The cichlid oral and pharyngeal jaws are evolutionarily and genetically coupled

Evolutionary constraints may significantly bias phenotypic change, while “breaking” from such constraints can lead to expanded ecological opportunity. Ray-finned fishes have broken functional constraints by developing two jaws (oral-pharyngeal), decoupling prey capture (oral jaw) from processing (pharyngeal jaw). It is hypothesized that the oral and pharyngeal jaws represent independent evolutionary modules and this facilitated diversification in feeding architectures. Here we test this hypothesis in African cichlids. Contrary to our expectation, we find integration between jaws at multiple evolutionary levels. Next, we document integration at the genetic level, and identify a candidate gene, smad7, within a pleiotropic locus for oral and pharyngeal jaw shape that exhibits correlated expression between the two tissues. Collectively, our data show that African cichlid evolutionary success has occurred within the context of a coupled jaw system, an attribute that may be driving adaptive evolution in this iconic group by facilitating rapid shifts between foraging habitats, providing an advantage in a stochastic environment such as the East African Rift-Valley.

Neural activation in the olfactory epithelium of East African cichlid in response to odorant exposure

Fish use olfaction to gain a variety of information. To know what they receive and how they receive is important for understanding the species. However, studies on fish olfactory or pheromone receptors are still few and most of them are based on the neural response from a cultured cell. Here, we established a method to detect a biological-derived neural response from the olfactory epithelium of East African cichlid, the most diversified fish lineage, by c-fos in situ hybridization. We tested the response of microvillous neurons, which are expected to be dominated by V2R-expressing neurons, against several odorants. We showed microvillous neurons respond to amino acids the most whereas they do not respond to conjugated steroids. We next tested the response of V2R receptors. Although some subfamilies of V2R responded to amino acids, other did not respond which contrasts with the traditional hypothesis. Especially, one V2R subfamily responded to arginine. Not all the copies in this subfamily responded to arginine, which indicates the ligand differentiation in the cichlid-specifically expanded subfamily. Finally, we tested the response of putative pheromone receptor V1R to male urine. For this experiment, we established a new method to collect urine from cichlid. We showed two V1R receptors responded to male urine. Moreover, we showed that V1R2 receptor responded to 4-hydroxyphenyl acetate and lithocholic acid. This study is expected to provide a basis for the study on the olfaction of East African cichlids.

Comparative osteology of three endemic cichlids (Iranocichla spp.) (Actinopterygii, Perciformes, Cichlidae) from southern Iran

Iranian cichlids are isolated disjunct populations of the African cichlids group, restricted to the Hormuz Strait region in the Hormuz basin, and are a very important group from both zoogeographic and phylogenetic points of view. Thus, the osteological structures, as one of the most reliable structures, of the three nominal species of Iranian cichlid fishes, Iranocichla hormuzensis Coad, 1982, from the Mehran River, Iranocichla persa Esmaeili, Sayyadzadeh et Seehausen, 2016, from Khorgo hot spring, and Iranocichla sp., from the Kol River were described and compared. Ten specimens of each species were cleared and stained with Alcian blue and Alizarin red. A digital camera was used for taking pictures and CorelDraw X6 software for preparing drawings. The three species have some differences in the caudal fin skeleton, shape of the urostyle, neural spine of preural 2, hypural spines 2, and 3, anterior part of parhypural and hypurapophysis, the size of neural spines of preural 2 and 3, hypural 3 and 4, and epurals. Although there are some differences among the species, osteology of these species is very conservative and other traits like behavioral and molecular should be used.

Four new species of Cichlidogyrus (Platyhelminthes, Monogenea, Dactylogyridae) from Lake Victoria haplochromine cichlid fishes, with the redescription of C. bifurcatus and C. longipenis

