Learned mate recognition and reproductive isolation in guppies

Research on learned species discrimination has focused on the consequences of early experience. However, in species where parental care is limited or absent, including most fish, juveniles have fewer opportunities to learn from adult conspecifics. We examined male mate recognition in Trinidadian guppies, Poecilia reticulata, and in their sister species, the swamp guppy, P. picta. Choice tests revealed that males from localities where their species is the only poeciliid present initially mated with conspecific and heterospecific females at random. In contrast, P. reticulata and P. picta found in sympatry preferred their own females. We then investigated the acquisition of mating discrimination by wild P. reticulata males from two allopatric populations. Males that were allowed to interact with females of both species learned within 4 days to distinguish conspecific partners, and within a week their species discrimination matched that of sympatric populations. This study confirms that learning is important in the acquisition of adult mating preferences and shows why learned mate preferences can be important in the last stages of speciation.     

Gyrodactylus pictae n. sp. (Monogenea: Gyrodactylidae) from the Trinidadian swamp guppy Poecilia picta Regan, with a discussion on species of Gyrodactylus von Nordmann, 1832 and their poeciliid hosts

Gyrodactylus pictae n. sp. is recorded from Poecilia picta in heterospecific shoals with the guppy P.reticulata in Northern Trinidad. G. pictae is morphologically similar to G. turnbulli Harris, 1986, but the hamuli and marginal hooks are slightly smaller and more gracile. The toe and the point of the marginal hook have a distinctly different shape, providing the best morphological characters for distinguishing the two species. The rDNA ITS1 and ITS2 sequences differ from those of G. turnbulli (the closest relative) by >5, suggesting that these two taxa are not sibling species. The origin of the two species on poeciliids of the subgenus Micropoecilia is discussed, and it is suggested that this may represent a case of host–parasite co-evolution.     

Poecilia picta,a Close Relative to the Guppy,Exhibits Red Male Coloration Polymorphism: A System for Phylogenetic Comparisons

Studies on the evolution of female preference and male color polymorphism frequently focus on single species since traits and preferences are thought to co-evolve. The guppy, Poecilia reticulata, has long been a premier model for such studies because female preferences and orange coloration are well known to covary, especially in upstream/downstream pairs of populations. However, focused single species studies lack the explanatory power of the comparative method, which requires detailed knowledge of multiple species with known evolutionary relationships. Here we describe a red color polymorphism in Poecilia picta, a close relative to guppies. We show that this polymorphism is restricted to males and is maintained in natural populations of mainland South America. Using tests of female preference we show female P. picta are not more attracted to red males, despite preferences for red/orange in closely related species, such as P. reticulata and P. parae. Male color patterns in these closely related species are different from P. picta in that they occur in discrete patches and are frequently Y chromosome-linked. P. reticulata have an almost infinite number of male patterns, while P. parae males occur in discrete morphs. We show the red male polymorphism in P. picta extends continuously throughout the body and is not a Y-linked trait despite the theoretical prediction that sexually-selected characters should often be linked to the heterogametic sex chromosome. The presence/absence of red male coloration of P. picta described here makes this an ideal system for phylogenetic comparisons that could reveal the evolutionary forces maintaining mate choice and color polymorphisms in this speciose group.     

Can Mixed-Species Groups Reduce Individual Parasite Load? A Field Test with Two Closely Related Poeciliid Fishes (Poecilia reticulata and Poecilia picta)

Predation and parasitism are two of the most important sources of mortality in nature. By forming groups, individuals can gain protection against predators but may increase their risk of being infected with contagious parasites. Animals might resolve this conflict by forming mixed-species groups thereby reducing the costs associated with parasites through a relative decrease in available hosts. We tested this hypothesis in a system with two closely related poeciliid fishes (Poecilia reticulata and Poecilia picta) and their host-specific monogenean ectoparasites (Gyrodactylus spp.) in Trinidad. Fish from three different rivers were sampled from single and mixed-species groups, measured and scanned for Gyrodactylus. The presence and abundance of Gyrodactylus were lower when fish of both species were part of mixed-species groups relative to single-species groups. This is consistent with the hypothesis that mixed-species groups provide a level of protection against contagious parasites. We discuss the importance of potentially confounding factors such as salinity and individual fish size.     

