Inhibition of optimal behavior by social transmission in the guppy depends on shoaling

Previous research has suggested social learning of foragingbehavior can inhibit learning of the optimal behavior pattern.Based on their transmission chain design, we used small groupsof guppies (Poecilia reticulata) to determine the degree towhich the optimal behavior pattern was inhibited by sociallylearned information. A founder group was trained to take a long, energetically costly route to a food source. The members ofthis group were gradually replaced with naive conspecifics.Replicating the findings of the earlier researchers, it wasclear that the behavior of the founders strongly influencedthe behavior of the naive fish, probably through a process oflocal enhancement. When tested as a group, the naive fish chosethe long route to the food source significantly more oftenthan chance. Each naive fish was also tested in isolation.When tested alone, there was a significant tendency to choosethe short route despite following the long route when testedas a group. These results suggest social learning does notinhibit learning of optimal behavior patterns but that a trade-offoccurs when tested in the group condition. It is possible thatthe advantages for an individual fish of swimming with theshoal, and thus following the socially learned route, may haveoutweighed the potential energetic costs of taking this longerroute.     

Ultraviolet vision and mate choice in the guppy (Poecilia reticulata)

Ultraviolet (UV) vision is well documented for many speciesof vertebrates. UV cues are known to be used in foraging, navigationand in mate choice. We conducted a series of behavioral experimentsto investigate the role of UV perception in mate choice inboth female and male guppies (Poecilia reticulata). In ourexperiments the visual appearance of potential mates was manipulatedusing either UV transmitting (UV+) or UV blocking (UV-) filters.Female guppies significantly preferred UV+ males. Male guppiestended to prefer UV- females, but their preferences were marginallynonsignificant. Further experiments investigating the roleof luminance, indicate that UV wavelengths are probably beingused for color discrimination rather than for detecting differencesin brightness. These experiments raise the possibility thatUV is used in mate assessment in different ways by male andfemale guppies. This may reflect the different strategies thatthe two sexes have in order to maximize reproductive success.To our knowledge, these are the first data showing that UVis used by any fish species in mate selection.     

Isolation and the development of shoaling in two populations of the guppy

Socially reared guppies Poecilia reticulata derived from two wild populations (Upper and Lower Aripo River, Trinidad) showed a significant relationship between body size and shoaling tendency, measured as the proportion of time spent in proximity to a bottle containing conspecifics. Larger females shoaled significantly more than smaller females. Fish from the high-shoaling population (Lower Aripo) showed significantly less shoaling behaviour when reared in isolation. In contrast, low-shoaling fish (Upper Aripo) demonstrated no significant change in their shoaling behaviour in response to social isolation.     

Male mating strategies under predation risk: do females call the shots?

Many authors have reported that, under elevated risk of predation,male guppies (Poecilia reticulata) alter their behavior fromcourtship to forced copulation (gonopodial thrusts not precededby sigmoid displays). This shift is presumed to benefit thebrightly colored male, whose intense courting activity mightotherwise increase his risk of detection and attack by predators.However, there is some evidence that females engaged in reproductiveactivity with males may be even more vulnerable to predatorsthan the males themselves, which suggests an alternative hypothesis:females in high-risk situations are less receptive to male courtship,and this leads males to change their behavior. We tested thishypothesis by providing either males and females separately,or both sexes concurrently, with information about elevatedpredation risk from a cichlid (Crenicichla sp.). We found thatwhen only females were provided with information about increased risk,males performed fewer courtship displays and fewer thrusts.They did not perform more forced copulations in any treatmentgroup. Nonetheless, our results suggest that the female's perceptionof predation risk can be at least as important as the male'sin changing male mating behavior.     

Sex differences in the trade-off between feeding and mating in the guppy

Male guppies Poecilia reticulata court almost continuously and often indulge in sneaky mating attempts. Female guppies, on the other hand, devote much of their time to foraging. This paper demonstrates that both sexes reduce their feeding rate in mixed-sex groups relative to single sex ones and suggests that the decrease is less pronounced for females than for males. Thus males make feeding and mating decisions that are dependent on hunger while female foraging behaviour is constrained by sexual harassment. The observed sex differences in behaviour are a consequence of asymmetrical mating costs in males and females.     

Characterization of tetranucleotide microsatellite markers in guppy (Poecilia reticulata)

Ten tetranucleotide microsatellite loci are characterized for guppy, Poecilia reticulata, an important model species in the study of adaptation and mating systems. Loci were isolated following a microsatellite enrichment procedure using probe‐labelled magnetic beads. These microsatellites were designed for use in examining gene flow, reproductive isolation, and parentage within natural guppy populations.     

