The metabolic, locomotor and sex-dependent effects of elevated temperature on Trinidadian guppies: limited capacity for acclimation.

Global warming poses a threat to many ectothermic organisms because of the harmful effects that elevated temperatures can have on resting metabolic rate (RMR) and body size. This study evaluated the thermal sensitivity of Trinidadian guppies (Poecilia reticulata) by describing the effects of developmental temperature on mass, burst speed and RMR, and investigated whether these tropical fish can developmentally acclimate to their thermal conditions. These traits were measured following exposure to one of three treatments: 70 days at 23, 25, 28 or 30°C (acclimated groups); 6 h at 23, 28 or 30°C following 70 days at 25°C (unacclimated groups); or 6 h at 25°C following 70 days in another 25°C tank (control group). Body mass was lower in warmer temperatures, particularly amongst females and individuals reared at 30°C. The burst speed of fish acclimated to each temperature did not differ and was marginally higher than that of unacclimated fish, indicative of complete compensation. Conversely, acclimated and unacclimated fish did not differ in their RMR at each temperature. Amongst the acclimated groups, RMR was significantly higher at 30°C, indicating that guppies may become thermally limited at this temperature as a result of less energy being available for growth, reproduction and locomotion. Like other tropical ectotherms, guppies appear to be unable to adjust their RMR through physiological acclimation and may consequently be susceptible to rising temperatures. Also, because larger females have higher fecundity, our data suggest that fecundity will be reduced in a warmer climate, potentially decreasing the viability of guppy populations.

Temporal variation at the MHC class IIb in wild populations of the guppy (Poecilia reticulata).

Understanding genetic diversity in natural populations is a fundamental objective of evolutionary biology. The immune genes of the major histocompatibility complex (MHC) are excellent candidates to study such diversity because they are highly polymorphic in populations. Although balancing selection may be responsible for maintaining diversity at these functionally important loci, temporal variation in selection pressure has rarely been examined. We examine temporal variation in MHC class IIB diversity in nine guppy (Poecilia reticulata) populations over two years. We found that five of the populations changed significantly more at the MHC than at neutral (microsatellite) loci as measured by F(ST), which suggests that the change at the MHC was due to selection and not neutral processes. Additionally, pairwise population differentiation measures at the MHC were higher in 2007 than in 2006, with the signature of selection changing from homogenizing to diversifying selection or neutral evolution. Interestingly, within the populations the magnitude of the change at the MHC between years was related to the change in the proportion of individuals infected by a common parasite, indicating a link between genetic structure and the parasite. Our data thereby implicate temporal variation in selective pressure as an important mechanism maintaining diversity at the MHC in wild populations.

Inter-population variation in multiple paternity and reproductive skew in the guppy.

We use microsatellite loci to detail the multiple paternity patterns in broods from 10 wild populations of the guppy (Poecilia reticulata) found in Northern Trinidad. The populations span two major drainages comprising the Caroni and the Oropouche, and include sites that are characterized by either high or low predation. Across the populations the frequency of multiple paternity is high with 95% (range: 70%-100%) of broods having multiple sires. Broods have an average of 3.5 sires (range: 1-9) and a mixed-model analysis suggests that broods from high predation sites have marginally more sires than do those from low predation sites, but this is true only in the Oropouche drainage. There is no difference in sire number between predation sites in the Caroni drainage. Brood size, but not female body length, is correlated with the number of sires and the correlation cannot be attributed solely to the stochastic process associated with sperm competition and a 'fair raffle'. Within broods there is significant skew in reproductive success among males, which may reflect variation in sperm competitiveness or female choice. There is, however, no difference in the skew among populations from different predation regimes or drainages. Finally, high predation populations were characterized by increased genetic variability at the microsatellite loci, suggesting a larger effective population size. We discuss explanations for the high degree of multiple paternity but the general lack of any major differences among broods from ecologically different populations.

Multiple paternity and kin recognition mechanisms in a guppy population.

Help directed toward kin (nepotism) is an important example of social behaviour. Such helping behaviour requires a mechanism to distinguish kin from nonkin. The prevailing kin recognition hypothesis is that when familiarity is a reliable cue of relatedness, other mechanisms of recognition will not evolve. However, when familiarity is an unreliable cue of relatedness, kin recognition by phenotype matching is instead predicted to evolve. Here we use genetic markers to show that guppies (Poecilia reticulata) from a population in a tributary of the Paria River in Trinidad are characterized by a high degree of multiple mating with 95% of broods having more than one sire and some dams having offspring sired by six males. These levels of multiple mating are the highest reported among live-bearing fishes. The mean relatedness of brood-mates was 0.36 (as compared to 0.5 for full-siblings). Therefore, familiarity does not seem to be a reliable mechanism to assess full-sibling relatedness. Using two-choice behavioural trials, we found that juveniles from this population use both phenotype matching and familiarity to distinguish kin from nonkin. However, we did not find strong evidence that the guppies use these mechanisms to form shoals of related individuals as adults, which is similar to results from other guppy populations in Trinidad. The use of both familiarity and phenotype matching is discussed in the context of the Paria River guppy population's mating system and ecology. Overall, these data provide support for the kin recognition hypothesis and increase our understanding of the evolution of kin recognition systems.

Multiple mating and sequential mate choice in guppies: females trade up.

The trade-up hypothesis outlines a behavioural strategy that females could use to maximize the genetic benefits to their offspring. The hypothesis proposes that females should be more willing to accept a mate when the new male encountered is a superior genetic source to previous mates. We provide a direct test of the trade-up hypothesis using guppies (Poecilia reticulata), and evaluate both behavioural and paternity data. Virgin female guppies were presented sequentially with two males of varying attractiveness, and their responsiveness to each male was quantified. Male attractiveness (ornamentation) was scored as the amount of orange coloration on their body. Females were generally less responsive to second-encountered males, yet responsiveness to second males was an increasing function of male ornamentation. These attractive second males also sired a greater proportion of the offspring. There was an overall tendency for last-male advantage in paternity, and this advantage was most exaggerated when the second male was more ornamented than the first. Finally, we found that our estimate of relative sperm number did not account for any significant variation in paternity. Our results suggest that female guppies may use pre-copulatory mechanisms to maximize the genetic quality of their offspring.

Multiple mating and sequential mate choice in guppies: females trade up

The trade-up hypothesis outlines a behavioural strategy that females could use to maximize the genetic benefits to their offspring. The hypothesis proposes that females should be more willing to accept a mate when the new male encountered is a superior genetic source to previous mates. We provide a direct test of the trade-up hypothesis using guppies (Poecilia reticulata), and evaluate both behavioural and paternity data. Virgin female guppies were presented sequentially with two males of varying attractiveness, and their responsiveness to each male was quantified. Male attractiveness (ornamentation) was scored as the amount of orange coloration on their body. Females were generally less responsive to second-encountered males, yet responsiveness to second males was an increasing function of male ornamentation. These attractive second males also sired a greater proportion of the offspring. There was an overall tendency for last-male advantage in paternity, and this advantage was most exaggerated when the second male was more ornamented than the first. Finally, we found that our estimate of relative sperm number did not account for any significant variation in paternity. Our results suggest that female guppies may use pre-copulatory mechanisms to maximize the genetic quality of their offspring.