Genomic architecture of ecologically divergent body shape in a pair of sympatric crater lake cichlid fishes

Determining the genetic bases of adaptations and their roles in speciation is a prominent issue in evolutionary biology. Cichlid fish species flocks are a prime example of recent rapid radiations, often associated with adaptive phenotypic divergence from a common ancestor within a short period of time. In several radiations of freshwater fishes, divergence in ecomorphological traits - including body shape, colour, lips and jaws - is thought to underlie their ecological differentiation, specialization and, ultimately, speciation. The Midas cichlid species complex (Amphilophus spp.) of Nicaragua provides one of the few known examples of sympatric speciation where species have rapidly evolved different but parallel morphologies in young crater lakes. This study identified significant QTL for body shape using SNPs generated via ddRAD sequencing and geometric morphometric analyses of a cross between two ecologically and morphologically divergent, sympatric cichlid species endemic to crater Lake Apoyo: an elongated limnetic species (Amphilophus zaliosus) and a high-bodied benthic species (Amphilophus astorquii). A total of 453 genome-wide informative SNPs were identified in 240 F-2 hybrids. These markers were used to construct a genetic map in which 25 linkage groups were resolved. Seventy-two segregating SNPs were linked to 11 QTL. By annotating the two most highly supported QTL-linked genomic regions, genes that might contribute to divergence in body shape along the benthic-limnetic axis in Midas cichlid sympatric adaptive radiations were identified. These results suggest that few genomic regions of large effect contribute to early stage divergence in Midas cichlids.

Shaping development through mechanical strain: The transcriptional basis of diet-induced phenotypic plasticity in a cichlid fish

Adaptive phenotypic plasticity, the ability of an organism to change its phenotype to match local environments, is increasingly recognized for its contribution to evolution. However, few empirical studies have explored the molecular basis of plastic traits. The East African cichlid fish Astatoreochromis alluaudi displays adaptive phenotypic plasticity in its pharyngeal jaw apparatus, a structure that is widely seen as an evolutionary key innovation that has contributed to the remarkable diversity of cichlid fishes. It has previously been shown that in response to different diets, the pharyngeal jaws change their size, shape and dentition: hard diets induce an adaptive robust molariform tooth phenotype with short jaws and strong internal bone structures, while soft diets induce a gracile papilliform tooth phenotype with elongated jaws and slender internal bone structures. To gain insight into the molecular underpinnings of these adaptations and enable future investigations of the role that phenotypic plasticity plays during the formation of adaptive radiations, the transcriptomes of the two divergent jaw phenotypes were examined. Our study identified a total of 187 genes whose expression differs in response to hard and soft diets, including immediate early genes, extracellular matrix genes and inflammatory factors. Transcriptome results are interpreted in light of expression of candidate genesmarkers for tooth size and shape, bone cells and mechanically sensitive pathways. This study opens up new avenues of research at new levels of biological organization into the roles of phenotypic plasticity during speciation and radiation of cichlid fishes.

The gut microbial community of midas cichlid fish in repeatedly evolved limnetic-benthic species pairs

Gut bacterial communities are now known to influence a range of fitness related aspects of organisms. But how different the microbial community is in closely related species, and if these differences can be interpreted as adaptive is still unclear. In this study we compared microbial communities in two sets of closely related sympatric crater lake cichlid fish species pairs that show similar adaptations along the limnetic-benthic axis. The gut microbial community composition differs in the species pair inhabiting the older of two crater lakes. One major difference, relative to other fish, is that in these cichlids that live in hypersaline crater lakes, the microbial community is largely made up of Oceanospirillales (52.28%) which are halotolerant or halophilic bacteria. This analysis opens up further avenues to identify candidate symbiotic or co-evolved bacteria playing a role in adaptation to similar diets and life-styles or even have a role in speciation. Future functional and phylosymbiotic analyses might help to address these issues.

