Extreme heterogeneity in sex chromosome differentiation and dosage compensation in livebearers.

Once recombination is halted between the X and Y chromosomes, sex chromosomes begin to differentiate and transition to heteromorphism. While there is a remarkable variation across clades in the degree of sex chromosome divergence, far less is known about the variation in sex chromosome differentiation within clades. Here, we combined whole-genome and transcriptome sequencing data to characterize the structure and conservation of sex chromosome systems across Poeciliidae, the livebearing clade that includes guppies. We found that the Poecilia reticulata XY system is much older than previously thought, being shared not only with its sister species, Poecilia wingei, but also with Poecilia picta, which diverged roughly 20 million years ago. Despite the shared ancestry, we uncovered an extreme heterogeneity across these species in the proportion of the sex chromosome with suppressed recombination, and the degree of Y chromosome decay. The sex chromosomes in P. reticulata and P. wingei are largely homomorphic, with recombination in the former persisting over a substantial fraction. However, the sex chromosomes in P. picta are completely nonrecombining and strikingly heteromorphic. Remarkably, the profound degradation of the ancestral Y chromosome in P. picta is counterbalanced by the evolution of functional chromosome-wide dosage compensation in this species, which has not been previously observed in teleost fish. Our results offer important insight into the initial stages of sex chromosome evolution and dosage compensation.

Local ancestry analysis reveals genomic convergence in extremophile fishes.

The molecular basis of convergent phenotypes is often unknown. However, convergence at a genomic level is predicted when there are large population sizes, gene flow among diverging lineages or strong genetic constraints. We used whole-genome resequencing to investigate genomic convergence in fishes ( Poecilia spp.) that have repeatedly colonized hydrogen sulfide (H2S)-rich environments in Mexico. We identified genomic similarities in both single nucleotide polymorphisms (SNPs) and structural variants (SVs) among independently derived sulfide spring populations, with approximately 1.2% of the genome being shared among sulfidic ecotypes. We compared these convergent genomic regions to candidate genes for H2S adaptation identified from transcriptomic analyses and found that a significant proportion of these candidate genes (8%) were also in regions where sulfidic individuals had similar SNPs, while only 1.7% were in regions where sulfidic individuals had similar SVs. Those candidate genes included genes involved in sulfide detoxification, the electron transport chain (the main toxicity target of H2S) and other processes putatively important for adaptation to sulfidic environments. Regional genomic similarity across independent populations exposed to the same source of selection is consistent with selection on standing variation or introgression of adaptive alleles across divergent lineages. However, combined with previous analyses, our data also support that adaptive changes in mitochondrially encoded subunits arose independently via selection on de novo mutations. Pressing questions remain on what conditions ultimately facilitate the independent rise of adaptive alleles at the same loci in separate populations, and thus, the degree to which evolution is repeatable or predictable. This article is part of the theme issue 'Convergent evolution in the genomics era: new insights and directions'.

Possible glimpses into early speciation: the effect of ovarian fluid on sperm velocity accords with post-copulatory isolation between two guppy populations.

Identifying mechanisms of reproductive isolation is key to understanding speciation. Among the putative mechanisms underlying reproductive isolation, sperm-female interactions (post-mating-prezygotic barriers) are arguably the hardest to identify, not least because these are likely to operate at the cellular or molecular level. Yet sperm-female interactions offer great potential to prevent the transfer of genetic information between different populations at the initial stages of speciation. Here, we provide a preliminary test for the presence of a putative post-mating-prezygotic barrier operating between three populations of Trinidadian guppies (Poecilia reticulata), an internally fertilizing fish that inhabits streams with different levels of connectivity across Trinidad. We experimentally evaluate the effect of female ovarian fluid on sperm velocity (a predictor of competitive fertilization success) according to whether males and females were from the same (native) or different (foreign) populations. Our results reveal the potential for ovarian fluid to act as a post-mating-prezygotic barrier between two populations from different drainages, but also that the strength of this barrier is different among populations. This result may explain the previous finding that, in some populations, sperm from native males have precedence over foreign sperm, which could eventually lead to reproductive isolation between these populations.

megasat: automated inference of microsatellite genotypes from sequence data.