Length‐weight relationships of four freshwater fish species from the coastal drainage system in Peru

The morphometric relationships of three native and one exotic freshwater fish species from the Lower San Juan and Lower Pisco River basins, central Coast of Peru, are presented. Specimens were collected in May and November 2010 using seine nets and electrofishing. Length‐Weight (LWR) relationships for Andinoacara stalsbergi, Trichomycterus punctulatus, Basilichthys archaeus and Poecilia reticulata are provided for the first time, contributing information towards the effort to conserve freshwater fishes.     

Effects of salinity on the survival and aggression of the invasive Rio Grande cichlid (Herichthys cyanoguttatus)

The spread of non-native Rio Grande cichlids (Herichthys cyanoguttatus) in southeast Louisiana includes brackish habitats. We studied the effects of three different salinity levels on the biology of juvenile H. cyanoguttatus for 13 months to determine the potential of this species to spread through local estuaries. The highest salinity tolerated was 30.0 psu, and these fish did not survive acclimation to the 32 psu treatment. Fish in brackish conditions grew slower than fish in freshwater conditions, indicating a potential long-term detriment to juvenile fish living in brackish conditions. Aggression levels were notably higher for fish in brackish (15 psu) vs. freshwater conditions. This persisted through the entire experiment even after acclimation back to freshwater. This study indicates that higher salinity habitats in Louisiana can be tolerated by this species. It also raises a question about the effect of higher salinities on aggressive behavior.     

Comparing the low‐salinity tolerance of Ulva species distributed in different environments

The green macroalgal genus Ulva (Ulvales, Ulvophyceae, Chlorophyta) is distributed worldwide from marine to freshwater environments. Comparative analyses of hyposalinity tolerance among marine, brackish, and freshwater Ulva species were performed by fluorescein diacetate viability counts. The subtidal marine species Ulva sp., collected from a depth of 30 m, showed the poorest tolerance to low salinity. This species died in 5 practical salinity units (PSU) artificial seawater or freshwater within 1 day. Its closely related species U. linza L. (an intertidal species) and U. prolifera Müller (a brackish species) showed varying tolerances to low salinity. After 7 days of freshwater exposure, the viability of U. linza L. decreased to approximately 20%, while U. prolifera Müller showed nearly 100% viability. The freshwater species U. limnetica Ichihara et Shimada, not yet found in coastal areas, was highly viable in seawater.     

Moderate salinity improves the availability of soil P by regulating P-cycling microbial communities in coastal wetlands

Accelerated sea-level rise is expected to cause the salinization of freshwater wetlands, but the responses to salinity of the availability of soil phosphorus (P) and of microbial genes involved in the cycling of P remain unexplored. We conducted a field experiment to investigate the effects of salinity on P cycling by soil microbial communities and their regulatory roles on P availability in coastal freshwater and brackish wetlands. Salinity was positively correlated with P availability, with higher concentrations of labile P but lower concentrations of moderately labile P in the brackish wetland. The diversity and richness of microbial communities involved in P cycling were higher in the brackish wetland than the freshwater wetland. Salinity substantially altered the composition of the P-cycling microbial community, in which those of the brackish wetland were separated from those of the freshwater wetland. Metagenomic sequence analysis indicated that functional genes involved in the solubilization of inorganic P and the subsequent transport and regulation of P were more abundant in coastal soils. The relative abundances of most of the target genes differed between the wetlands, with higher abundances of P-solubilization (gcd and ppa) and -mineralization (phoD, phy, and ugpQ) genes and lower abundances of P-transport genes (pstB, ugpA, ugpB, ugpE, and pit) in the brackish wetland. A significant positive correlation between the concentration of labile P and the abundances of the target genes suggested that salinity may, at least in part, improve P availability by regulating the P-cycling microbial community. Our results suggest that the P-cycling microbial community abundance and P availability respond positively to moderate increases in salinity by promoting the microbial solubilization and mineralization of soil P. Changes in microbial communities and microbially mediated P cycling may represent microbial strategies to adapt to moderate salinity levels, which in turn control soil function and nutrient balance.     