Mortality estimates of the four major Cichlid fishes of Umuoseriche Lake, Imo State, Nigeria

The mortality of the four major cichlid fishes of Urnuoseriche Lake is the subject of this paper. Mortality I as estimated by five techniques, vary amongst the cichlid fishes, viz, Tilapia carbrae, Tilapia mariac, Tilapia zilli cend (hrornoditilapfa guntheri. The highest mortality rate was recorded for T mariac where the total mortality (Z) was 2.06; and natural mortality (M) was 1.8949. This species was also the most highly exploited species of fish with an exploitation ratio of0.566 (56.6%) and exploitation rate of 0.494. The least exploited cichlid fish is (. gun/hen where an exploitation ratio of 0.43209%) and exploitation rate of 0.2225 was recorded. In C'. guntheni, total mortality was 0.726 and natural mortality was 0.413 1. In T zilli, total mortality was 1.0547 wile exploitation ratio was 0.3674 (3 6.74%) and an exploitation rate was 0.2394. In T cahrae. total mortality was 1.8662: exploitation ratio was 0.4786 with an exploitation rate of 0.4045. (7 page document)

The distribution and feeding habits of a cichlid fish Oreochromis niloticus (Linnaeus) in Opa Reservoir, Ile-Ife

The distribution of Oreochromis niloticus was studied in Opa reservoir (Nigeria) using a graded set of gillnets while the food and feeding habits were studied using a castnet to collect the fish samples. About 90% of the fish specimens were caught near the reservoir bottom while about 69% of the specimens were caught within the inshore area of the reservoir. The species fed mainly on detritus, algae and higher plants. Feeding rhythm in O. niloticus started around 6.00 a.m. and reached a peak by 3.00 p.m. but then declined gradually until 6.00 p.m. These results can be utilized for the proper management of the fish species in the reservoir

Recent research in the African Great Lakes: fisheries, biodiversity and cichlid evolution

The East African Great Lakes are now well known for (1) their fisheries, of vital importance for their rapidly rising riparian human populations, and (2) as biodiversity hotspots with spectacular endemic faunas, of which the flocks of cichlid fishes unique to each of the three largest lakes, Tanganyika, Malawi and Victoria, offer unique opportunities to investigate how new species evolve and coexist. Since the early 1990s research involving over a hundred scientists, financed by many international bodies, has produced numerous reports and publications in widely scattered journals. This article summarizes their main discoveries and examines the status of, and prospects for, the fisheries, as well as current ideas on how their rich endemic fish faunas have evolved. It first considers fisheries projects in each of the three lakes: the deep rift valley lakes Tanganyika and Malawi and the huge Victoria, all of which share their waters between several East African countries. Secondly it considers the biodiversity surveys of each lake, based on underwater (SCUBA) observations of fish ecology and behaviour which have revealed threats to their fish faunas, and considers what conservation measures are needed. Thirdly, using the lakes as laboratories, what have the international investigations (including DNA techniques and follow-up aquarium experiments) now revealed about the origins and relationships of their cichlid species flocks and mechanisms of evolution?

Age, growth, and mortality of the Mayan cichlid (Cichlasoma urophthalmus) from the southeastern Everglades

Mayan cichlids (Cichlasoma urophthalmus) were collected monthly from March 1996 to October 1997 with hook-and-line gear at Taylor River, Florida, an area within the Crocodile Sanctuary of Everglades National Park, where human activities such as fishing are prohibited. Fish were aged by examining thin-sectioned otoliths, and past size-at-age information was generated by using back-calculation techniques. Marginal increment analysis showed that opaque growth zones were annuli deposited between January and May. The size of age-1 fish was estimated to be 33–66 mm standard length (mean=45.5 mm) and was supported by monthly length-frequency data of young-of-year fish collected with drop traps over a seven-year period. Mayan cichlids up to seven years old were observed. Male cichlids grew slower but achieved a larger size than females. Growth was asymptotic and was modeled by the von Bertalanffy growth equation Lt=263.6(1–exp[–0.166(t–0.001)]) for males (r2=0.82, n=581) and Lt=215.6 (1–exp[–0.197(t–0.058)]) for females (r2= 0.77, n=639). Separate estimates of total annual mortality were relatively consistent (0.44–0.60) and indicated moderate mortality at higher age classes, even in the absence of fishing mortality. Our data indicated that Mayan cichlids grow slower and live longer in Florida than previously reported from native Mexican habitats. Because the growth of Mayan cichlids in Florida periodically slowed and thus produced visible annuli, it may be possible to age introduced populations of other subtropical and tropical cichlids in a similar way.