megasat is software that enables genotyping of microsatellite loci using next-generation sequencing data. Microsatellites are amplified in large multiplexes, and then sequenced in pooled amplicons. megasat reads sequence files and automatically scores microsatellite genotypes. It uses fuzzy matches to allow for sequencing errors and applies decision rules to account for amplification artefacts, including nontarget amplification products, replication slippage during PCR (amplification stutter) and differential amplification of alleles. An important feature of megasat is the generation of histograms of the length-frequency distributions of amplification products for each locus and each individual. These histograms, analogous to electropherograms traditionally used to score microsatellite genotypes, enable rapid evaluation and editing of automatically scored genotypes. megasat is written in Perl, runs on Windows, Mac OS X and Linux systems, and includes a simple graphical user interface. We demonstrate megasat using data from guppy, Poecilia reticulata. We genotype 1024 guppies at 43 microsatellites per run on an Illumina MiSeq sequencer. We evaluated the accuracy of automatically called genotypes using two methods, based on pedigree and repeat genotyping data, and obtained estimates of mean genotyping error rates of 0.021 and 0.012. In both estimates, three loci accounted for a disproportionate fraction of genotyping errors; conversely, 26 loci were scored with 0-1 detected error (error rate ≤0.007). Our results show that with appropriate selection of loci, automated genotyping of microsatellite loci can be achieved with very high throughput, low genotyping error and very low genotyping costs.

Molecular and taxonomic characterisation of introduced specimens of Poecilia reticulata in the lower Paraguay River basin (Cyprinodontiformes: Poeciliidae)

Poeciliids comprise around 300 species inhabiting the fresh and brackish waters of the Americas and Africa. Poecilia reticulata is native to Northeastern South America and Trinidad and Tobago. In this paper, introduced specimens of P. reticulata collected in the lower Paraguay River in Argentina, were characterized by means of molecular and taxonomic approaches. We further explore, by means of DNA Barcoding, the singularity of the genetic identity of these specimens. Ocurrence of P. reticulata in the lower Paraguay River represents the first record of this species in Argentina. Thirteen individuals of P. reticulata were collected. DNA barcoding showed that all five specimens sampled belong to a single mitochondrial lineage, which was also present in 11 countries from five continents. The distance-based tree clearly grouped separetely four different clusters of P. reticulata when including public data. Genetic distance between the most divergent P. reticulata almost paralleled distance between this species and Poecilia mexicana and P. vivipara. Established populations from Paraguay could be one of the plausible sources for the introduced populations recorded in the lower Paraguay River. The presence of P. reticulata in an open waterway with known drainage to a natural stream is of major concern.(AU)

Poecilídeos compreendem cerca de 300 espécies que habitam águas doces e salobras das Américas e África. Poecilia reticulata é nativa do nordeste da América do Sul e Trinidad e Tobago. Neste trabalho, espécimes introduzidos de P. reticulata coletados no baixo rio Paraguai na Argentina, foram caracterizados por meio de abordagens moleculares e taxonômicas. Exploramos ainda, por meio de DNA Barcoding, a singularidade da identidade genética destes espécimes. A ocorrência de P. reticulata no baixo rio Paraguai representa o primeiro registro dessa especie na Argentina. Treze indivíduos de P. reticulata foram coletados. O Barcoding mostrou que todos os espécimes pertencem a uma única linhagem mitocondrial, a qual está presente em 11 países dos cinco continentes. A árvore de distâncias agrupou separadamente quatro clusters diferentes de P. reticulata quando incluindo dados públicos. A distância genética entre os agrupamentos mais divergentes de P. reticulata quase igualou a distância entre esta espécie e Poecilia mexicana e P. vivipara. As populações estabelecidas no Paraguai poderiam ser uma das fontes plausíveis para as populações introduzidas registradas no baixo rio Paraguai. A presença de P. reticulata em um canal aberto com drenagem conhecida para um córrego natural é de grande preocupação.(AU)

Phylogenetic analyses of the subgenus Mollienesia (Poecilia, Poeciliidae, Teleostei) reveal taxonomic inconsistencies, cryptic biodiversity, and spatio-temporal aspects of diversification in Middle America.

