Kin structure and shoal composition dynamics in the guppy, Poecilia reticulata

Although kin associations between fish can develop in the laboratory, the importance of kin structure in natural social groups remains uncertain. Guppies ( Poecilia reticulata ) possess many behavioural and life-history traits that promote kin structure, and so constitute ideal candidates for testing its general significance amongst fish. We collected 25 adult only shoals from ∼150 m stretches along two Trinidadian rivers. The 272 sampled fish were genotyped from a pool of seven hypervariable microsatellite loci. Although mark-recapture experiments have previously detected male-biased dispersal, which is predicted from the guppy mating system and which may have important implications for population differentiation, we found no sex-biases in dispersal between shoals. Because sex-biased dispersal was not evident, males and females were pooled in analyses of kin structure. Relatedness levels within each shoal did not differ significantly from zero, indicating that kin selection does not contribute to gross structure. Instead, our data is more consistent with the hypothesis that individual-level selection underlies shoal composition dynamics. These dynamics have important implications for the evolution of cooperative behaviour in guppies.     

Intrinsic reproductive isolation between Trinidadian populations of the guppy, Poecilia reticulata

Although Trinidadian populations of the guppy, Poecilia reticulata, show considerable adaptive genetic differentiation, they have been assumed to show little or no reproductive isolation. We tested this assumption by crossing Caroni (Tacarigua River) and Oropuche (Oropuche R.) drainage populations from Trinidad's Northern Range, and by examining multiple aspects of reproductive compatibility in the F1, F2 and BC1 generations. In open-aquarium experiments, F1 males performed fewer numbers of mating behaviours relative to parental population controls. This is the first documentation of hybrid behavioural sterility within a species, and it suggests that such sterility may feasibly be involved in causing speciation. The crosses also uncovered hybrid breakdown for embryo viability, brood size and sperm counts. In contrast, no reductions in female fertility were detected, indicating that guppies obey Haldane's rule for sterility. Intrinsic isolation currently presents a much stronger obstacle to gene flow than behavioural isolation, and our results indicate that Trinidadian populations constitute a useful model for investigating incipient speciation.     

Male display rate reveals ejaculate characteristics in the Trinidadian guppy,Poecilia reticulata.

Previous work (Reynolds and Gross 1992) has demonstrated that mate choice enables female guppies, Poecilia reticulata, to obtain heritable benefits (''good genes'') for their offspring. It is not yet known whether males also signal their functional fertility, that is, their viability as fathers, to female guppies. Our study tested the hypothesis that a male''s behaviour reveals characteristics of his ejaculate. We uncovered a strong correlation between display rate and sperm number in male guppies derived from two wild Trinidadian populations. This result indicates that information on the functional fertility of potential mates is available to females. However, our investigation also found a significant relationship between the frequency of sneaky mating and sperm number. Female guppies in wild populations, particularly those experiencing a high predation regime, are constantly subjected to sneaky mating attempts. As the males responsible for these sneaky matings have highest reserves of sperm they may have the capacity to undermine female choice. The implications of these findings for sperm competition are discussed.     

Predator inspection,shoaling and foraging under predation hazard in the Trinidadian guppy,Poecilia reticulata

Synopsis Guppies,Poecilia reticulata, living in stream pools in Trinidad, West Indies, approached a potential fish predator (a cichlid fish model) in a tentative, saltatory manner, mainly as singletons or in pairs. Such behavior is referred to as predator inspection behavior. Inspectors approached the trunk and tail of the predator model more frequently, more closely and in larger groups than they approached the predator's head, which is presumably the most dangerous area around the predator. However, guppies were not observed in significantly larger shoals in the stream when the predator model was present. In a stream enclosure, guppies inspected the predator model more frequently when it was stationary compared to when it was moving, and made closer inspections to the posterior regions of the predator than to its head. Therefore, the guppies apparently regarded the predator model as a potential threat and modified their behavior accordingly when inspecting it. Guppies exhibited a lower feeding rate in the presence of the predator, suggesting a trade-off between foraging gains and safety against predation. Our results further suggest that predator inspection behavior may account for some of this reduction in foraging. These findings are discussed in the context of the benefits and costs of predator inspection behavior.     

