Genetic assimilation and the evolution of direction of genital asymmetry in anablepid fishes.

Phylogenetic comparative studies suggest that the direction of deviation from bilateral symmetry (sidedness) might evolve through genetic assimilation; however, the changes in sidedness inheritance remain largely unknown. We investigated the evolution of genital asymmetry in fish of the family Anablepidae, in which males' intromittent organ (the gonopodium, a modified anal fin) bends asymmetrically to the left or the right. In most species, males show a 1 : 1 ratio of left-to-right-sided gonopodia. However, we found that in three species left-sided males are significantly more abundant than right-sided ones. We mapped sidedness onto a new molecular phylogeny, finding that this left-sided bias likely evolved independently three times. Our breeding experiment in a species with an excess of left-sided males showed that sires produced more left-sided offspring independently of their own sidedness. We propose that sidedness might be inherited as a threshold trait, with different thresholds across species. This resolves the apparent paradox that, while there is evidence for the evolution of sidedness, commonly there is a lack of support for its heritability and no response to artificial selection. Focusing on the heritability of the left : right ratio of offspring, rather than on individual sidedness, is key for understanding how the direction of asymmetry becomes genetically assimilated.

An integrative approach method reveals the presence of a previously unreported species of Imparfinis Eigenmann and Norris 1900 (Siluriformes: Heptapteridae) in Argentina.

Specimens of Imparfinis were recently collected in north-western Argentina from the Bermejo River basin (Salta and Jujuy Provinces), del Valle River (Salta Province) and Horcones River (Santiago del Estero Province). An integrative approach to taxonomy, combining a detailed morphological study and molecular phylogenetic analyses, was applied to determine the species identity of these specimens. A principal components analysis of morphological data clustered the specimens from north-western Argentina and from the Amazon basin, indicating a close morphological resemblance. Also, a molecular phylogenetic analysis showed populations of I. guttatus from Argentina and Peru forming a clade. According to the conducted haplotype network analysis these populations are distinct in two mutations. Thus, in the absence of morphological or molecular data indicating the contrary, the combined method supports the identity of the specimens from the tributaries of the Paraguay River in Argentina as I. guttatus, whose type locality is in the upper Beni River basin in Bolivia. This contribution is also the first record for this species from Argentina. The disjunct distribution of I. guttatus provides new evidence reinforcing the hypothesis for the origin of the Paraguayan ichthyofauna. We also provide an approach to the phylogenetic relationships of Imparfinis in Heptapteridae.

The direction of genital asymmetry is expressed stochastically in internally fertilizing anablepid fishes.

Animal genitalia vary considerably across taxa, with divergence in many morphological traits, including striking departures from symmetry. Different mechanisms have been proposed to explain this diversity, mostly assuming that at least some of the phenotypic variation is heritable. However, heritability of the direction of genital asymmetry has been rarely determined. Anablepidae are internally fertilizing fish where the anal fin of males has been modified into an intromittent organ that transfers sperm into the gonopore of females. Males of anablepid fishes exhibit asymmetric genitalia, and both left- and right-sided individuals are commonly found at similar proportions within populations (i.e. antisymmetry). Although this polymorphism was described over a century ago, there have been no attempts to determine if genital asymmetry has a genetic basis and whether the different morphs are accumulating genetic differences, as might be expected since in some species females have also asymmetric gonopores and thereby can only be fertilized by compatible asymmetric males. We address this issue by combining breeding experiments with genome-wide data (ddRAD markers) in representative species of the two anablepid genera with asymmetric genitalia: Anableps and Jenynsia. Breeding experiments showed that all offspring were asymmetric, but their morphotype (i.e. right- or left-sided) was independent of parental morphotype, implying that the direction of asymmetry does not have a strong genetic component. Consistent with this conclusion, association analyses based on approximately 25 000 SNPs did not identify markers significantly associated with the direction of genital asymmetry and there was no evidence of population structure between left- and right-sided individuals. These results suggest that the direction of genital asymmetry in anablepid fishes might be stochastic, a commonly observed pattern in species with antisymmetry in morphological traits.

[Which factors determine the altitudinal distribution of tropical Andean riverine fishes]? / Qué factores determinan la distribución altitudinal de los peces de ríos tropicales andinos?