The subgenus Mollienesia is a diverse group of freshwater fishes, including species that have served as important models across multiple biological disciplines. Nonetheless, the taxonomic history of this group has been conflictive and convoluted, in part because the evolutionary relationships have not been rigorously resolved. We conducted a comprehensive molecular phylogenetic analysis of the subgenus Mollienesia to identify taxonomic discrepancies and potentially identify undescribed species, estimate ancestral areas of origin and estimate dates of divergence, as well as explore biogeographical patterns. Our findings confirm the presence of three main clades composed of the P. latipinna, P. sphenops, and P. mexicana species complexes. Unlike previously hypothesized morphology-based analyses, species found on the Caribbean Islands are not part of Mollienesia, but are more closely related to species of the subgenus Limia. Our study also revealed several taxonomic inconsistencies and distinct lineages in the P. mexicana species complex that may represent undescribed species. The diversity in the subgenus Mollienesia is a result of dynamic geologic activity leading to vicariant events, dispersal across geologic blocks, and ecological speciation.

Phylogeny and biogeography of Poecilia (Cyprinodontiformes: Poeciliinae) across Central and South America based on mitochondrial and nuclear DNA markers.

Poeciliids are a diverse group of small Neotropical fishes, and despite considerable research attention as models in ecology and evolutionary biology, our understanding of their biogeographic and phylogenetic relationships is still limited. We investigated the phylogenetic relationships of South and Central American Poecilia, by examining 2395 base pairs of mitochondrial DNA (ATPase 8/6, COI) and nuclear DNA (S7) for 18 species across six subgenera. Fifty-eight novel sequences were acquired from newly collected specimens and 20 sequences were obtained from previously published material. Analyses of concatenated and partitioned mitochondrial DNA and nuclear DNA sets resulted in a well-supported phylogeny that resolved several monophyletic groups corresponding to previously hypothesized subgenera and species complexes. A divergence-dating analysis supported the hypothesis of the genus Poecilia dispersing into Central America in the early Pliocene (ancestors of Psychropoecilia+Allopoecilia+Mollienesia: 7.3-2.0Mya) from predominantly South America. Subsequently, one lineage (subgenus Allopoecilia: 5.1-1.3Mya) expanded deeper into South America from Lower-Central America, and one lineage expanded from Nuclear-Central America into South America (subgenus Mollienesia: 0.71-0.14Mya). The subgenus Mollienesia diverged into three monophyletic groups that can be identified by nuptial male dorsal fin morphology and inner jaw dentition. A subclade of the unicuspid short-fins (subgenus Mollienesia) was the lineage that expanded into South America during the middle Pleistocene. Species in this subclade are now distributed across northern South America, where they are partially sympatric with Allopoecilia. However the P. (A.) caucana complex was not monophyletic, with P. (A.) wandae clustering in the Mollienesia subclade that expanded into South America. It is apparent that characters (body size, scale count, pigmentation, and gonopodium morphology) used to define the P. (A.) caucana complex are homoplastic and suggestive of rapid convergence in northern South America. Our improved taxon sampling and divergence-time calibration allowed for insights into the timing and direction of dispersals, and provides an improved understanding of the biogeographic history of an enigmatic group of fishes. Furthermore, we provided strong evidence for the monophyly of the subgenus Mollienesia and further substantiated its species complexes; therefore, we advise a taxonomic re-evaluation for the P. (A.) caucana complex to maintain monophyly of both Mollienesia and Allopoecilia.

Color vision varies more among populations than among species of live-bearing fish from South America.

BACKGROUND: Sensory Bias models for the evolution of mate preference place a great emphasis on the role of sensory system variation in mate preferences. However, the extent to which sensory systems vary across- versus within-species remains largely unknown. Here we assessed whether color vision varies in natural locations where guppies (Poecilia reticulata) and their two closest relatives, Poecilia parae and Poecilia picta, occur in extreme sympatry and school together. All three species base mate preferences on male coloration but differ in the colors preferred. RESULTS: Measuring opsin gene expression, we found that within sympatric locations these species have similar color vision and that color vision differed more across populations of conspecifics. In addition, all three species differ across populations in the frequency of the same opsin coding polymorphism that influences visual tuning. CONCLUSIONS: Together, this shows sensory systems vary considerably across populations and supports the possibility that sensory system variation is involved in population divergence of mate preference.

Assessing species boundaries using multilocus species delimitation in a morphologically conserved group of neotropical freshwater fishes, the Poecilia sphenops species complex (Poeciliidae).

