Decoupled jaws promote trophic diversity in cichlid fishes

Functional decoupling of oral and pharyngeal jaws is widely considered to have expanded the ecological repertoire of cichlid fishes. But, the degree to which the evolution of these jaw systems is decoupled and whether decoupling has impacted trophic diversification remains unknown. Focusing on the large Neotropical radiation of cichlids, we ask whether oral and pharyngeal jaw evolution is correlated and how their evolutionary rates respond to feeding ecology. In support of decoupling, we find relaxed evolutionary integration between the two jaw systems, resulting in novel trait combinations that potentially facilitate feeding mode diversification. These outcomes are made possible by escaping the mechanical trade-off between force transmission and mobility, which characterizes a single jaw system that functions in isolation. In spite of the structural independence of the two jaw systems, results using a Bayesian, state-dependent, relaxed-clock model of multivariate Brownian motion indicate strongly aligned evolutionary responses to feeding ecology. So, although decoupling of prey capture and processing functions released constraints on jaw evolution and promoted trophic diversity in cichlids, the natural diversity of consumed prey has also induced a moderate degree of evolutionary integration between the jaw systems, reminiscent of the original mechanical trade-off between force and mobility.     

闽江口鱼类功能多样性

根据2016年春季(5月)、夏季(8月)、秋季(11月)和冬季(2月)在闽江口海域进行底拖网作业的渔业资源调查样品,应用鱼类形态功能特征点数据分析了福建闽江口海域的鱼类功能多样性现状及时空变异.结果表明: 功能多样性与物种丰度具有相同的变化趋势,各站位鱼类物种丰度分别为:春季(17.8±5.1);夏季(22.4±5.6);秋季(17.7±6.3);冬季(12.1±2.3).鱼类集群功能多样性指数夏季最高,为(33.94±28.70),冬季最低(9.93±8.83),春季为(11.30±7.55),秋季为(19.05±19.32).群落内功能种呈现显著的季节演替,春季为太的黄鲫(占比为26%),夏季为六指马鲅(26%),秋季为龙头鱼,占比高达69%,显著高于其他物种的总和,冬季为棘头梅童鱼(38%).鱼类功能的季节性变动主要受其生活史及与其他物种之间的种间关系影响.     

淡水鱼类功能多样性及其研究方法

目前,群落功能多样性备受生态学界关注,被认为是能解决生态问题的一种重要途径。我国对于群落功能多样性主要集中在植物群落和微生物群落,而在鱼类群落方面的研究几乎是空白。我国鱼类资源正面临着严重威胁,包括水坝建设导致的鱼类通道受阻、水库形成造成鱼类产卵场功能消失、过度捕捞、水质恶化和富营养化加重、外来种入侵等因素,导致渔业资源急剧衰退,水生生态系统功能下降。以淡水鱼类群落为例,对鱼类功能多样性的数据获取及处理分析与评价、测定指标及计算方法与研究难点等进行综述,以期为鱼类资源保护提供新的理论依据和切入点。     

Genetic interactions controlling sex and color establish the potential for sexual conflict in Lake Malawi cichlid fishes

Sex-determining systems may evolve rapidly and contribute to lineage diversification. In fact, recent work has suggested an integral role of sex chromosome evolution in models of speciation. We use quantitative trait loci analysis of restriction site-associated DNA -tag single nucleotide polymorphisms to identify multiple loci responsible for sex determination and reproductively adaptive color phenotypes in Lake Malawi cichlids. We detect a complex epistatic sex system consisting of a major female heterogametic ZW locus on chromosome 5, two separate male heterogametic XY loci on chromosome 7, and two additional interacting loci on chromosomes 3 and 20. Our data support the known chromosomal linkage between orange blotch color and ZW, as well as novel genetic associations between male blue nuptial color and two sex determining regions (an XY and ZW locus). These results provide further empirical evidence for a complex antagonistic sex–color system in this species flock and suggest a possible role for, and effect of, polygenic sex-determining systems in rapid evolutionary diversification.     