The nature of the aerobic gastrointestinal bacteria of cichlid fish Sarotherodon mossambicus (Peters) and Tilapia nilotica (Linnaeus) grown under captivity

Bacteriological examination of the gastrointestinal microflora of 2 freshwater cichlid fish species (Sarotherodon mossambicus and Tilapia nilotica ) was performed, resulting in the bacteria enumeration of total viable counts of 1.06x10⁷/g and 7.75x10⁷/g of gastro-bacteria intestinal tract plus contents (wet weight) respectively, by aerobic incubation at 30+1°C. The majority (78%) of the total gut isolates from both fish species was Gram positive mesophilic which is characteristic of the higher ambient temperature in the tropics. These isolates were fastidious in their nutritional requirements and together with the rest are isogenous to bacteria autochthonous to soil and water. The occurrence of such organisms is attributed to the feeding habits of these fish. The gastrointestinal bacteria isolated in this study are transient residents but not "indigenous" in these cichlid fish.

Scale characteristics in relation to size in cichlid fish, Oreochromis mossambicus

Teleostean scale characteristics like length, width, number of circuli, primary radii, secondary radii and ctenii were studied to delineate its relationship with total length in a euryhaline fish, Oreochromis mossambicus (perciformes: cichlidae). Statistical analysis proved a positive correlation between the scale characteristics and fish size. The characteristics of midlateral scales indicated that these scales appeared first during squamation chronology. Further observations also indicated that differences in salinity could modify the morphometries of scales.

The biology, ecology, distribution and conservation of some surviving native non-cichlid fish species of Lakes Victoria, Kyoga and Nabugabo

Prior to introduction of non-native fish species into Lakes Victor i a, Kyoga and Nabugabo, the three lakes suppor ted diverse fish fauna representing 13 families consisting of six cichlid genera and fifteen non-cichlid genera. There were about 50 non-cichlid species and over 300 cichlids consisting of mainly haplochromines (Graham 1929, worthington 1929, Greenwood 1960). Many of the species were commercially and scientifically important and provided a rich variety of protein source to choose from. Following introduction of the Nile perch and several tilapiines species, most of the native species were drastically reduced and some have apparently disappeared. The few remaining species appear to be restricted in distribution due to the presence of the Nile perch. They are mainly confined to refugia such as marginal macrophytes, rocky outcrops and small satellite lakes which are separated from the areas of introduction by swamps

Comparative electrophoretic patterns of lactase dehydrogenase and malate dehydrogenase in five Lake Victoria cichlid species

Biochemical techniques designed to compare species on the basis of protein differences were started by NUTTALL (1904) who used immunological methods to compare the serum of humans with that of other primates. Since then more refined techniques have led to better results at the protein level in taxonomy, The analyses of proteins are considered to be the simplest indirect approach to understanding the structure and function of the genetic material, deoxyribonucleic acid (DNA). Interest in these analyses arises because of the close relationship between protein structure and gene structure. Thus by comparing the properties of homologous proteins from different taxa one is in essence comparins their genes (GORMAN er al., 1971). It is now an established fact that genetic information coded in molecules of DNA is translated through a series of reactions in the structure of proteins which form the principal morphological units of the animal body at the molecular level of organization (SIBLEY, 1952). A convenient method of comparing molecular differences between species is to measure the electrophoretic mobility of proteins in a starch gel medium (ASPINWALL and TSUYUKI, 1968) or acrylamide gel (RAYMOND and WEINTRAUB, 1959; BOUCK and BALL, 1968). Proteins with enzymatic properties can be compared on the basis of catalytic activity in the presence or absence of inhibitors (KAPLAN et al., 1959); BAILEY et al., t 1970). A combination of gel electrophoresis and histochemical enzyme detection techniques (HUNTER and MARKERT, 1957) makes it possible to combine electrophoretic mobility anti catalytic activity comparison, Enzyme patterns exhibited in starch gel or acrylamide gel have been used to classify different species. BOUCK and BALL (1968)working with lactate dehydrogenase in species of Trout found that each Trout species had LDH pattern characterbtic of that species. ASPINIWALL and TSUYUKI (1968) used muscle protein electrophoretic patterns to identify hybrid fishes. TSUYUKI and ROBERTS (1963) and TSUYUKI et al. (1964-65) found that myogen protein patterns in fishes were species specific. The myogen patterns within one family were remarkably parallel with the existing morphometric classification and these patterns constituted a single criterion by which the fishes could be identified. The fish used in these investigations were collected from shallow waters (10 metres) of Lake Victoria in two areas, Jinja and Kisumu, using gillnets and beach-seines. The study included ten specimens of each of the following specIes: (l) Haplochromis michaeli (2) Haploehromis obems (3) Astatoreochromis ulluaudi (4) Tilapia zillii and (5) Tilapia nilotica.