Accurately delimiting species is fundamentally important for understanding species diversity and distributions and devising effective strategies to conserve biodiversity. However, species delimitation is problematic in many taxa, including 'non-adaptive radiations' containing morphologically cryptic lineages. Fortunately, coalescent-based species delimitation methods hold promise for objectively estimating species limits in such radiations, using multilocus genetic data. Using coalescent-based approaches, we delimit species and infer evolutionary relationships in a morphologically conserved group of Central American freshwater fishes, the Poecilia sphenops species complex. Phylogenetic analyses of multiple genetic markers (sequences of two mitochondrial DNA genes and five nuclear loci) from 10/15 species and genetic lineages recognized in the group support the P. sphenops species complex as monophyletic with respect to outgroups, with eight mitochondrial 'major-lineages' diverged by ≥2% pairwise genetic distances. From general mixed Yule-coalescent models, we discovered (conservatively) 10 species within our concatenated mitochondrial DNA dataset, 9 of which were strongly supported by subsequent multilocus Bayesian species delimitation and species tree analyses. Results suggested species-level diversity is underestimated or overestimated by at least ~15% in different lineages in the complex. Nonparametric statistics and coalescent simulations indicate genealogical discordance among our gene tree results has mainly derived from interspecific hybridization in the nuclear genome. However, mitochondrial DNA show little evidence for introgression, and our species delimitation results appear robust to effects of this process. Overall, our findings support the utility of combining multiple lines of genetic evidence and broad phylogeographical sampling to discover and validate species using coalescent-based methods. Our study also highlights the importance of testing for hybridization versus incomplete lineage sorting, which aids inference of not only species limits but also evolutionary processes influencing genetic diversity.

Evidence for cryptic speciation in directly transmitted gyrodactylid parasites of Trinidadian guppies.

Cryptic species complexes are common among parasites, which tend to have large populations and are subject to rapid evolution. Such complexes may arise through host-parasite co-evolution and/or host switching. For parasites that reproduce directly on their host, there might be increased opportunities for sympatric speciation, either by exploiting different hosts or different micro-habitats within the same host. The genus Gyrodactylus is a specious group of viviparous monogeneans. These ectoparasites transfer between teleosts during social contact and cause significant host mortality. Their impact on the guppy (Poecilia reticulata), an iconic evolutionary and ecological model species, is well established and yet the population genetics and phylogenetics of these parasites remains understudied. Using mtDNA sequencing of the host and its parasites, we provide evidence of cryptic speciation in Gyrodactylus bullatarudis, G. poeciliae and G. turnbulli. For the COII gene, genetic divergence of lineages within each parasite species ranged between 5.7 and 17.2%, which is typical of the divergence observed between described species in this genus. Different lineages of G. turnbulli and G. poeciliae appear geographically isolated, which could imply allopatric speciation. In addition, for G. poeciliae, co-evolution with a different host species cannot be discarded due to its host range. This parasite was originally described on P. caucana, but for the first time here it is also recorded on the guppy. The two cryptic lineages of G. bullatarudis showed considerable geographic overlap. G. bullatarudis has a known wide host range and it can also utilize a killifish (Anablepsoides hartii) as a temporary host. This killifish is capable of migrating overland and it could act as a transmission vector between otherwise isolated populations. Additional genetic markers are needed to confirm the presence of these cryptic Gyrodactylus species complexes, potentially leading to more in-depth genetic, ecological and evolutionary analyses on this multi-host-parasite system.

Transcriptome assemblies for studying sex-biased gene expression in the guppy, Poecilia reticulata.

BACKGROUND: Sexually dimorphic phenotypes are generally associated with differential gene expression between the sexes. The study of molecular evolution and genomic location of these differentially expressed, or sex-biased, genes is important for understanding inter-sexual divergence under sex-specific selection pressures. Teleost fish provide a unique opportunity to examine this divergence in the presence of variable sex-determination mechanisms of recent origin. The guppy, Poecilia reticulata, displays sexual dimorphism in size, ornaments, and behavior, traits shaped by natural and sexual selection in the wild. RESULTS: To gain insight into molecular mechanisms underlying the guppy's sexual dimorphism, we assembled a reference transcriptome combining genome-independent as well as genome-guided assemblies and analyzed sex-biased gene expression between different tissues of adult male and female guppies. We found tissue-associated sex-biased expression of genes related to pigmentation, signal transduction, and spermatogenesis in males; and growth, cell-division, extra-cellular matrix organization, nutrient transport, and folliculogenesis in females. While most sex-biased genes were randomly distributed across linkage groups, we observed accumulation of ovary-biased genes on the sex linkage group, LG12. Both testis-biased and ovary-biased genes showed a significantly higher rate of non-synonymous to synonymous substitutions (dN/dS) compared to unbiased genes. However, in somatic tissues only female-biased genes, including those co-expressed in multiple tissues, showed elevated ratios of non-synonymous substitutions. CONCLUSIONS: Our work identifies a set of annotated gene products that are candidate factors affecting sexual dimorphism in guppies. The differential genomic distribution of gonad-biased genes provides evidence for sex-specific selection pressures acting on the nascent sex chromosomes of the guppy. The elevated rates of evolution of testis-biased and female-biased genes indicate differing evolution under distinct selection pressures on the reproductive versus non-reproductive tissues.

