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Rudimentary structures are variable, which is most impressively demonstrated by the eyes and dark body pigmentation of species evolving in caves. As these can often still be crossed with their ancestral surface sister forms, variability is usually attributed to hybridization. For two invertebrate and two vertebrate cave species, I show here that in spite of genetic separation between surface and cave forms, variability of the rudimentary structures is nonetheless present. This probably results from the loss of stabilizing selection, because neutral deleterious mutations are no longer eliminated. In contrast, compensatory traits in caves do not differ from their surface forms with respect to variability, because both are subject to selection. In the phylogenetically young Astyanax cave populations, traits compensatory to cave life have achieved different stages of adaptation compared to the phylogenetically old populations. They may show the same degree of differentiation, be intermediate or do not differ from the surface ancestor. This is attributed to different intensities of selection. [ABSTRACT FROM AUTHOR]

Cave animals and species flocks exhibit common evolutionary principles. In caves, all traits dependent on the information derived from light lose their biological function. Mutations destructive for such traits, but neutral for the organism as a whole, can persist and accumulate until a trait has vanished. Adaptive radiations start in ecosystems containing open niches. Here, selection on niche-specific traits, such as the viscerocranium in fish, is relaxed owing to the absence of competing species, and viscerocranial variability arises. It is transitorily high in recent and phylogenetically younger flocks, providing new phenotypes. It lessens and is completely lost after directional selection promotes the fixation of phenotypes that are best adapted. In cave animals and species flocks, single traits manifest phenotypic variability owing to relaxed selection. Like the eye in cave species, the viscerocranium can be classified a module, the development of which is encoded in gene regulatory networks. Mutations in these genes can result in new phenotypes. Regarding functionality, these mutations might be destructive and eliminated by selection, neutral and thus persisting, or beneficial and promoted to fixation by directional selection. Given the ancient heritage of teleostean fish, these gene regulatory networks might be prone to mutations at the same loci or to developmental reactions resulting in similar phenotypes in closely related or taxonomically and geographically distant species. [ABSTRACT FROM AUTHOR]

The forces driving regression of biologically functionless traits remain disputed. There is ongoing debate regarding whether selection, as opposed to disuse and neutral mutations, is involved in this process. Cave species are of particular relevance for study in this regard because in continuous darkness all traits that depend on information from light, such as eyes, dark pigmentation and certain behaviours, abruptly lose their function. Recently, strong selection driving reduction has again been proposed, which relied on modelling analyses based on assumptions such as immigration of surface alleles into the cave forms or no fitness difference existing between Astyanax surface and cave fish. The validity of these assumptions, often applied to reject neutral processes in functionless traits, is questioned in this review. Morphological variation in a trait resulting from genetic variability is typical of biologically functionless traits and is particularly notable in phylogenetically young cave species. It is the most evident indicator of loss of selection, which normally enforces uniformity to guarantee optimal functionality. Phenotypic and genotypic variability in Astyanax cave fish eyes does not derive from genetic introgression by the surface form, but from regressive mutations not being eliminated by selection. This matches well with the principles of Kimura's neutral theory of molecular evolution. [ABSTRACT FROM AUTHOR]

Astyanax is a species-rich, non-monophyletic genus composed of several supraspecific taxa that are poorly delimited. The Astyanax fasciatus complex is one of these taxa and shows high taxonomic complexity. To elucidate the evolutionary history of the A. fasciatus complex from southern South America, we conducted cytogenetic, molecular and morphological analyses in specimens from the Uruguay River basin. Cytogenetic characters demonstrated two closely related operational taxonomic units: Astyanax sp. 1 (8 m +22 sm +10 st +6 a), Astyanax sp. 2 (8 m +24 sm +10 st +4 a) and natural hybrids (8 m +23 sm +8 st +5 a). 5S ribosomal DNA sites were found in two pairs of m chromosomes and one pair of a chromosomes in Astyanax sp. 1, two pairs of a chromosomes and one pair of m chromosomes in Astyanax sp. 2, and three m chromosomes and three a chromosomes in hybrids. As 51 sites were found in three chromosomes in Astyanax sp. 1 and in five chromosomes in Astyanax sp. 2 and hybrids. Mitochondrial sequence analyses did not separate the two units and hybrids. Morphological analyses revealed differences between Astyanax sp. 2 and hybrids. This secondary contact with gene flow between lineages that diverged long ago might slow or reverse the differentiation/speciation process. These results help us to understand the evolutionary history of this highly complex clade of Astyanax in southern South America. [ABSTRACT FROM AUTHOR]