Probing aggressive motivation in a cichlid fish

The duration of startles provides an inverse measure of motivation to resume the previous activity. Here, we use a novel method in which one convict cichlid fish (Amatitlania nigrofasciata) of a competing pair was startled independently of the opponent. Fish were given various opponents and the mean startle duration determined. This mean was negatively correlated with the mean use of highly escalated 'frontal activities' such as biting and frontal display, but not the less escalated lateral displays or tail beating. Thus the startle duration was a reliable surrogate measure of the most escalated components of aggressive interactions. That is, it provided a motivational probe for aggressiveness of individual fish. Fight motivation is often determined in terms of fight duration or physiological costs for losers, who reveal the costs they are prepared to pay. We discuss various potential advantages of the motivational probe over previous measures, particularly with respect to winners and losers and different times during the interactions.

Rapid sympatric ecological differentiation of crater lake cichlid fishes within historic times

Background: After a volcano erupts, a lake may form in the cooled crater and become an isolated aquatic ecosystem. This makes fishes in crater lakes informative for understanding sympatric evolution and ecological diversification in barren environments. From a geological and limnological perspective, such research offers insight about the process of crater lake ecosystem establishment and speciation. In the present study we use genetic and coalescence approaches to infer the colonization history of Midas cichlid fishes (Amphilophus cf. citrinellus) that inhabit a very young crater lake in Nicaragua-the ca. 1800 year-old Lake Apoyeque. This lake holds two sympatric, endemic morphs of Midas cichlid: one with large, hypertrophied lips (~20% of the total population) and another with thin lips. Here we test the associated ecological, morphological and genetic diversification of these two morphs and their potential to represent incipient speciation.
Results: Gene coalescence analyses [11 microsatellite loci and mitochondrial DNA (mtDNA) sequences] suggest that crater lake Apoyeque was colonized in a single event from the large neighbouring great lake Managua only about 100 years ago. This founding in historic times is also reflected in the extremely low nuclear and mitochondrial genetic diversity in Apoyeque. We found that sympatric adult thin- and thick-lipped fishes occupy distinct ecological trophic niches. Diet, body shape, head width, pharyngeal jaw size and shape and stable isotope values all differ significantly between the two lip-morphs. The eco-morphological features pharyngeal jaw shape, body shape, stomach contents and stable isotopes (d15N) all show a bimodal distribution of traits, which is compatible with the expectations of an initial stage of ecological speciation under disruptive selection. Genetic differentiation between the thin- and thick-lipped population is weak at mtDNA sequence (FST = 0.018) and absent at nuclear microsatellite loci (FST < 0.001).
Conclusions: This study provides empirical evidence of eco-morphological differentiation occurring very quickly after the colonization of a new and vacant habitat. Exceptionally low levels of neutral genetic diversity and inference from coalescence indicates that the Midas cichlid population in Apoyeque is much younger (ca. 100 years or generations old) than the crater itself (ca. 1 800 years old). This suggests either that the crater remained empty for many hundreds of years after its formation or that remnant volcanic activity prevented the establishment of a stable fish population during the early life of the crater lake. Based on our findings of eco-morphological variation in the Apoyeque Midas cichlids, and known patterns of adaptation in Midas cichlids in general, we suggest that this population may be in a very early stage of speciation (incipient species), promoted by disruptive selection and ecological diversification.