Selection from parasites favours immunogenetic diversity but not divergence among locally adapted host populations.

The unprecedented polymorphism in the major histocompatibility complex (MHC) genes is thought to be maintained by balancing selection from parasites. However, do parasites also drive divergence at MHC loci between host populations, or do the effects of balancing selection maintain similarities among populations? We examined MHC variation in populations of the livebearing fish Poecilia mexicana and characterized their parasite communities. Poecilia mexicana populations in the Cueva del Azufre system are locally adapted to darkness and the presence of toxic hydrogen sulphide, representing highly divergent ecotypes or incipient species. Parasite communities differed significantly across populations, and populations with higher parasite loads had higher levels of diversity at class II MHC genes. However, despite different parasite communities, marked divergence in adaptive traits and in neutral genetic markers, we found MHC alleles to be remarkably similar among host populations. Our findings indicate that balancing selection from parasites maintains immunogenetic diversity of hosts, but this process does not promote MHC divergence in this system. On the contrary, we suggest that balancing selection on immunogenetic loci may outweigh divergent selection causing divergence, thereby hindering host divergence and speciation. Our findings support the hypothesis that balancing selection maintains MHC similarities among lineages during and after speciation (trans-species evolution).

Comparative analysis of the gonadal transcriptomes of the all-female species Poecilia formosa and its maternal ancestor Poecilia mexicana.

BACKGROUND: The Amazon molly, Poecilia formosa (Teleostei: Poeciliinae) is an unisexual, all-female species. It evolved through the hybridisation of two closely related sexual species and exhibits clonal reproduction by sperm dependent parthenogenesis (or gynogenesis) where the sperm of a parental species is only used to activate embryogenesis of the apomictic, diploid eggs but does not contribute genetic material to the offspring.Here we provide and describe the first de novo assembled transcriptome of the Amazon molly in comparison with its maternal ancestor, the Atlantic molly Poecilia mexicana. The transcriptome data were produced through sequencing of single end libraries (100 bp) with the Illumina sequencing technique. RESULTS: 83,504,382 reads for the Amazon molly and 81,625,840 for the Atlantic molly were assembled into 127,283 and 78,961 contigs for the Amazon molly and the Atlantic molly, respectively. 63% resp. 57% of the contigs could be annotated with gene ontology terms after sequence similarity comparisons. Furthermore, we were able to identify genes normally involved in reproduction and especially in meiosis also in the transcriptome dataset of the apomictic reproducing Amazon molly. CONCLUSIONS: We assembled and annotated the transcriptome of a non-model organism, the Amazon molly, without a reference genome (de novo). The obtained dataset is a fundamental resource for future research in functional and expression analysis. Also, the presence of 30 meiosis-specific genes within a species where no meiosis is known to take place is remarkable and raises new questions for future research.

Population structure of guppies in north-eastern Venezuela, the area of putative incipient speciation.