The salinity preference of members of the genus Macroplea (Coleoptera, Chrysomelidae, Donaciinae), fully aquatic leaf beetles that occur in brackish water

Reed beetles (Donaciinae) of the genus Macroplea Samouelle, 1819 live permanently submerged. Literature indicates that Macroplea mutica occurs in brackish water, whereas Macroplea appendiculata is restricted to freshwater. The salinity preference of these two species was tested in a linear and a circular device that offered a continuous salinity gradient. The distribution of animals in the devices was monitored over at least 3 h in each of the 21 experiments. Both species preferred freshwater (salinity 0) over brackish water (salinity 10). In particular, this holds true for specimens collected in brackish water. Likewise, immediate reactions could be observed when during such experiments the direction of the gradient was reversed. While M. mutica can be regarded as a truly marine insect, this marine environment does not strictly reflect its fundamental niche with respect to salinity preference. This is in line with accumulating evidence that M. mutica can be found in freshwater habitats (and M. appendiculata in brackish water). This indicates that the species’ distribution might be influenced by other factors like host plant preference or dispersal mechanisms. It is discussed if—in spite of similar fundamental niches—differences in salinity tolerance (and hence performance in brackish water) may have contributed to speciation in the genus Macroplea.     

Novel microsatellites from the green swordtail (Xiphophorus hellerii) also display polymorphism in guppy (Poecilia reticulata)

The green swordtail (Xiphophorus hellerii; Poeciliidae) is a popular ornamental freshwater fish species. Fourteen microsatellites were isolated from a green swordtail genomic library enriched for CA‐repeats. Thirteen microsatellites were polymorphic in green swordtail; interestingly four of them were tetranucleotide‐type. Cross‐species amplification showed that 10 of the 14 microsatellites were polymorphic in guppy (Poecilia reticulata; Poeciliidae) as well. No significant directional difference of allele length was seen between the two species at any of the loci tested.     

Comparative phylogeography and species boundaries in Echinolittorina snails in the central Indo-West Pacific

Aim This study aimed to test monophyly and geographical boundaries in five marine intertidal snail species from the central Indo‐West Pacific. We tested the prediction that phylogenetic breaks between the Indian and Pacific Ocean basins should be more pronounced in continental than oceanic settings, and sought common geographical patterns of interspecific boundaries and intraspecific phylogenetic breaks in the region. Location The tropical seas of the Indo‐West Pacific. Methods We sequenced over 1200 bp of the mitochondrial cytochrome oxidase subunit I gene (COI) from 18–92 individuals sampled from throughout the ranges of each of five species of Echinolittorina (Littorinidae): three members of the Echinolittorina trochoides species complex; Echinolittorina reticulata; and Echinolittorina vidua, together with sister species, in order to test species boundaries. In addition, 630 bp of the nuclear 28S rRNA gene were sequenced from E. reticulata and its sister Echinolittorina millegrana. Phylogenetic structure was assessed using neighbour‐joining and parsimony analyses. Results COI data confirmed species boundaries and geographical distributions for all species except the pair E. reticulata and E. millegrana, which were nevertheless reciprocally monophyletic for 28S rRNA. The species from ecologically ‘continental’ habitats (E. trochoides A and E. vidua, but not E. trochoides B) mostly showed strong interoceanic breaks (with age estimates 0.58–4.4 Ma), while the ecologically ‘oceanic’E. trochoides D and E. reticulata did not. The sister species E. trochoides A and B occupy the shores of the continental shelves of Southeast Asia and Australasia respectively; between them lies the oceanic ‘eastern Indonesian corridor’ occupied by E. trochoides D and E. reticulata. The widespread continental species E. vidua showed a complex pattern of deep division into six haplotype clades with apparently parapatric distributions. Main conclusions Our results show that ecological differences (in this case continental vs. oceanic habitat) influence both intraspecific phylogenetic structure and interspecific boundaries in these snails of intertidal rocky shores. Two of the three species restricted to continental shelves show phylogenetic breaks between the Indian and Pacific Oceans, consistent with vicariant separation during Plio‐Pleistocene low sea levels. The two oceanic species do not show breaks, suggesting that they maintained interoceanic connections through the eastern Indonesian corridor. The geographical location of the interspecific boundary between continental E. trochoides A and oceanic E. trochoides D mirrors intraspecific breaks reported in other species. The sister relationship of E. trochoides A and B in Asia and Australasia, respectively, is an example of a ‘marine Wallace's line’ distribution, and we suggest that it is the result of separation of two continental species by a barrier of unsuitable oceanic habitat.     