Beta diversity of rock-restricted cichlid fishes in Lake Malawi: importance of environmental and spatial factors

The rock-restricted cichlid fish assemblages of Lake Malawi exhibit high spatial diversity in their species composition and relative abundance. However the extent to which this is due to the effects of local environmental differences, dispersal limitation of constituent taxa, and the assignment of allopatric populations to species is uncertain. We examined the factors associated with diversity within an assemblage from the north-western shores, encompassing a spatial scale of 170 km. For both the whole assemblage, and all constituent species-complexes, spatial variance in community structure was significantly dependent upon both geographic distances between locations and local habitat variables. Pronounced effects of distance indicate limited dispersal, but our results also show that that the spatial variance explained by geographic distance alone was strongly linked to proportion of allopatric populations within a species-complex with species status. Thus, the taxonomic status of allopatric populations underlies, at least partially, the biogeographical structure of this assemblage. Substrate composition and habitat depth were also significant determinants of community structure, although spatial variance attributed to these variables was less than that associated with distance alone. Substantial unexplained variance may be a consequence of the effects of unmeasured habitat variables, high ecological similarity between co-occurring species, stochastic influences on population abundance, and the effects of local adaptation. Despite low spatial variance explained by the assessed environmental variables, significant environmental influence on cichlid assemblage structure across a wide spatial scale indicates that even slight future environmental changes may have the capacity to significantly alter species composition.     

Adaptive sequence evolution in a color gene involved in the formation of the characteristic egg-dummies of male haplochromine cichlid fishes

Background  

The exceptionally diverse species flocks of cichlid fishes in East Africa are prime examples of parallel adaptive radiations. About 80% of East Africa's more than 1 800 endemic cichlid species, and all species of the flocks of Lakes Victoria and Malawi, belong to a particularly rapidly evolving lineage, the haplochromines. One characteristic feature of the haplochromines is their possession of egg-dummies on the males' anal fins. These egg-spots mimic real eggs and play an important role in the mating system of these maternal mouthbrooding fish.     

Captive breeding for the conservation of cichlid fishes

Cichlid fishes are by far the largest familial group of endangered vertebrates, especially the haplochromines. This paper concerns the organization and management of captive breeding of haplochromine cichlids. The setting up of a small-scale laboratory programme for the conservation of endangered species is described in terms of funding, staffing, installation and livestock husbandry. Breeding is discussed in the context of the selection of broodstock, basic reproductive biology, rearing, disease, pathological disorders and the arrangements necessary for the transfer and documentation of progeny. There are already indications in Africa and elsewhere that the dramatic decline and demise of cichlid taxa in Lake Victoria will not be an isolated phenomenon. There is no prospect that the captive breeding of cichlids can alone resolve such large-scale problems in fisheries management and ecology, or prevent the loss of taxa in nature. Nevertheless, captive breeding provides conservation options which are otherwise limited or unavailable regarding the saving of individual 'heritage' species, restocking to the wild, fundamental laboratory research and, finally, public education on the grave issue of actual and prospective mass extinctions of cichlids and other rare fishes.     

Introgressive hybridization between color morphs in a population of cichlid fishes twelve years after human-induced secondary admixis

In the extremely species-rich haplochromine cichlid fishes of the East African Great Lakes, prezygotic isolation between closely related species is often maintained by color-assortative mating. In 1998, local fisherman working for the ornamental fish trade released different color morphs of the cichlid genus Tropheus into a small harbor basin in the southern part of Lake Tanganyika. This artificial amalgamation of color morphs provides a unique possibility to study mating patterns in cichlids in a natural environment over time. In a precursor study, we analyzed genotypes and phenotypes of almost 500 individuals sampled between 1999 and 2001 and uncovered a marked degree of color-assortative mating, which depended on the level of color pattern dissimilarity between morphs. Twelve years after introduction of nonindigenous morphs, we again sampled Tropheus individuals from the harbor basin and an adjacent, originally pure population and analyzed phenotypes (coloration) and genotypes (mitochondrial control region and 9 microsatellite loci) to assess the current status of the admixed population. Principal component analyses of color score data and population assignment tests demonstrate an increasing level of introgressive hybridization between morphs but also some ongoing color-assortative mating within morphs. The observed mating pattern might have been influenced by fluctuating environmental conditions such as periodic algal blooms or increased sedimentation causing turbid conditions in an otherwise clear lake.     