Altitudinal gradients represent an appropriate system to assess whether there is a relationship between richness patterns, environmental variables, and the ecological processes that determine the species type and number inhabiting a given area. In mountain streams freshwater fishes, the most prevalent relationship is a monotonic decrease in species richness with elevation. The objective of this study was to evaluate four hypotheses that can explain the negative relationship between local fish species richness and altitude, 1) the hypothesis of decreasing energy availability, 2) the hypothesis of increasing climate severity, 3) the hypothesis of habitat diversity, and 4) the hypothesis of isolation by physical severity of the environment. Fish and macro-invertebrates were collected following standard methods from 83 sites (between 200-4 000 meters) of two river basins in the Bolivian Amazon. The first hypothesis was tested by analyzing relationships between the density of macro-invertebrates, the richness of invertivorous fish species and altitude; while the second and third hypotheses were assessed by a multiple regression analysis (GLM) between fish species richness and several local and regional factors. Besides, assemblage dissimilarity between sites along the altitudinal gradient was analyzed using ßsim and ßness indices. Fish richness decreases linearly with increasing altitude. The density of macro-invertebrates tends to increase at higher altitudes, contrary to invertivorous fish species richness, suggesting that energy availability is not a limiting factor for fish species colonization. The GLM explained 86 % of the variation in fish species richness, with a significant contribution of water temperature, maximum slope in the river mainstem, and stream width. There is a higher species turnover (ßsim) between sites at low elevation. Inversely, ßness shows higher values in the upper parts, corresponding to change in assemblages mainly due to species loss. Taken together, these results suggest that climatic and physical severities create strong barriers to colonization, further explaining the decrease in fish richness along the altitudinal gradient.

Asymmetry in genitalia is in sync with lateralized mating behavior but not with the lateralization of other behaviors.

Asymmetries in bilateral organisms attract a lot of curiosity given that they are conspicuous departures from the norm. They allow the investigation of the integration at different levels of biological organization. Here we study whether and how behavioral and asymmetrical anatomical traits co-evolved and work together. We ask if asymmetry is determined locally for each trait or at a whole individual level in a species bearing conspicuous asymmetrical genitalia. Asymmetric genitalia evolved in many species; however, in most cases the direction of asymmetry is fixed. Therefore, it has been rarely determined if there is an association between the direction of asymmetry in genitalia and other traits. In onesided livebearer fish of the genus Jenynsia (Cyprinodontiformes, Anablepidae), the anal fin of males is modified into a gonopodium, an intromittent organ that serves to inseminate females. The gonopodium shows a conspicuous asymmetry, with its tip bending either to the left or the right. By surveying 13 natural populations of Jenynsia lineata, we found that both genital morphs are equally common in wild populations. In a series of experiments in a laboratory population, we discovered asymmetry and lateralization for multiple other traits; yet, the degree of integration varied highly among them. Lateralization in exploratory behavior in response to different stimuli was not associated with genital morphology. Interestingly, the direction of genital asymmetry was positively correlated with sidedness of mating preference and the number of neuromasts in the lateral line. This suggests integration of functionally linked asymmetric traits; however, there is no evidence that asymmetry is determined at the whole individual level in our study species.

Molecular and morphological convergence to sulfide-tolerant fishes in a new species of Jenynsia (Cyprinodontiformes: Anablepidae), the first extremophile member of the family.

Freshwater sulfide springs have extreme environmental conditions that only few vertebrate species can tolerate. These species often develop a series of morphological and molecular adaptations to cope with the challenges of life under the toxic and hypoxic conditions of sulfide springs. In this paper, we described a new fish species of the genus Jenynsia, Anablepidae, from a sulfide spring in Northwestern Argentina, the first in the family known from such extreme environment. Jenynsia sulfurica n. sp. is diagnosable by the lack of scales on the pre-pelvic area or the presence of a single row of scales, continuous or not, from the isthmus to the bases of the pelvic fins. Additionally, it presents a series of morphological and molecular characteristics that appear convergent with those seen in other fish species (e.g., Poeciliids) inhabiting sulfide springs. Most notably, J. sulfurica has an enlarged head and postorbital area compared to other fish of the genus and a prognathous lower jaw with a hypertrophied lip, thought to facilitate respiration at the air-water interface. Analyses of cox1 sequence showed that J. sulfurica has two unique mutations resulting in amino acid substitutions convergent to those seen in Poeciliids from sulfide springs and known to provide a physiological mechanism related to living in sulfide environments. A phylogenetic analysis, including molecular and morphological characters, placed J. sulfurica as sister taxa to J. alternimaculata, a species found in nearby, non-sulfide habitats directly connected to the sulfide springs. Thus, it can be inferred that the selection imposed by the presence of H2S has resulted in the divergence between these two species and has potentially served as a barrier to gene flow.

Description of a new species of the Neotropical cichlid genus Gymnogeophagus Miranda Ribeiro, 1918 (Teleostei: Cichliformes) from the Middle Paraná basin, Misiones, Argentina.