An experimental test of antagonistic effects of competition and parasitism on host performance in semi‐natural mesocosms

The mechanisms by which parasites can mediate the interactions between species have received increased interest in recent years. Nonetheless, most research has focused on the role of shared parasites as mediators of interspecific competition. Here, we explore the relative effects of Gyrodactylus specialist ectoparasites of Trinidadian guppies Poecilia reticulata on competition between their host and juveniles of the killifish Rivulus hartii. In mesocosms that replicate natural streams, we exposed guppies to only competitors, to only parasites, to both parasites and competitors, or the absence of both. Consistent with previous studies, we found that female guppies grew significantly less where only Gyrodactylus were present, and this was regardless of infection status or parasite load. Surprisingly, this effect of Gyrodactylus on the growth of female guppies was greatly reduced when both parasites and competitors were present in the mesocosms. We conclude that guppies can mediate the effects of Gyrodactylus on competition with Rivulus, by adaptively fine‐tuning their phenotype when simultaneously facing multiple enemies.     

COMPETITION AND THE EVOLUTION OF AGGRESSIVE BEHAVIOR IN TWO SPECIES OF TERRESTRIAL SALAMANDERS

The effects of competition on the evolution of interspecific interference mechanisms were studied by comparing the aggressive behavior of two terrestrial salamander species from two localities that differ in the intensity of interspecific competition. Plethodon jordani and P. glutinosus are closely related, ecologically similar species that are sympatric at intermediate elevations in the southern Appalachian Mountains. Previous removal and transplant experiments showed that interspecific competition is more intense in the northeastern Great Smoky Mountains, where the species are narrowly sympatric, than in the nearby Balsam Mountains, where sympatry is broader. In laboratory encounters, P. glutinosus from the Great Smoky Mountains were more aggressive to heterospecific and conspecific intruders than were P. glutinosus from the Balsam Mountains. For P. jordani, however, the variation in interspecific and intraspecific aggressive behavior among individuals within populations was as great as the variation between populations. Alpha-selection (i.e., improved competitive ability by the acquisition of interspecific interference mechanisms) has occurred in populations of P. glutinosus under conditions of intense interspecific competition. The evolution of aggressive behavior appears to have been influenced by the intensity of intraspecific competition as well.     

The role of climate for the range limits of parapatric European land salamanders

Abrupt range limits of parapatric species may serve as a model system to understand the factors that determine species’ range borders. Theory suggests that parapatric range limits can be caused by abiotic conditions along environmental gradients, biotic interactions or a combination of both. Geographic ranges of the parapatric salamanders, Salamandra salamandra and S. atra, meet in small contact zones in the European Alps and to date, the cause of parapatry and the restricted range of S. atra remain elusive. We combine multivariate approaches and climatic data analysis to explore niche differentiation among the two salamanders with respect to the available climatic environment at their contact zones. Our purpose is to evaluate whether climatic conditions explain the species’ sharp range limits or if biotic interactions may play a role for range delimitation. Analyses were carried out in three contact zones in Switzerland to assess possible geographic variation. Our results indicate that both species occur at localities with different climatic conditions as well as the presence of a strong climatic gradient across the species’ range limits. Although the species’ climatic niches differ moderately (with a wider niche breadth for S. atra), interspecific niche overlap is found. Comparisons among the contact zones confirm geographic variation in the species’ climatic niches as well as in the conditions within the geographically available space. Our results suggest that the change in climatic conditions along the recognized gradient represents a determining factor for species’ range limits within contact zones. However, our analyses of geographic variation in climatic conditions reveal that both salamander species can occur in a much wider range of conditions than observed within contact zones. This finding and the interspecific climatic niche overlap within each contact zone provides indirect evidence that biotic interactions (likely competition) between the two species may also determine their range limits.     