BACKGROUND: Geographic barriers to gene flow and divergence among populations in sexual traits are two important causes of genetic isolation which may lead to speciation. Genetic isolation may be facilitated if these two mechanisms act synergistically. The guppy from the Cumaná region (within the Cariaco drainage) of eastern Venezuela has been previously described as a case of incipient speciation driven by sexual selection, significantly differentiated in sexual colouration and body shape from the common guppy, Poecilia reticulata. The latter occurs widely in northern Venezuela, including the south-eastern side of Cordillera de la Costa, where it inhabits streams belonging to the San Juan drainage. Here, we present molecular and morphological analyses of differentiation among guppy populations in the Cariaco and San Juan drainages. Our analyses are based on a 953 bp long mtDNA fragment, a set of 15 microsatellites (519 fish from 20 populations), and four phenotypic traits. RESULTS: Both microsatellite and mtDNA data showed that guppies inhabiting the two drainages are characterised by a significant genetic differentiation, but a higher proportion of the genetic variance was distributed among populations within regions. Most guppies in the Cariaco drainage had mtDNA from a distinct lineage, but we also found evidence for widespread introgression of mtDNA from the San Juan drainage into the Cariaco drainage. Phenotypically, populations in the two regions differed significantly only in the number of black crescents. Phenotypic clustering did not support existence of two distinct groupings, but indicated a degree of distinctiveness of Central Cumaná (CC) population. However, CC population showed little differentiation at the neutral markers from the proximate populations within the Cariaco drainage. CONCLUSIONS: Our findings are consistent with only partial genetic isolation between the two geographic regions and indicate that the geographic barrier of Cordillera de la Costa has not played an important role in strengthening the incomplete pre-zygotic reproductive barrier between Cumaná and common guppy. Significant phenotypic differentiation between genetically similar (in terms of neutral variation) populations suggests that mate choice can maintain divergence at sexually selected traits despite gene flow. However, neither genetic nor phenotypic clustering supported delineation of two species within the region.

Multiple pigment cell types contribute to the black, blue, and orange ornaments of male guppies (Poecilia reticulata).

The fitness of male guppies (Poecilia reticulata) highly depends on the size and number of their black, blue, and orange ornaments. Recently, progress has been made regarding the genetic mechanisms underlying male guppy pigment pattern formation, but we still know little about the pigment cell organization within these ornaments. Here, we investigate the pigment cell distribution within the black, blue, and orange trunk spots and selected fin color patterns of guppy males from three genetically divergent strains using transmission electron microscopy. We identified three types of pigment cells and found that at least two of these contribute to each color trait. Further, two pigment cell layers, one in the dermis and the other in the hypodermis, contribute to each trunk spot. The pigment cell organization within the black and orange trunk spots was similar between strains. The presence of iridophores in each of the investigated color traits is consistent with a key role for this pigment cell type in guppy color pattern formation.

Trophic ecology of the exotic Lerma livebearer Poeciliopsis infans (Cyprinodontiformes: Poeciliidae) in the Lago de Pátzcuaro, Central Mexico.

Exotic fish species has caused several impacts on aquatic biodiversity. The Lago de Pátzcuaro has some well-studied exotic species, except the Lerma livebearer Poeciliopsis infans. This fish species was introduced into the Lago de Pátzcuaro before 1997 and the aspects of its biology are still unknown. In this study we assessed aspects of the trophic ecology of this exotic fish, P infans, using gut content and stable isotope analysis to understand its capacity to tolerate anthropogenic environmental degradation in the Lago de Pátzcuaro. We also determined its trophic guild position (TP) using the TrophLab Program and stable isotope. Niche breadth was calculated by standardized Levins' Index (Bi). Fish was captured with a seine during wet and dry seasons at six environmentally different sites and gut contents were obtained. We analyzed a total of 239 gut contents of P. infans. The contribution of each food item in the diet was quantified using frequency of occurrence and area percentage. The importance of each prey item was determined according to the index of relative importance (IRI), and the omnivory index (OI) was used to assess the feeding behavior. Fish were categorized by size and the diet was compared between fish sizes and sites. Dorsal muscle tissue and water hyacinth tissue was obtained for nitrogen isotope signature he fish can behave as a specialist (Bi = 0.39) or generalist (Bi = 0.68) and as a primary consumer (TROPH = 2.2; TP = 2.3) with a feeding strategy that was the same at dif- f erent sizes, seasons and sites. None of the evaluated sites showed good environmental quality. We argue that P infans can tolerate changes in water quality and feeding items availability, because it can exploit resources in multiple trophic webs. However, this species could be dependent on habitat complexity, especially in the aquatic vegetation cover.