Cave colonization offers a natural laboratory to study an extreme environmental shift, and diverse cave species from around the world often have converged on robust morphological, physiological and behavioural traits. The Mexican tetra (Astyanax mexicanus) has repeatedly colonized caves in the Sierra de El Abra and Sierra de Guatemala regions of north-east Mexico ~0.20–1 Mya, indicating an ability to adapt to the cave environment. The time frame for the evolution of these traits in any cave animal, however, is poorly understood. Astyanax mexicanus from the Río Grande in South Texas were brought to Central Texas beginning in the early 1900s and colonized underground environments. Here, we investigate whether phenotypic and behavioural differences have occurred rapidly between a surface population and a geographically proximate cave population, probably of recent origin. Fish from the cave and surface populations differ significantly in morphological traits, including coloration, lateral line expansion and dorsal fin placement. Striking behavioural shifts in aggression, feeding and wall-following have also occurred. Together, our results suggest that morphological and behavioural changes accompanying cave colonization can be established rapidly, and this system offers an exciting and unique opportunity for isolating the genetic and environmental contributions to colonization of extreme environments. [ABSTRACT FROM AUTHOR]

Similar phenotypic traits can evolve independently in closely related lineages when these lineages are exposed to similar selective pressures (i.e. parallel evolution), while unique histories (i.e. historical contingencies) can also shape phenotypic variation. Here, we addressed the importance of parallel evolution and historical contingency on the morphological variation in sympatric morphs (deep-body and elongate-body) of the genus Astyanax from two different systems (Lake Catemaco and Río San Juan) based on analysis of trophic ecomorphology and geometric morphometrics of body shape. Parallel evolution due to adaptive divergence associated with morphs better explained the variation in trophic ecomorphology, while body shape variation was better explained by historical contingencies. Related to trophic ecomorphology evolution, the elongate-body morph from both systems showed a longer snout, longer ascending process of premaxilla, fewer cuspids on the anterior tooth of premaxilla and acute premaxillary angles associated with parallel evolution (with opposite patterns in the deep-body morph). A higher degree of similarity in body shape between sympatric morphs than between morphs from allopatric systems accounted for the morphological evolution related to historical contingencies. This study highlights how parallel evolution and historical contingency differentially influenced several aspects of phenotype (trophic morphology and body shape, respectively) in the morphological evolution of the genus Astyanax. [ABSTRACT FROM AUTHOR]

Change in ecological conditions, as seen in surface and cave populations of Astyanax (Teleostei), has caused the divergent evolution of a large number of traits like eyes, coloration, taste, lateral line, and different kinds of behaviour like schooling, sleep or feeding posture. Because of the interfertility of surface and cave forms these fish are an exceptional object to study the morphological and genetic basis of the evolution of such complex regressive and constructive traits. Classical crossing analyses and genomic studies are contributing to growing understanding. Both kinds of traits mostly rely on multiple genetic bases and the phenotypic manifestation in the various crosses is similar. The gene effect underlying the phenotypic manifestation may exhibit an exponential increase at differing amounts in the various traits and crosses. Missing or presence of such genetic interaction helps determine whether the variability of eyes or pigmentation exhibited by Astyanax cave fish populations like Micos, is due to a more recent origin or to secondary hybridization with the surface fish. Neither crossing analysis nor QTL mapping revealed that eye reduction is pleiotropically antagonistically related to the increase of taste buds or lateral line sense. Independent inheritance of traits suggests that Astyanax cave fish are subjected to mosaic evolution. [ABSTRACT FROM AUTHOR]

Two hypotheses exist to explain ontogenetic eye reduction in Astyanax cave fish: first, after lens induction by the primordial eye cup, the lens plays the role of a central regulator of eye and retina regression or, second, the retina itself is an independent unit of eye development. A comparative study of five blind cave fish populations and their surface sister form was performed to investigate the differences of ontogenetic eye regression between the cave populations during different stages of development. The study revealed that, in addition to the initial formation of smaller primordia, eye regression is also caused during later ontogeny by different relative growth and specific histological characteristics. Whereas the cave fish lens never properly differentiates, the regressive process of the retina is transitorily interrupted by ongoing differentiation. In the newly-discovered Molino cave population, even visual cells with well-organized outer segments develop, which are secondarily reduced at a later ontogenetic stage. This result shows that the retina and lens are independent developmental units within the eye ball. Presumably, the genetic systems responsible for both show independent inheritance, which is also corroborated by hybrids of F2-crosses between the cave and surface fish, in which lens and retina development do not correlate. During ontogeny, the eye size differs between the cave populations. In Pachón cave fish, the relatively large eye size correlates with an ancient introgression from a surface population, which may have delayed eye regression. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 92, 287–296. [ABSTRACT FROM AUTHOR]

More than 20 populations of the cave-dwelling characid Astyanax occur within a restricted karst area in Mexico. The fish possess reduced eyes without lenses and visual cells. It is still an open question as to whether this condition evolved convergently after multiple entries of the surface ancestor into the different caves or whether a single cave ancestor, already characterized by reduced eyes, spread secondarily within them. In the crosses between specific populations, specimens appear that deviate considerably from those of the parents. They possess larger and better-developed eyes with histologically intact lenses and visual cells; they thus have the structural potential for vision. This indicates that in different cave populations, different mutations in the eye gene system have occurred. In cases where these non-functional rudimentary genes are recombined in hybrid specimens, gene expression may be restored. This is the result of separate evolution of several Astyanax cave populations. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 80, 545−554. [ABSTRACT FROM AUTHOR]