Isolation and differentiation of Rivulus hartii across Trinidad and neighboring islands

Diversification of freshwater fishes on islands is considered unlikely because the traits that enable successful colonization—specifically, broad salinity tolerances and the potential for oceanic dispersal—may also constrain post‐colonization genetic differentiation. Some secondary freshwater fish, however, exhibit pronounced genetic differentiation and geographic structure on islands, whereas others do not. It is unclear what conditions give rise to contrasting patterns of differentiation because few comparative reconstructions of population history have been carried out for insular freshwater fishes. In this study, we examined the phylogeography of Hart’s killifish (Rivulus hartii) across Trinidad, with reference to neighboring islands and northern South America, to test hypotheses of colonization and differentiation derived from comparable work on co‐occurring guppies (Poecilia reticulata). Geographic patterns of mitochondrial DNA haplotype variation and microsatellite genotype variation provide evidence of genetic differentiation of R. hartii among islands and across Trinidad. Our findings are largely consistent with patterns of geographically structured ancestry and admixture found in Trinidadian guppies, which suggests that both species share a history of colonization and differentiation and that post‐colonization diversification may be more common among members of insular freshwater fish assemblages than has been previously thought.     

Growth pattern and survival in populations of Poecilia vivipara (Teleostei; Poeciliidae) inhabiting an environmental gradient: a common garden study

We performed a common garden experiment to assess the existence of genetic differences on growth and body size between two populations of Poecilia vivipara inhabiting extremes of an environmental gradient caused by water salinity in lagoons of Northern Rio de Janeiro State, Brazil: the Campelo lagoon (freshwater) and Açu lagoon (brackish/saltwater). The two populations show extreme differences in average phenotypes for body size, shape and life history (freshwater populations with smaller body size, lower fecundity and larger reproductive allotment). Pregnant females were brought to the lab and the offspring from both groups were kept in a common recirculating system with freshwater. Standard length and survival were measured weekly over a period of 200 days and growth models were fitted and selected with information criteria. The offspring originally from the brackish water lagoon presented larger asymptotic length, higher maximum growth rate but lower survival than the offspring originally from the freshwater lagoon. Potential confounding variables such as density differences due to mortality and maternal effects (offspring size) were included as covariates in comparisons of growth rates between groups. The results are consistent with phenotypic differences among populations having some genetic basis, and with the existence of a trade-off between growth and maintenance due to the high growth/low survival observed in the group that changed from salt to freshwater. Comparisons of captive and natural populations suggest that the influence of environmental factors, such as salinity, food availability, fish density and predation should also be considered relevant to explain phenotypic variation in this system.     

The biogeography of parasitism in sticklebacks: distance,habitat differences and the similarity in parasite occurrence and abundance

Similarity in parasite community composition often decreases with both increasing geographic distance and environmental dissimilarity between localities, though it is unknown whether similarity in local abundance of selected parasite species follows similar rules. We tested this using data on metazoan parasites in 126 stickleback (Gasterosteus aculeatus) populations, with locations from Eurasia, eastern North America, and western North America treated separately. Similarity values were regressed against pairwise distances between localities; after correcting for distance, the effect of environmental dissimilarity was assessed by splitting similarity values into those between pairs of localities with either similar, moderately different or very different salinity (freshwater, marine or brackish). For selected parasite species, pairwise similarity in abundance (mean no. parasites per host) were computed across all localities, and treated as above. Similarity in parasite community composition decreased with increasing distance between localities in all three geographic regions. A significant effect of environmental difference was found in all regions: for a given distance between two sites, their parasite communities were more similar if they were of the same salinity. Slopes for distance decay in similarity were consistently higher for eastern North America than for Eurasia. Among the 12 parasite species for which sufficient data were available, only 4 showed the expected relationship, i.e. the greater the geographical separation between host populations, the greater the difference in parasite abundance; also, significant effects of environmental differences in salinity were only found for 3 of these species. Our findings show that parasite communities of sticklebacks are structured by geographical distance and local salinity conditions. The results indicate that strong effects at the community level do not translate into corresponding effects at the population level, suggesting that parasite dispersal and population dynamics are controlled by different processes.     

Effect of salinity on cable bacteria species composition and diversity

Cable bacteria (CB) are Desulfobulbaceae that couple sulphide oxidation to oxygen reduction over centimetre distances by mediating electric currents. Recently, it was suggested that the CB clade is composed of two genera, Ca. Electronema and Ca. Electrothrix, with distinct freshwater and marine habitats respectively. However, only a few studies have reported CB from freshwater sediment, making this distinction uncertain. Here, we report novel data to show that salinity is a controlling factor for the diversity and the species composition within CB populations. CB sampled from a freshwater site (salinity 0.3) grouped into Ca. Electronema and could not grow under brackish conditions (salinity 21), whereas CB from a brackish site (salinity 21) grouped into Ca. Electrothrix and decreased by 93% in activity under freshwater conditions. On a regional scale (Baltic Sea), salinity significantly influenced species richness and composition. However, other environmental factors, such as temperature and quantity and quality of organic matter were also important to explain the observed variation. A global survey of 16S rRNA gene amplicon sequencing revealed that the two genera did not co-occur likely because of competitive exclusion and identified a possible third genus.     