Multiple selection pressures influence Trinidadian guppy (Poecilia reticulata) antipredator behavior

Animals alter their behavior to avoid a variety of differenttypes of predators. Trinidadian guppies (Poecilia reticulata)have been an important system for examining the evolution ofantipredator behavior because geographically isolated populationsexperience different amounts of aquatic predation. Althoughthe influence of aquatic predators has been well documented,selective pressures from other types of predators are less wellunderstood. We examined the response of wild-caught individualsfrom upstream and downstream populations to an aquatic predatorand a simulated aerial predator. As previously documented, fishfrom the downstream population responded more strongly to theaquatic predator than did fish from the upstream population,inspecting for longer periods of time. Guppies also exhibiteda strong behavioral response to the potential aerial predator.Although both populations displayed a similar magnitude of response,they differed in the specific behaviors used. Upstream fishtended to freeze on the tank bottom, whereas downstream fishtended to hide under shelter. Field observations suggest thatthese strategies are related to habitat features specific toeach site. The behaviors used against aerial predators differedsubstantially from the behaviors used against aquatic predators,suggesting that different types of predators exert conflictingselection pressures. This research demonstrates the importanceof considering multiple selection pressures acting on an organismwhen trying to understand the evolutionary history of behavioraland morphological traits.     

Isolation and characterization of polymorphic microsatellites in the Trinidadian guppy (Poecilia reticulata)

We isolated microsatellite loci for the Trinidadian guppy Poecilia reticulata from the Oropuche and Caroni drainages in Trinidad, using a modified enrichment protocol that provided 93% enriched libraries. The eight loci presented here are polymorphic, and have between seven and 18 alleles in 33–48 individuals initially screened. Observed heterozygosities in this sample range from 0.28 to 0.88. These loci significantly increase the number of polymorphic microsatellite markers available for paternity and population analysis in this species.     

Cross-generational environmental effects and the evolution of offspring size in the Trinidadian guppy Poecilia reticulata

Abstract The existence of adaptive phenotypic plasticity demands that we study the evolution of reaction norms, rather than just the evolution of fixed traits. This approach requires the examination of functional relationships among traits not only in a single environment but across environments and between traits and plasticity itself. In this study, I examined the interplay of plasticity and local adaptation of offspring size in the Trinidadian guppy, Poecilia reticulata. Guppies respond to food restriction by growing and reproducing less but also by producing larger offspring. This plastic difference in offspring size is of the same order of magnitude as evolved genetic differences among populations. Larger offspring sizes are thought to have evolved as an adaptation to the competitive environment faced by newborn guppies in some environments. If plastic responses to maternal food limitation can achieve the same fitness benefit, then why has guppy offspring size evolved at all? To explore this question, I examined the plastic response to food level of females from two natural populations that experience different selective environments. My goals were to examine whether the plastic responses to food level varied between populations, test the consequences of maternal manipulation of offspring size for offspring fitness, and assess whether costs of plasticity exist that could account for the evolution of mean offspring size across populations. In each population, full‐sib sisters were exposed to either a low‐ or high‐food treatment. Females from both populations produced larger, leaner offspring in response to food limitation. However, the population that was thought to have a history of selection for larger offspring was less plastic in its investment per offspring in response to maternal mass, maternal food level, and fecundity than the population under selection for small offspring size. To test the consequences of maternal manipulation of offspring size for offspring fitness, I raised the offspring of low‐ and high‐food mothers in either low‐ or high‐food environments. No maternal effects were detected at high food levels, supporting the prediction that mothers should increase fecundity rather than offspring size in noncompetitive environments. For offspring raised under low food levels, maternal effects on juvenile size and male size at maturity varied significantly between populations, reflecting their initial differences in maternal manipulation of offspring size; nevertheless, in both populations, increased investment per offspring increased offspring fitness. Several correlates of plasticity in investment per offspring that could affect the evolution of offspring size in guppies were identified. Under low‐food conditions, mothers from more plastic families invested more in future reproduction and less in their own soma. Similarly, offspring from more plastic families were smaller as juveniles and female offspring reproduced earlier. These correlations suggest that a fixed, high level of investment per offspring might be favored over a plastic response in a chronically low‐resource environment or in an environment that selects for lower reproductive effort     

Direct and indirect ecosystem effects of evolutionary adaptation in the Trinidadian guppy (Poecilia reticulata)

Ecological and evolutionary processes may interact on the same timescale, but we are just beginning to understand how. Several studies have examined the net effects of adaptive evolution on ecosystem properties. However, we do not know whether these effects are confined to direct interactions or whether they propagate further through indirect ecological pathways. Even less well understood is how the combination of direct and indirect ecological effects of the phenotype promotes or inhibits evolutionary change. We coupled mesocosm experiments and ecosystem modeling to evaluate the ecological effects of local adaptation in Trinidadian guppies (Poecilia reticulata). The experiments show that guppies adapted to life with and without predators alter the ecosystem directly through differences in diet. The ecosystem model reveals that the small total indirect effect of the phenotype observed in the experiments is likely a combination of several large indirect effects that act in opposing directions. The model further suggests that these indirect effects can reverse the direction of selection that direct effects alone exert back on phenotypic variation. We conclude that phenotypic divergence can have major effects deep in the web of indirect ecological interactions and that even small total indirect effects can radically change the dynamics of adaptation.     

