Cichlinae. The simultaneous analysis of 6309 characters for 90 terminals, including representatives of all major cichlid lineages and all Neotropical genera, resulted in the first well‐supported and resolved generic‐level phylogeny for Neotropical cichlids. The Neotropical subfamily Cichlinae was recovered as monophyletic and partitioned into seven tribes: Astronotini, Chaetobranchini, Cichlasomatini, Cichlini, Geophagini, Heroini, and Retroculini. Chaetobranchini + Geophagini (including the “crenicichlines”) was resolved as the sister group of Heroini + Cichlasomatini (including Acaronia). The monogeneric Astronotini was recovered as the sister group of these four tribes. Finally, a clade composed of Cichlini + Retroculini was resolved as the sister group to all other cichlines. The analysis included the recently described ?Proterocara argentina, the oldest known cichlid fossil (Eocene), which was placed in an apical position within Geophagini, further supporting a Gondwanan origin for Cichlidae. These phylogenetic results were used as the basis for generating a monophyletic cichline taxonomy. © The Willi Hennig Society 2008.     

A revision of the genus Labidochromis (Teleostei: Cichlidae) from Lake Malawi

The diagnosis of the genus Labidochromis is revised to include species with bicuspid outer teeth. Sixteen species, of which 13 are new, are described and illustrated and the validity of five others is discussed. The interrelationships between members of the genus and the relationship of the genus to other genera are discussed. No clearly defined sister group to the genus has been found either within or outside the Lake Malawi cichlid species flock. A key based on the coloration of known species is included.     

Environmental determination of sex in Apistogrammai (Cichlidae) and two other freshwater fishes (Teleostei)

Environmental sex determination by temperature could be revealed significantly in 33 Apistogramma-species and in Poecilia melanogaster . In some, but not all, Apistogramma-species pH also influences the sex ratio, whereas neither temperature nor pH affect the sex ratio of Pseudocrenilabrus multicolor victoriae . The sex in offspring of A. trifasciata is determined within a sensitive period of about 30 to at least 40 days after spawning.     

Food composition and feeding habits of Astatotilapia flaviijosephi (Lortet) in Lake Kinneret (Israel)

Food composition and feeding habits of 102 specimens of Astatotilapia (Haplochromis) flaviijosephi (Lortet, 1883), collected in Lake Kinneret during 1973–1975 and 1981–1982, was studied. A. flaviijosephi is relatively abundant in the littoral zone of the lake. Females feed mostly on chironomid larvae, oligochaetes and amphipods whilst males consume mostly gastropods. Small fry are rarely preyed upon or collected by both sexes. The adaptive relation between the teeth and the type of food and the ecological status of A. flaviijosephi in the Kinneret ecosystem is discussed.     

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.     

Characterization of sex chromosomes in three deeply diverged species of Pseudocrenilabrinae (Teleostei: Cichlidae)

The African cichlid radiations have created thousands of new cichlid species with a wide diversity of trophic morphologies, behaviors, sensory systems, and pigment patterns. In addition, recent research has uncovered a surprising number of young sex chromosome systems within African cichlids. Here, we refine methods to describe the differentiation of young sex chromosomes from whole genome comparisons. We identified a novel XY sex chromosome system on linkage group 14 in Oreochromis mossambicus, confirmed a linkage group 1 XY system in Coptodon zillii, and also defined the limits of our methodology by examining a ZW system on linkage group 3 in Pelmatolapia mariae. These data further demonstrate that cichlids are an excellent model system for understanding the early stages of sex chromosome evolution.