African cichlids are model systems for evolutionary studies and for host-parasite interactions, because of their adaptive radiations and because they harbour many species of monogenean parasites with high host-specificity. Here, we sampled five locations in southern Lake Victoria, the youngest of the African Great Lakes. We surveyed gillinfecting monogeneans from 18 cichlid species belonging to the Lake Victoria radiation superflock and two cichlid species representing two older and distantly related lineages. We found one species of Gyrodactylus (Gyrodactylidae, Monogenea), Gyrodactylus sturmbaueri Vanhove, Snoeks, Volckaert & Huyse, 2011, and seven species of Cichlidogyrus (Dactylogyridae, Monogenea). Four species are herein described: Cichlidogyrus pseudodossoui n. sp., C. nyanza n. sp., C. furu n. sp., C. vetusmolendarius n. sp.. Another species is reported but not formally described (because of few specimens and morphological similarity with C. furu n. sp.). Two other species are redescribed: Cichlidogyrus bifurcatus Paperna, 1960 and C. longipenis Paperna & Thurston, 1969. Our results confirm that the monogenean fauna of Victorian littoral cichlids displays lower species richness and lower host-specificity than that of Lake Tanganyika littoral cichlids. In C. furu n. sp., hooks V are clearly longer than the other hooks, highlighting the need to re-evaluate the current classification system of haptoral configurations that considers hook pairs III-VII as rather uniform. Some morphological features of C. bifurcatus, C. longipenis and C. nyanza n. sp. suggest that these are closely related to other congeners that infect haplochromines. We also found morphological indications that representatives of Cichlidogyrus colonised Lake Victoria haplochromines or their ancestors at least twice, which is in line with the Lake Victoria superflock being colonized by two cichlid tribes (Haplochromini and Oreochromini).DisclaimerThis preprint is disclaimed for purposes of Zoological Nomenclature in accordance with the International Code of Zoological Nomenclature, Fourth Edition Articles 8.2 and 8.3 (ICZN 1999). No new names or nomenclatural changes are available from statements in this preprint.Résumé - Quatre espèces nouvelles de Cichlidogyrus (Platyhelminthes, Monogenea, Dactylogyridae) parasites d’haplochrominé (Cichlidae) du lac Victoria, avec la redescription de C. bifurcatus and C. longipenisA cause des radiations adaptatives qu’ils ont subies, les cichlidés africain sont des systèmes modèles pour étudier l’évolution, mais aussi les relations hôtes/parasites, car ils hébergent de nombreuses espèces de Monogènes parasites qui présentent une spécificité étroite vis-à-vis de leurs hôtes. Dans ce travail, nous avons échantillonné cinq localités dans le Sud du lac Victoria, le plus jeune des grands lacs d’Afrique de l’Est. Nous avons examiné les Monogènes présents sur les branchies de 18 espèces de Cichlidés appartenant à la radiation adaptative « superflock » du lac Victoria et de deux espèces représentant deux lignées anciennes et non étroitement apparentées. Nous avons trouvé une espèce de Gyrodactylus (Gyrodactylidae, Monogenea), Gyrodactylus sturmbaueri Vanhove, Snoeks, Volckaert & Huyse, 2011 et sept espèces de Cichlidogyrus (Dactylogyridae, Monogenea). Quatre espèces nouvelles sont décrites dans le présent travail : Cichlidogyrus pseudodossoui n. sp., C. nyanza n. sp., C. furu n. sp., C. vetusmolendarius n. sp.. Une est signalée mais non décrite formellement (trop peux d’individus recueillis, morphologiquement proche de C. furu n. sp.). Deux autres sont redécrites : Cichlidogyrus bifurcatus Paperna, 1960 and C. longipenis Paperna & Thurston, 1969. Nos résultats confirment que la faune des Monogènes des Cichlidés du lac Victoria fait preuve d’une richesse spécifique et d’une spécificité moins importante que celle du lac Tanganyika. Chez C. furu n. sp. la paire de crochet V étant nettement plus longue que les autres, il faudra reconsidérer le système de classification actuel des types de hapteurs chez les Cichlidogyrus, qui considère que tous les crochets (III à VII) ont la même taille. Quelques caractéristiques morphologiques de C. bifurcatus, C. longipenis et C. nyanza n. sp. pourraient être la preuve d’une ascendance commune avec des congénères présents chez d’autres Haplochrominés. De même, certains caractères indiqueraient que des représentants des Cichlidogyrus ont colonisé les Haplochrominés du lac Victoria, ou leurs ancêtres, au moins à deux reprises, ce qui est cohérent avec une colonisation du lac par deux lignées de cichlidés distinctes (Haplochromini and Oreochromini).