Comparative size,biomass, elemental composition (C,N, H), and energy concentration of caridean shrimp eggs

Summary

In the evolution of decapod crustaceans, interspecific variation in egg size is considered as an important life-history trait that is linked with the duration of embryonic and larval development, the number and type of larval stages, and with juvenile size. Aiming to provide a quantitative characterization of reproductive traits in related decapod taxa differing in lifestyle (freshwater, estuarine, marine) and geographic-climatic distribution (tropical-temperate), we compared size, biomass, and elemental composition of eggs of caridean shrimps from three families: seven species of Palaemonidae (three congeners of Macrobrachium: M. olfersii, M. carcinus, M. acanthurus; four species of Palaemon: P. northropi, P. pandaliformis, P. elegans, P. adspersus), two Atyidae (Potimirim potimirim, Atya scabra), and one Pandalid (Pandalus montagui). Egg size was measured as larger and smaller diameter (D₁ D₂), volume was calculated from D₁ and D₂, and biomass was measured as dry mass (W), carbon (C), nitrogen (N), hydrogen (H), and energy (E, estimated from C) contents. The smallest size and lowest biomass were found in the eggs of two freshwater atyids (both originating from Brazil); the largest size occurred in a marine species, P. montagui (from the North Sea); and intermediate values in freshwater, estuarine, and marine palaemonid species (from Brazil and the Baltic Sea, respectively). Among the Palaemon species, the most limnic (P. pandaliformis) showed a significantly larger egg size and volume (P<0.001) than the estuarine and marine congeners, P. elegans, P. adspersus and P. northropi. This suggests that the generally postulated relationship between egg size and lifestyle (freshwater vs. estuarine or marine) may appear at a generic but not at the family level. On the other hand, individual biomass (in μg or Joules per egg) of early eggs was significantly higher in P. elegans and P. adspersus, indicating interspecific variability in biomass and energy concentration (in μg or Joules per unit volume, mm³). Generally lower biomass concentrations in early eggs of freshwater shrimps may be caused by a higher average water content. Eggs in late embryonic stages were generally larger than earlier eggs of the same species, reflecting an increase in the water content, while an increasing D₁:D₂ quotient indicated an increasingly elongated egg shape. The biomass per egg decreased during embryonic development due to metabolic degradation of organic reserves. As a consequence of inverse ontogenetic changes in size and organic biomass of developing eggs, the mass-specific biomass values (C, N, H in percent of W; E in Joules per mg W) and volume-specific concentrations (μg or Joules per unit volume) decreased. This change was consistently stronger in the C, H, and E contents than in other measures of biomass (W, N). In consequence, the C:N mass ratio also decreased, suggesting that lipid degradation rather than protein utilization was the principal fuel for embryonic development. Our results indicate high intra- (mainly developmental) and interspecific variation in reproductive traits of closely related species. While volume-specific biomass and energy concentrations of early eggs appear to be associated with variation in habitat salinity (freshwater, brackish, marine), individual egg size and biomass may be related more with the climatic-geographic distribution (temperate, tropical) of different taxa.     

Patterns of species assemblages and geographical distributions associated with mandible size differences in coastal tiger beetles in Japan

Nine species of tiger beetle (Cicindelidae) occur in coastal habitats in Japan, with two to four species co-occurring at each locality. To examine the patterns of coexistence and geographical distribution, the mandible size of co-occurring species at 17 localities in Japan was examined, based on the assumption that competition for food is an important factor in determining these patterns. The interspecific overlap of mandible length was absent or very low in localities with two or three species, whereas it was more or less evident in localities with four species. For four large coastal species, the geographical distributions of two species with similar mandible lengths are either allopatric or parapatric, whereas those of two species with different mandible lengths largely overlap. These results strongly suggest that size-assortment in mandible length is important in determining species assemblage and distribution in coastal tiger beetles in Japan.An erratum to this article can be found at