Age and spread of the haplochromine cichlid fishes in Africa

The Haplochromini are by far the most species-rich cichlid fish tribe that originated along with the so-called primary radiation of the Lake Tanganyika cichlid species flock, i.e. at the same time during which the majority of the endemic Lake Tanganyika cichlid tribes emerged. Unlike the other tribes, the haplochromines are not restricted to Lake Tanganyika but distributed throughout Africa, except for the northwestern part of the continent. Haplochromine cichlids seeded the adaptive radiation of cichlid fishes in Lakes Malawi, Kivu, Victoria, Turkana, as well as in the now extinct paleo-Lake Makgadikgadi. Here we present a comprehensive phylogenetic and phylogeographic analysis of haplochromine cichlids that is based upon DNA sequences of two mitochondrial gene segments of riverine taxa covering all major African biogeographic regions where haplochromines are found. Our analysis revealed that six lineages of haplochromines originated within a short period of time, about 5.3-4.4 MYA. These haplochromine lineages show a highly complex phylogeographic pattern, probably severely influenced by climate- and/or geology-induced changes of the environment, with river capture events most likely playing an important role for species dispersal.     

Secondary contact seeds phenotypic novelty in cichlid fishes

Theory proposes that genomic admixture between formerly reproductively isolated populations can generate phenotypic novelty for selection to act upon. Secondary contact may therefore be a significant promoter of phenotypic novelty that allows species to overcome environmental challenges and adapt to novel environments, including during adaptive radiation. To date, this has largely been considered from the perspective of interspecific hybridization at contact zones. However, it is also possible that this process occurs more commonly between natural populations of a single species, and thus its importance in adaptive evolution may have been underestimated. In this study, we tested the consequences of genomic introgression during apparent secondary contact between phenotypically similar lineages of the riverine cichlid fish Astatotilapia calliptera. We provide population genetic evidence of a secondary contact zone in the wild, and then demonstrate using mate-choice experiments that both lineages can reproduce together successfully in laboratory conditions. Finally, we show that genomically admixed individuals display extreme phenotypes not observed in the parental lineages. Collectively, the evidence shows that secondary contact can drive the evolution of phenotypic novelty, suggesting that pulses of secondary contact may repeatedly seed genetic novelty, which when coupled with ecological opportunity could promote rapid adaptive evolution in natural circumstances.     

Ecomorphological diversification and convergence in fluvial cichlid fishes

Synopsis We compared ecological and morphological patterns among cichlid faunas from three different biotic regions: the Río Tortuguero/Rio Sarapiquf in Costa Rica, the Río Apure drainage in Venezuela, and the Upper Zambezi drainage in Zambia. Cichlids comprise 19 percent of the fish fauna in the Tortuguero and Upper Zambezi drainages and 6.5 percent in the Apure drainage. Cichlid faunas exhibited similar patterns of habitat and food resource utilization, although vegetation-dwelling is more common and detritivory and herbivory are rarer in the Apure fauna. We hypothesize that South American ostariophysan fishes were more preadapted than cichlids to exploit detritivore and herbivore niches. The Zambezi cichlid fauna shows less ecomorphological diversification than the other two faunas, even though the degree of dietary diversification is similar among faunas. Chaetobranchus flavescens from the Venezuelan fauna is the only species that specializes on zooplankton as an adult, and algae grazing (Neetroplus nematopus) and specialized fruit feeding (Cichlasoma tuba) were represented only in the Costa Rican fauna. Based on the most recent hypothesized phylogeny for the family Cichlidae, we identified numerous interfaunal ecomorphological and feeding niche convergences. Patterns of ecomorphological divergence in relation to cladogenesis indicate a faster rate of evolutionary niche diversification in Central American cichlids compared with the two other faunas.     