Trophic ecology of the exotic Lerma livebearer Poeciliopsis infans (Cyprinodontiformes: Poeciliidae) in the Lago de Pátzcuaro, Central Mexico / Ecología trófica del pez exótico Guatapote del Lerma Poeciliopsis infans (Cyprinodontiformes: Poeciliidae) en el Lago de Pátzcuaro, Región Central de México

Exotic fish species has caused several impacts on aquatic biodiversity. The Lago de Pátzcuaro has some well-studied exotic species, except the Lerma livebearer Poeciliopsis infans. This fish species was introduced into the Lago de Pátzcuaro before 1997 and the aspects of its biology are still unknown. In this study we assessed aspects of the trophic ecology of this exotic fish, P. infans, using gut content and stable isotope analysis to understand its capacity to tolerate anthropogenic environmental degradation in the Lago de Pátzcuaro. We also determined its trophic guild position (TP) using the TrophLab Program and stable isotope. Niche breadth was calculated by standardized Levins’ Index (Bi). Fish was captured with a seine during wet and dry seasons at six environmentally different sites and gut contents were obtained. We analyzed a total of 239 gut contents of P. infans. The contribution of each food item in the diet was quantified using frequency of occurrence and area percentage. The importance of each prey item was determined according to the index of relative importance (IRI), and the omnivory index (OI) was used to assess the feeding behavior. Fish were categorized by size and the diet was compared between fish sizes and sites. Dorsal muscle tissue and water hyacinth tissue was obtained for nitrogen isotope signature (δ15N) analysis. Additionally, we measured water and habitat quality to evaluate environmental conditions at each site. We concluded that P. infans is an omnivore (OI=0.28) that consumes mainly detritus (44%), epiphytic diatoms (37%), and secondary on terrestrial insects (6%) and zooplankton (10%). The fish can behave as a specialist (Bi=0.39) or generalist (Bi=0.68) and as a primary consumer (TROPH=2.2; TP=2.3) with a feeding strategy that was the same at different sizes, seasons and sites. None of the evaluated sites showed good environmental quality. We argue that P. infans can tolerate changes in water quality and feeding items availability, because it can exploit resources in multiple trophic webs. However, this species could be dependent on habitat complexity, especially in the aquatic vegetation cover.

Las especies de peces exóticas han causado numerosos impactos sobre la biodiversidad acuática. El Lago de Pátzcuaro tiene especies exóticas bien estudiadas, excepto el Guatapote del Lerma, Poeciliopsis infans. Esta especie fue introducida en el Lago de Pátzcuaro antes de 1997 y los aspectos de su biología son aún desconocidos. Se estudiaron aspectos de la ecología trófica de este pez exótico, P. infans, mediante el uso de análisis de contenidos del tracto digestivo para entender su capacidad para tolerar la degradación ambiental antrópica en el Lago de Pátzcuaro. Lo que involucró la determinación del gremio y posición trófica (TP) con el programa TrophLab e isotopos estables. La amplitud de dieta fue calculada con el índice estandarizado de Levin (Bi). Un total de 239 tractos digestivos de P.infans fueron analizados. Los peces fueron capturados con una red tipo chinchorro durante la temporada de lluvias y estiaje en seis sitios ambientalmente distintos. La contribución de cada componente de la dieta fue cuantificada con la frecuencia de ocurrencia y el porcentaje de área. La importancia relativa de cada artículo alimentario fue determinada de acuerdo al índice de importancia relativa (IIR) y el índice de omnivoría (OI) fue usado para estimar la conducta en la forma de alimentarse. La dieta fue comparada entre tallas y entre sitios de estudio. El tejido de músculo dorsal y el tejido de lirio fueron obtenidos para determinar valores de δ15N. De manera adicional, la calidad del agua y del hábitat fue medida para evaluar la condición ambiental en cada sitio. P. infans es un omnívoro (OI=0.28) que consume principalmente detritus (44%), diatomeas epifíticas (37%) y de forma secundaria insectos terrestres (6%) y zooplancton (10%); puede comportarse como especialista (Bi=0.39) o generalista (Bi=0.68) y consumidor primario (TROPH=2.2; TP=2.3). La estrategia alimenticia fue similar entre tallas, temporadas y sitios. Ningún sitio mostró buena calidad ambiental. Los resultados del presente estudio permiten argumentar que P. infans puede tolerar cambios en la calidad del agua y en la disponibilidad de artículos alimenticios, porque puede usar recursos provenientes de múltiples redes tróficas. Sin embargo, esta especie podría ser dependiente de la complejidad del hábitat, especialmente de la cobertura de la vegetación acuática.

The rediscovery of a long described species reveals additional complexity in speciation patterns of poeciliid fishes in sulfide springs.