Gymnogeophagus jaryi, new species, is described from Southern tributaries of the Middle Paraná basin in Misiones. It can be distinguished from all other members of the genus, except from G. australis and G. caaguazuensis, by the presence of a hyaline to grey anterior portion of the dorsal fin. Gymnogeophagus jaryi differs from G. caaguazuensis by a longer caudal peduncle, caudal fin not lyrate, central portion of scales on dorsal portion of trunk light iridescent blue and by white spots in soft portion of dorsal fin in adult males, and from G. australis by the light iridescent blue coloration of central portion of scales on the dorsal portion of trunk and tail, and by the lack of scales on the soft portion of the dorsal fin. Additionally, it can be diagnosed by the following unique combination of characters: 10-11 dorsal-fin branched rays, 27-30 E1 scales, absence of lips thickening, and, in males, by the possession of a hump in adults, caudal fin not lyrate, presence of large white spots forming transversal stripes distally and in anterior area of the dorsal fin's soft portion, central area of scales on the dorsal portion of the trunk light iridescent blue, lack of scales on the base of the dorsal fin's soft portion, absence of a conspicuous and oblique dark band from the eye to the anterior border of the head, anterior portion of dorsal fin hyaline to grey, scales of the midlateral spot each bearing a semicircular light blue blotch, head hump starting at the horizontal through the eyes, concave anterior profile in lateral view, base of unpaired fins yellow, and whitish hyaline spots on caudal fin. The new species, based on mtDNA phylogeny, is the sister species of G. caaguazuensis from the Paraguay basin and is closely related to G. australis.

Living in the waterfalls: A new species of Trichomycterus (Siluriformes: Trichomycteridae) from Tabay stream, Misiones, Argentina.

A new species assigned to the genus Trichomycterus from the area of the waterfalls of Tabay stream, Paraná River basin, Misiones, Argentina, is described. Trichomycterus ytororo sp. nov. is distinguished from all other species in the genus by the presence of 31-35 dorsal procurrent caudal-fin rays and the combination of some external characters such as: coloration, number of pectoral-fin rays and pores of the laterosensory canals. The new taxon belongs to a presumably monophyletic group of species composed of T. crassicaudatus, T. igobi, and T. stawiarski based on the presence of 24 or more thickly ossified and rigid procurrent caudal-fin rays with a slender distal tip extending along the tips of at least ten neural spines.

A new species of Farlowella (Siluriformes: Loricariidae) from the upper Bermejo River, La Plata River basin, northwestern Argentina

A new species of Farlowella is described from the Bermejo River basin, in Salta and Jujuy provinces, northwestern Argentina. The new species belongs to the Farlowella nattereri species group. The new species is diagnosed by the following combination of characters: marbled rostrum, five rows of lateral plates series, relatively short snout (snout-mouth length less than 50.0% of head length), complete half-moon shaped spot on caudal fin, and short predorsal distance (37.8-41.8% of standard length).(AU)

Se describe una nueva especie de Farlowella de la cuenca del Río Bermejo, en las provincias de Salta y Jujuy, en el noroeste de Argentina. La nueva especie pertenece al grupo de Farlowella nattereri. La nueva especie se diagnostica con la siguiente combinación de caracteres: rostro veteado, cinco hileras de placas laterales en el cuerpo, hocico relativamente corto (longitud hocico-boca menor a 50.0% longitud cabeza), mancha en forma de media luna completa en la aleta caudal, y distancia predorsal corta (37.8-41.8% longitud estándar).(AU)

¿Qué factores determinan la distribución altitudinal de los peces de ríos tropicales andinos? / Which factors determine the altitudinal distribution of tropical Andean riverine fishes?