Evolution of regulatory networks associated with traits under selection in cichlids

Background Seminal studies of vertebrate protein evolution speculated that gene regulatory changes can drive anatomical innovations. However, very little is known about gene regulatory network (GRN) evolution associated with phenotypic effect across ecologically diverse species. Here we use a novel approach for comparative GRN analysis in vertebrate species to study GRN evolution in representative species of the most striking examples of adaptive radiations, the East African cichlids. We previously demonstrated how the explosive phenotypic diversification of East African cichlids can be attributed to diverse molecular mechanisms, including accelerated regulatory sequence evolution and gene expression divergence. Results To investigate these mechanisms across species at a genome-wide scale, we develop a novel computational pipeline that predicts regulators for co-extant and ancestral co-expression modules along a phylogeny, and candidate regulatory regions associated with traits under selection in cichlids. As a case study, we apply our approach to a well-studied adaptive trait—the visual system—for which we report striking cases of network rewiring for visual opsin genes, identify discrete regulatory variants, and investigate their association with cichlid visual system evolution. In regulatory regions of visual opsin genes, in vitro assays confirm that transcription factor binding site mutations disrupt regulatory edges across species and segregate according to lake species phylogeny and ecology, suggesting GRN rewiring in radiating cichlids. Conclusions Our approach reveals numerous novel potential candidate regulators and regulatory regions across cichlid genomes, including some novel and some previously reported associations to known adaptive evolutionary traits.

Molecular Plasticity in Animal Pigmentation: Emerging Processes Underlying Color Changes

Synopsis Animal coloration has been rigorously studied and has provided morphological implications for fitness with influences over social behavior, predator–prey interactions, and sexual selection. In vertebrates, its study has developed our understanding across diverse fields ranging from behavior to molecular biology. In the search for underlying molecular mechanisms, many have taken advantage of pedigree-based and genome-wide association screens to reveal the genetic architecture responsible for pattern variation that occurs in early development. However, genetic differences do not provide a full picture of the dynamic changes in coloration that are most prevalent across vertebrates at the molecular level. Changes in coloration that occur in adulthood via phenotypic plasticity rely on various social, visual, and dietary cues independent of genetic variation. Here, I will review the contributions of pigment cell biology to animal color changes and recent studies describing their molecular underpinnings and function. In this regard, conserved epigenetic processes such as DNA methylation play a role in lending plasticity to gene regulation as it relates to chromatophore function. Lastly, I will present African cichlids as emerging models for the study of pigmentation and molecular plasticity for animal color changes. I posit that these processes, in a dialog with environmental stimuli, are important regulators of variation and the selective advantages that accompany a change in coloration for vertebrate animals.

Comparing speciation rates in lakes, rivers, and the sea

ABSTRACTThe diversity of species inhabiting freshwater relative to marine habitats is striking, given that freshwater habitats make up less than 1% of Earth’s water. The most commonly proposed mechanism for this pattern is that freshwater habitats are more fragmented than marine habitats, resulting in increased opportunities for allopatric speciation in freshwater and thus increased speciation rates relative to marine habitats. However, some authors suggest that speciation is faster in sympatry than allopatry, as illustrated by lacustrine radiations such as African cichlids. Sympatric speciation may be common in lakes, while allopatric speciation may be dominant in rivers. If allopatric speciation is inherently slower than sympatric speciation, we might expect differences in rates of speciation between lineages primarily confined to riverine versus lacustrine habitats. These differences within freshwater habitats may be important to consider when comparing diversification rates between freshwater and marine lineages. Here I compared rates of speciation associated with ray-finned fishes in marine, riverine and lacustrine habitats. I also used geographic range data to compare rates of speciation associated with allopatric and sympatric speciation modes in freshwater fishes. I found that lakes had faster rates of speciation than other aquatic habitats. However, most freshwater fish diversity arose through allopatric speciation in rivers, despite their slower speciation rate. Speciation rates were more strongly correlated with habitat (riverine or lacustrine) than present-day range overlap (allopatry or sympatry). Surprisingly, riverine and marine habitats had similar rates of speciation. Models of evolution suggest that lacustrine habitats are evolutionary unstable, and biased transition rates explain the dearth of lacustrine species. Collectively, these results suggest that while allopatric speciation is important for generating freshwater diversity, this mechanism may not be the cause of faster speciation rates between freshwater and marine habitats.