Exocrine pancreatic carcinogenesis in the guppy Poecilia reticulata

Exocrine pancreatic neoplasms developed in the guppy Poecilia reticulata following exposure to the direct-acting carcinogen methylazoxymethanol acetate (MAM-Ac). Fish 6 to 10 d old were exposed to nominal, non-toxic concentrations of 4 and 10 mg MAM-Ac l(-1) for 2 h and then transferred to carcinogen-free water for grow-out. Whole specimens were sampled monthly up to 9 mo post-exposure to follow the histologic progression of the lesions. No neoplasms occurred in 119 control specimens examined. Pancreatic acinar cell adenomas and carcinomas occurred in 42 of 243 (17%) of the specimens exposed to MAM-Ac. As in earlier studies, specimens exposed to the low MAM-Ac concentration exhibited a higher pancreatic neoplasm incidence (27.8%) than those exposed to the high concentration (7.8%). Acinar cell adenomas accounted for 27 of the 42 neoplasms. Adenomas exhibited a high degree of acinar cell differentiation and some contained foci of atypical acinar cells that were less differentiated and more basophilic than were surrounding adenoma cells. Carcinomas occurred in 15 specimens and exhibited a range of cellular patterns. Although no distant metastases were found, carcinomas tended to invade neighboring tissues and organs. The occurrence of carcinogen-induced pancreatic neoplasms in guppies strengthens the usefulness of small fish species in carcinogen testing and provides an additional model for studying pancreatic neoplasia.     

Salinity tolerance of the guppy, Poecilia reticulata Peters

Experiments were conducted to determine the adaptability of the guppy to various salt concentrations. The guppy, Poecilia reticulata , (total length 11–40 mm) were subjected to abrupt and gradual changes from fresh water (salinity=0.1%) to various salinities (%). No mortality occurred when the fish were transferred from fresh water to 50% sea water (19.5%). Through gradual adaptation from 50% sea water to 80% of the fish were able to tolerate 100% (39%) sea water for 7 days. After a 7 day stay in sea water, fish were readapted to fresh water during a 3 h period. Through gradual adaptation fish were also able to tolerate salinities ranging from 39.0% (100% sea water) to 58.5%. After a 30 day stay in 150% sea water (58.5%), fish were readapted to fresh water over a 5 h period. The results indicate that they were well able to tolerate the abrupt change from 100 or 150% sea water to fresh water. Females that stayed in 150% sea water (58.5%) for 30 days had embryos in their gonads.     

Predator prey interactions: predator choice for prey group size and composition in Crenicichla alta, a natural predator of the Trinidadian guppy, Poecilia reticulata

Young salmonids may use substratum as hiding stations and/or shelter and they depend on invertebrates, which develop on substratum, for their feeding. For several decades, human activities have contributed to increase siltation in streams, and negative consequences on trout production have sometimes been highlighted. In the research devoted to the understanding of that negative effect, most studies have focused on embryo‐larval survival, and consequences of substrate embeddedness on later stages have rarely been investigated. In the present work we attempt at studying the impact of embeddedness on brown trout juveniles. In an experimental channel, trout growth was compared in embedded and non‐embedded sections. Growth was reduced with embeddedness due to change in trophic conditions and/or in habitat. To investigate the direct role of substratum for fish, trouts behaviour was observed from an under water observation room in two cages offering embedded and non‐embedded substrate conditions but similar trophic conditions. Competition appeared heavier in the embedded cage where dominated fishes stayed almost motionless. The effect of substratum quality on intra‐specific competition is discussed in relation with visual isolation and territory size.     