Feeding schools among Lake Malawi cichlid fishes

Synopsis Seven species of rock-frequenting cichlids, endemic to Lake Malawi, formed feeding-schools. Petrotilapia genalutea was the nuclear species comprising 92% of the school members (n = 2690 individuals from 50 schools). Fifty-eight per cent of all schools encountered consisted only of P. genalutea. School size ranged from 10 to 300 members (x = 81 ± 74 SD, median = 62). Schooling behaviour was facultative. Ninety-seven per cent of the feeding sites used by the schools occurred within the territories of highly aggressive, territorial species. These sites support considerably greater standing crops of algae, the food of the schooling species, than undefended sites. Individuals of schooling species were unable to gain access to the aggresively defended areas unless they joined schools.     

Food resources of shrimp-eating cichlid fishes in Lake Tanganyika

Species, size composition, and diurnal variation in abundance of shrimp (Atyidae) were examined and compared with food habits of 14 species of shrimp-eating cichlids in Lake Tanganyika. In the daytime when the cichlids foraged actively, only one species of shrimp, Limnocaridina latipes, was present on rock surfaces. Eleven of the 14 species of shrimp-eaters examined preyed mainly on L. latipes; the shrimp-eaters foraging only on rock surfaces preyed exclusively on L. latipes, whereas shrimp-eaters also taking foods from the interstices between or under stones and debris preyed on various species of shrimp. Although three kinds of shrimp-eaters, each with a unique prey catching technique and preferred shrimp size, were recognized, all shrimp-eaters preyed on shrimp of greatly overlapping sizes. This fact confirms that partitioning of food resources takes place via different capturing techniques. Moreover, the utilization of the same food resource in a different manner may promote group foraging in some shrimp-eaters, because the shrimp is a suitable animal to hunt in groups and is a quickly renewable food of potential abundance.     

Correlated evolution of body and fin morphology in the cichlid fishes

Body and fin shapes are chief determinants of swimming performance in fishes. Different configurations of body and fin shapes can suit different locomotor specializations. The success of any configuration is dependent upon the hydrodynamic interactions between body and fins. Despite the importance of body–fin interactions for swimming, there are few data indicating whether body and fin configurations evolve in concert, or whether these structures vary independently. The cichlid fishes are a diverse family whose well‐studied phylogenetic relationships make them ideal for the study of macroevolution of ecomorphology. This study measured body, and caudal and median fin morphology from radiographs of 131 cichlid genera, using morphometrics and phylogenetic comparative methods to determine whether these traits exhibit correlated evolution. Partial least squares canonical analysis revealed that body, caudal fin, dorsal fin, and anal fin shapes all exhibited strong correlated evolution consistent with locomotor ecomorphology. Major patterns included the evolution of deep body profiles with long fins, suggestive of maneuvering specialization; and the evolution of narrow, elongate caudal peduncles with concave tails, a combination that characterizes economical cruisers. These results demonstrate that body shape evolution does not occur independently of other traits, but among a suite of other morphological changes that augment locomotor specialization.     

Genome mapping of the orange blotch colour pattern in cichlid fishes

The dramatic variation of cichlid fish colour pattern is thought to function in mate choice, evolve by sexual selection, and contribute to explosive speciation. Here, we combine linkage mapping and population genetic analyses to identify a single region of the cichlid genome responsible for the orange blotch (OB) colour phenotype. In each analysis, OB is tightly linked to the c-ski1 gene. Additionally, we use comparative mapping information from the Takifugu rubripes and human genomes to suggest positional candidate loci for OB. Our work should engender a more comprehensive understanding of the molecular ecology of OB and its role in cichlid speciation. Moreover, we have assembled the components of a method to focus upon the genetic basis of evolutionarily and ecologically significant phenotypes.     