ResumenLos gradientes altitudinales representan un sistema apropiado para evaluar si existe una relación general entre los patrones de riqueza, variables ambientales, y los procesos ecológicos que determinan cuantas especies co-ocurren en un área dada.En cuanto a los peces de agua dulce en arroyos de montaña la relación prevalente es un decrecimiento monotónico en riqueza de especies con la altura. El objetivo de este estudio es evaluar cuatro hipótesis que pueden explicar la relación negativa entre la riqueza local de especies de peces y la altura, 1) hipótesis de la disminución de la energía disponible 2) hipótesis del aumento del rigor climático 3) hipótesis de la diversidad de hábitats e 4) hipótesis de la severidad de las condiciones físicas del hábitat que se presentan río arriba, relacionada a las dificultades para colonizar las partes altas de los ríos. Se recolectaron peces y macro-invertebrados en 83 sitios entre 200 a 4 000 m en dos cuencas en la Amazonía boliviana. La primera hipótesis fue evaluada mediante regresiones entre la densidad de macro-invertebrados, los peces invertívoros y la altura. Para evaluar la segunda y tercera hipótesis se realizó un análisis de regresión múltiple (GLM), entre la riqueza y varios factores locales y regionales. Se analizó también la disimilitud de los ensamblajes de peces sobre el gradiente altitudinal utilizando los índices βsim y βnes. Se encontró que la riqueza de peces disminuye de manera lineal con el aumento de la altura. La densidad de macroinvertebrados aumenta con la altura, contrariamente a la riqueza de peces invertívoros, sugiriendo que la energía no constituye un factor limitante para la colonización de las especies de peces en zonas altas. El GLM explicó un 86 % de la variación total de la riqueza, con un aporte significativo de la temperatura del agua, las pendientes máximas en el río principal y el ancho de río. El βsim presentó valores altos en niveles de altura bajos, mostrando mayor recambio de especies entre los sitios cercanos a la fuente de colonización. Inversamente, el βnes mostró valores mayores en las partes altas, correspondientes a cambios por pérdida de especies. Estos resultados sugieren que el rigor climático junto con la severidad física del hábitat crean barreras a la colonización y explican los cambios de riqueza en el gradiente altitudinal.

AbstractAltitudinal gradients represent an appropriate system to assess whether there is a relationship between richness patterns, environmental variables, and the ecological processes that determine the species type and number inhabiting a given area. In mountain streams freshwater fishes, the most prevalent relationship is a monotonic decrease in species richness with elevation. The objective of this study was to evaluate four hypotheses that can explain the negative relationship between local fish species richness and altitude, 1) the hypothesis of decreasing energy availability, 2) the hypothesis of increasing climate severity, 3) the hypothesis of habitat diversity, and 4) the hypothesis of isolation by physical severity of the environment. Fish and macro-invertebrates were collected following standard methods from 83 sites (between 200-4 000 meters) of two river basins in the Bolivian Amazon. The first hypothesis was tested by analyzing relationships between the density of macro-invertebrates, the richness of invertivorous fish species and altitude; while the second and third hypotheses were assessed by a multiple regression analysis (GLM) between fish species richness and several local and regional factors. Besides, assemblage dissimilarity between sites along the altitudinal gradient was analyzed using βsim and βness indices. Fish richness decreases linearly with increasing altitude. The density of macro-invertebrates tends to increase at higher altitudes, contrary to invertivorous fish species richness, suggesting that energy availability is not a limiting factor for fish species colonization. The GLM explained 86 % of the variation in fish species richness, with a significant contribution of water temperature, maximum slope in the river mainstem, and stream width. There is a higher species turnover (βsim) between sites at low elevation. Inversely, βness shows higher values in the upper parts, corresponding to change in assemblages mainly due to species loss. Taken together, these results suggest that climatic and physical severities create strong barriers to colonization, further explaining the decrease in fish richness along the altitudinal gradient.

Jenynsia luxata, a new species from Northwestern Argentina, with additional observations of J. maculata regan and phylogeny of the genus (cyprinodontiformes: anablepidae)

Jenynsia luxata, a new species from northwestern Argentina, is described. This species is diagnosable from all other Jenynsia by the medial processes of left and right pelvic bones relatively reduced and separated from each other. The new species resembles J. multidentata, but it is further distinguished from this species by the absence of a swelling between the urogenital opening and the anterior base of the anal fin in females and details of coloration. Phylogenetic analyses, both under implied and equal weighting, recover the subgenera Plesiojenynsia and Jenynsia as monophyletic units. New information on previously missing characters of Jenynsia maculata is added. These data and phylogenetic characters coded for the new species herein described contribute to a better resolution of the phylogenetic relationships within the subgenus Jenynsia, which is herein supported by additional synapomorphies relative to previous phylogenies.

Jenynsia luxata, una nueva especie del noroeste de Argentina, es descripta. Esta especie es diagnosticable por presentar los procesos mediales de los huesos pélvicos izquierdo y derecho relativamente reducidos y separados. La nueva especie se parece a J. multidentata, pero se distingue de ésta por la ausencia de un abultamiento entre la abertura urogenital y la base de la aleta anal en hembras y por detalles en el patrón de coloración. Los análisis filogenéticos, tanto bajo pesos implicados como iguales, recuperan los subgéneros Plesiojenynsia y Jenynsia como unidades monofiléticas. Se aporta nueva información sobre caracteres de J. maculata previamente codificados como entradas faltantes. Esos datos y los caracteres filogenéticos codificados para la nueva especie aquí descripta contribuyen a una mayor resolución de las relaciones filogenéticas dentro del subgénero Jenynsia, que está aquí soportado por sinapomorfías adicionales en relación a las filogenias previas.