Aerial Jumping in the Trinidadian Guppy (Poecilia reticulata)

Many fishes are able to jump out of the water and launch themselves into the air. Such behavior has been connected with prey capture, migration and predator avoidance. We found that jumping behavior of the guppy Poecilia reticulata is not associated with any of the above. The fish jump spontaneously, without being triggered by overt sensory cues, is not migratory and does not attempt to capture aerial food items. Here, we use high speed video imaging to analyze the kinematics of the jumping behavior P. reticulata. Fish jump from a still position by slowly backing up while using its pectoral fins, followed by strong body trusts which lead to launching into the air several body lengths. The liftoff phase of the jump is fast and fish will continue with whole body thrusts and tail beats, even when out of the water. This behavior occurs when fish are in a group or in isolation. Geography has had substantial effects on guppy evolution, with waterfalls reducing gene flow and constraining dispersal. We suggest that jumping has evolved in guppies as a behavioral phenotype for dispersal.     

Marked variation in parasite resistance between two wild populations of the Trinidadian guppy, Poecilia reticulata (Pisces: Poeciliidae)

This is the first study to demonstrate significant differences between two natural Trinidadian guppy populations in susceptibility to a pathogenic monogenean parasite, Gyrodactylus turnbulli . Following experimental infection with an isogenic laboratory culture of G. turnbulli , fish from the Upper Aripo (UA) lost parasites more slowly and carried up to three times as many parasites at peak infection than did those from the Lower Aripo (LA). The UA population appeared to be more susceptible than the LA fish, even though fish of both populations were naïve to this particular laboratory strain of G. turnbulli and had not encountered any gyrodactylid infection for at least 65 days. The parasite infection reduced the feeding response and feeding activity of UA and LA fish to a similar extent. Our findings suggest that this ectoparasite may have an important impact on the evolutionary biology of guppies (and possibly of other teleosts), particularly as the prevalence of Gyrodactylus infection in natural populations may be as high as 50%. We discuss the role of parasite infections on natural and sexual selection, the good genes model and the implications for immunogenetics in small genetically isolated host populations.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79 , 645–651.     

Phenotypic correlates of spermatozoon quality in the guppy, Poecilia reticulata

The phenotype-linked fertility hypothesis suggests that a femalecan benefit directly from mate choice when the cues she usesindicate the quantity and/or quality of his spermatozoa. Wetested the link between sperm quality and male body size andcoloration in the resource-free mating system of the guppy,a tropical fish characterized by strong female choice. Largermales possessed larger testes and are therefore predicted toproduce larger numbers of spermatozoa than smaller males. Largermale guppies also produced longer spermatozoa than smaller males.Degree of carotenoid coloration did not predict either the quantityor the quality of a male's spermatozoa. These results are consistentwith a previous study that showed that female guppies in thestudy population prefer larger males to brightly colored males.The male-size directed increase in spermatozoon size may bethe result of interplay between sperm competition and the coevolutionof spermatozoon traits with the female reproductive tract.     

Schooling preferences for familiar fish in the guppy, Poecilia reticulata

Although schooling behaviour is an important antipredator response of guppies ( Poecilia reticulata ) in Trinidad it is not known whether fish elect to school with particular individuals. Familiarity is a simple mechanism that could lead to non-random associations within schools. In this paper we test the idea that prior experience of particular individuals influences the choice of partners during schooling. We show (in a laboratory study) that female guppies prefer to associate with familiar individuals. However, we are unable to detect any schooling preference for unfamiliar females from the same population.     

Intergenerational effects of CO2‐induced stream acidification in the Trinidadian guppy (Poecilia reticulata)

Rising atmospheric carbon dioxide levels are driving decreases in aquatic pH. As a result, there has been a surge in the number of studies examining the impact of acidification on aquatic fauna over the past decade. Thus far, both positive and negative impacts on the growth of fish have been reported, creating a disparity in results. Food availability and single‐generation exposure have been proposed as some of the reasons for these variable results, where unrealistically high food treatments lead to fish overcoming the energetic costs associated with acclimating to decreased pH. Likewise, exposure of fish to lower pH for only one generation may not capture the likely ecological response to acidification that wild populations might experience over two or more generations. Here we compare somatic growth rates of laboratory populations of the Trinidadian guppy (Poecilia reticulata) exposed to pH levels that represent the average and lowest levels observed in streams in its native range. Specifically, we test the role of maternal acclimation and resource availability on the response of freshwater fishes to acidification. Acidification had a negative impact on growth at more natural, low food treatments. With high food availability, fish whose mothers were acclimated to the acidified treatment showed no reduction in growth, compared to controls. Compensatory growth was observed in both control–acidified (maternal–natal environment) and acidified–control groups, where fish that did not experience intergenerational effects achieved the same size in response to acidification as those that did, after an initial period of stunted growth. These results suggest that future studies on the effects of shifting mean of aquatic pH on fishes should take account of intergenerational effects and compensatory growth, as otherwise effects of acidification may be overestimated.     