Alternative life-history styles of some African cichlid fishes

Synopsis About 50% of African cichlid species have been described. Little is known about cichlid ecology, behaviour or about the evolution and interactions of communities. Nevertheless, trends which provide an insight into cichlid life histories, into evolutionary alternatives and into behavioural alternatives which may be followed during the life time of these fishes are emerging. Cichlids which spend their entire life history in a single habitat belong to species flocks that have spectated greatly. In contrast, those groups which live in a variety of habitats during their life history have spectated little. Despite the trophic specializations that have occurred among cichlids, many and perhaps all species, have the ability to feed upon alternative food resources. They do so by switching or by markedly modifying their behaviour and all are particularly opportunistic. The extensive adaptive radiation of cichlids with regard to those attributes of morphology and behaviour that are associated with survival and growth, is apparently not matched by similar radiations in reproductive behaviour. The perceived evolutionary conservatism with regard to reproductive behaviour is attributed to the effects of stabilizing selection, but this still needs to be tested. The constraints of stabilizing selection appear to be lifted with regard to coloration. If colour is an important component of the specific-mate recognition system then it should also be subjected to stabilizing selection so the readiness with which colour variation occurs within and between populations needs explanation. Parental care in cichlids has followed a variety of evolutionary alternatives, all of which are geared to improve the chances of survival of offspring in their specific micro-habitats.     

On the evolutionary pathway of parental care in mouth-brooding cichlid fishes

Evolutionary theory predicts that differences in parental care patterns among species arose from interspecific differences in the costs and benefits of care for each sex. In Galilee St Peter''s fish, Sarotherodon galilaeus (Cichlidae), male care, female care and biparental care all occur in the same population. We exploit this unusual variability to isolate conditions favouring biparental versus uniparental mouth-brooding by males or females. We first review a game-theoretic model of parental care evolution, predictions of which we test experimentally in this paper. Manipulations of the operational sex ratio show that males and females desert their offspring more frequently when the costs of care are high (in terms of lost mating opportunities). Breeding trials with males of different sizes show that small fathers desert more frequently than large fathers. We attribute this to the associated difference in the fitness benefit of biparental care relative to female-only care. Our experimental results confirm that in St Peter''s fish the probability of caring is determined facultatively according to current conditions at each spawn. The experiments and model together suggest that interspecific variation in remating opportunities and clutch size may be responsible for differences in care patterns within the sub-family Tilapiini. Our results support the hypothesis that biparental mouth-brooding was the ancestral state of both male and female uniparental mouth-brooding in cichlid fishes.     

Evolutionary history of Lake Tanganyika's scale-eating cichlid fishes

Although Lake Tanganyika is not the most species-rich of the Great East African Lakes it comprises by far the greatest diversity of cichlid fishes in terms of morphology, ecology, and breeding styles. Our study focuses on the Tanganyikan cichlid tribe Perissodini, which exhibits one of the most peculiar feeding strategies found in cichlids-scale-eating. Their evolutionary history was reconstructed from 1416 bp DNA sequence of two mitochondrial genes (ND2 and partial control region) and from 612 AFLP markers. We confirm the inclusion of the zooplanktivorous genus Haplotaxodon in the tribe Perissodini, and species status of Haplotaxodon trifasciatus. Within the Perissodini, the major lineages emerged within a short period roughly 1.5-2 MYA, which makes their radiation slightly younger than that of other Tanganyikan cichlid tribes. Most scale-eaters evolved in deep-water habitat, perhaps associated with the previously documented radiations of other deep-water dwelling cichlid lineages, and colonized the shallow habitat only recently.