The process of ecological speciation drives the evolution of locally adapted and reproductively isolated populations in response to divergent natural selection. In Southern Mexico, several lineages of the freshwater fish species of the genus Poecilia have independently colonized toxic, hydrogen sulfide-rich springs. Even though ecological speciation processes are increasingly well understood in this system, aligning the taxonomy of these fish with evolutionary processes has lagged behind. While some sulfide spring populations are classified as ecotypes of Poecilia mexicana, others, like P. sulphuraria, have been described as highly endemic species. Our study particularly focused on elucidating the taxonomy of the long described sulfide spring endemic, Poecilia thermalis Steindachner 1863, and investigates if similar evolutionary patterns of phenotypic trait divergence and reproductive isolation are present as observed in other sulfidic species of Poecilia. We applied a geometric morphometric approach to assess body shape similarity to other sulfidic and non-sulfidic fish of the genus Poecilia. We also conducted phylogenetic and population genetic analyses to establish the phylogenetic relationships of P. thermalis and used a population genetic approach to determine levels of gene flow among Poecilia from sulfidic and non-sulfidic sites. Our results indicate that P. thermalis' body shape has evolved in convergence with other sulfide spring populations in the genus. Phylogenetic analyses placed P. thermalis as most closely related to one population of P. sulphuraria, and population genetic analyses demonstrated that P. thermalis is genetically isolated from both P. mexicana ecotypes and P. sulphuraria. Based on these findings, we make taxonomic recommendations for P. thermalis. Overall, our study verifies the role of hydrogen sulfide as a main factor shaping convergent, phenotypic evolution and the emergence of reproductive isolation between Poecilia populations residing in adjacent sulfidic and non-sulfidic environments.

Phylogeny and biogeography of the Poecilia sphenops species complex (Actinopterygii, Poeciliidae) in Central America.

We inferred the phylogenetic relationships among members of the Poecilia sphenops species complex to resolve the colonization process and radiation of this group in Central America. We analyzed 2550 base pairs (bp) of mitochondrial DNA (mtDNA), including ATP synthase 6 and 8, cytochrome oxidase subunit I and NADH dehydrogenase subunit 2 genes, and 906bp of the nuclear S7 ribosomal protein of 86 ingroup individuals from 61 localities spanning most of its distribution from Mexico to Panama. Our mitochondrial data rendered a well-supported phylogeny for the P. sphenops complex that differed with the nuclear data set topology, which did not recover the monophyly of the P. mexicana mitochondrial lineage. Coalescent-based simulations tests indicated that, although hybridization cannot be completely ruled out, this incongruence is most likely due to incomplete lineage sorting in this group, which also showed the widest geographic distribution. A single colonization event of Central America from South America was estimated to have occurred between the early Paleocene and Oligocene (53-22millionyears ago). Subsequently, two largely differentiated evolutionary lineages diverged around the Early Oligocene-Miocene (38-13million years ago), which are considered two separate species complexes: P. sphenops and P. mexicana, which can also be distinguished by their tricuspid and unicuspid inner jaw teeth, respectively. Ultimately, within lineage diversification occurred mainly during the Miocene (22-5million years ago). All major cladogenetic events predated the final closure of the Isthmus of Panama. The allopatric distribution of lineages together with the long basal internodes suggest that vicariance and long term isolations could be the main evolutionary forces promoting radiation in this group, although dispersal through water barriers might also have occurred. Lastly, our results suggest the need to review the current species distribution and taxonomy of the P. sphenops complex sensu lato.

Relative host body condition and food availability influence epidemic dynamics: a Poecilia reticulata-Gyrodactylus turnbulli host-parasite model.

Understanding disease transmission is important to species management and human health. Host body condition, nutrition and disease susceptibility interact in a complex manner, and while the individual effects of these variables are well known, our understanding of how they interact and translate to population dynamics is limited. Our objective was to determine whether host relative body condition influences epidemic dynamics, and how this relationship is affected by food availability. Poecilia reticulata (guppies) of roughly similar size were selected and assembled randomly into populations of 10 guppies assigned to 3 different food availability treatments, and the relative condition index (Kn) of each fish was calculated. We infected 1 individual per group ('source' fish) with Gyrodactyus turnbulli and counted parasites on each fish every other day for 10 days. Epidemic parameters for each population were analysed using generalized linear models. High host Kn-particularly that of the 'source' fish-exerted a positive effect on incidence, peak parasite burden, and the degree of parasite aggregation. Low food availability increased the strength of the associations with peak burden and aggregation. Our findings suggest that host Kn and food availability interact to influence epidemic dynamics, and that the condition of the individual that brings the parasite into the host population has a profound impact on the spread of infection.