Inheritance of the snakeskin color pattern in the guppy, Poecilia reticulata

We made single-pair reciprocal crosses between the Green Snakeskin and Yellow Snakeskin domesticated strains of the guppy, Poecilia reticulata. The two snakeskin strains differ by a single autosomal gene, with the Green Snakeskin strain having the wild-type background coloration caused by the dominant gene (B), whereas the Yellow Snakeskin is homozygous for the recessive blond allele (bb). The snakeskin body and tail patterns characterizing males of these two strains are determined by two genes--Ssb and Sst--that are closely linked on the Y chromosome. The greenish-yellow tail color of the Green Snakeskin strain is mediated by an X-linked dominant gene, Grt. The recessive wild-type allele, Grt+, gives the hyaline tail color. In the Yellow Snakeskin strain, the Grt gene is expressed as a golden-yellow color as a result of the presence of the bb homozygous condition. The putative genotypes of the males and females of the Green Snakeskin strain are BB XGrt YSsb,Sst and BB XGrt XGrt, respectively. Males and females of the Yellow Snakeskin strain have the putative genotypes bb XGrt YSsb,Sst and bb XGrtXGrt, respectively. As a result of crossing over between the X and Y chromosomes, a few males and females of these two snakeskin strains may carry one or both snakeskin pattern genes (Ssb and Sst) on the X chromosome.     

Identification of major histocompatibility complex genes in the guppy,Poecilia reticulata

The guppy, Poecilia reticulata, a teleostean fish of the order Cyprinodontiformes, has been used extensively in studies of host-parasite interactions, courtship behavior, and mating preference, as well as in ecological and evolutionary genetics. A related species was among the first poikilotherm vertebrates to be used in the study of histocompatibility genes. All these studies could benefit from the identification and characterization of the guppy major histocompatibility complex (Mhc) genes. Here, both class I and class II genes of the guppy are described. The number of expressed loci, as determined by representation of clones in a cDNA library, sequencing, and Southern blot analysis, may be low in both Mhc classes: combined evidence suggests that there may be one expressed class II locus only and one or two expressed class I loci. The variability of aquaristic guppy stocks is very low: only three and two genes have been detected at the class I and class II loci, respectively, in the stocks examined. This genetic paucity is most likely the consequence of breeding practices employed by aquarists and commercial establishments. Limited sampling of wild guppy populations revealed extensive Mhc polymorphism at loci of both classes in nature. Comparison of guppy Mhc sequences with those of other vertebrates has revealed the existence of a set of insertions/deletions which can be used as characters in cladistic analysis to infer phylogenetic relationships among vertebrate taxa and the Mhc genes themselves. These indels are particularly frequent in the regions coding for the loops of 1 and 2 domains of class I proteins.The nucleotide sequence data reported in this publication have been submitted to the EMBL nucleotide sequence database and have been assigned the accession numbers Z54076-Z54095     

Recognition of chemical stimuli by guppy, Poecilia reticulata Peters

An alimentary conditioned reflex to beta-phenylethanol and a subsequent differentiation of cumarine were elaborated in conditions of simultaneous paired choice in guppies. It was shown that beta-phenylethanol in a concentration not over 7 X 10(-6) M is perceived through olfaction. The conditioned reflex persists for a long time: twenty five to fifty days after section of the olfactory nerves (the conditioned reflex is restored after their regeneration) and at least seventy days in intact fishes.     

Two-phase exploration of a novel environment in the guppy, Poecilia reticulata

Swimming speed and percentage of time fish swam along the boundary of an aquarium gradually decreased with the time following release of juvenile guppies into unfamiliar surroundings. When swimming speed was plotted against time on a log scale, two distinct phases were identified. The high activity phase lasted c, 3 h in 30-day-old guppies, 6 h in 10-day-olds and not less than S h in 3-day-olds. Observed trends in swimming in guppies are interpreted as a component of exploratory behaviour. Alternating the surroundings confirmed the exploratory role of such swimming and suggested that a period of general exploration was followed by a period of more detailed investigation of habitat. An interaction between exploratory and avoidance (fear) responses is discussed.