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.     

Social foraging: individual learning and cultural transmission of innovations

We present two stochastic models of individual and social learningthat count the number of individuals exhibiting a learned, resource-producingtrait in a group of social foragers. The novelty of our modelingresults from incorporating the empirically based assumptionthat rates of both individual and social learning should dependon the frequency of the learned trait within the group. Whenresources occur as clumps shared by group members, a naive individual'sacquisition of the skill required for clump discovery/productionshould involve opposing processes of frequency dependence. Theopportunity to learn via cultural transmission should increasewith the trait's frequency, but the opportunity for learningindividually should decrease as the trait's frequency increases.The results of the model suggest that the evolution of the capacityfor cultural transmission may be promoted in environments wherescrounging at resource clumps inhibits rates of individual learning.     

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.     

Foraging innovation is inversely related to competitive ability in male but not in female guppies

Foraging success is likely to affect hunger level and motivationto locate and exploit novel food sources in animals. We exploredthe relationship between scramble competition for limited foodand foraging innovation in the guppy (Poecilia reticulata),predicting that poor competitors would be more likely to innovatewhen presented with novel foraging tasks. Among males, we foundthat latency to complete novel foraging tasks was correlated bothwith weight gain and number of food items consumed, suggestingthat poor competitors are more likely to innovate. However,among females there was no relationship between innovative tendencyand either weight gain or foraging success. We suggest thatthis sex difference may reflect parental investment asymmetriesin males and females, and we predict similar sex differencesin other species.     

Social transmission of nectar-robbing behaviour in bumble-bees

Social transmission of acquired foraging techniques is rarely considered outside of a vertebrate context. Here, however, we show that nectar robbing by bumble-bees (Bombus terrestris)-an invertebrate behaviour of considerable ecological significance-has the potential to spread through a population at the accelerated rates typical of social transmission. Nectar robbing occurs when individuals either bite through the base of a flower to 'steal' nectar (primary robbing) or use robbing holes that others have made (secondary robbing). We found that experience of foraging from robbed flowers significantly promoted the development of primary robbing in previously legitimate foragers, thus implying that the acquisition of nectar robbing by one individual will facilitate its adoption in others. Our findings suggest that the positive feedback effects of social transmission may potentially play an ecologically important role in the relationship between plants and pollinators.     

Genes, copying, and female mate choice: shifting thresholds

Recent experimental work on guppies (Poecilia reticulata) hasexamined the strength of genetic and cultural (copying) factorsin determining female mate choice. Using females from a populationwith a heritable preference for the amount of orange body colorpossessed by males, prior work discovered that a threshold differencein orange color among males existed below which females wouldchoose a less orange male if they observed another female choosethat male, but above which they consistently preferred the moreorange of the males, regardless of whether they viewed anotherfemale prefer the less orange male. I tested whether this thresholdcan be shifted by increasing the amount of mate-copying informationavailable to a female. I demonstrate that when a female hasthe opportunity to see two different model females independentlyprefer the less orange of two males or a single female neara drab male for a longer period of time (twice as long as inprior work), the observer female prefers this drab male evenwhen males dramatically differ in orange coloration.     

Heterosis for neonatal survival in the guppy

Neonatal survival rate ranged from 91.8 to 100.0% in 12 populations of the guppy Poecilia reticulata , and was higher in naturalized Japanese stream populations than in domestic strains. Mean heterozygosity at five allozyme and four microsatellite loci varied between 0.112 and 0.430 and was significantly correlated with neonatal survival rate among populations, suggest ing that inbreeding decreases neonatal survival. Diallel and reciprocal crosses among four domestic strains demonstrated heterosis for neonatal survival. These results indicate that neonatal survival is genetically affected by heterosis and its antithesis, inbreeding depression. The relationship between neonatal survival and the mean heterozygosity suggests that overall heterozygosity is important for neonatal survival of the guppy.     

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.     

Primers for 12 polymorphic microsatellite DNA loci from the guppy (Poecilia reticulata)

Twelve polymorphic microsatellite DNA loci were isolated from genomic guppy DNA. We assessed the level of genetic diversity for these loci using individuals from five native Trinidadian populations. All the loci were polymorphic although there were considerable differences among loci and populations in allele frequencies. Within populations, the number of alleles ranged from 1 to 22 and observed heterozygosities ranged from 0.000 to 0.990. Allele frequencies differed substantially between populations suggesting divergence.     

The complex ecology of genitalia: Gonopodium length and allometry in the Trinidadian guppy

Male genitalia present an extraordinary pattern of rapid divergence in animals with internal fertilization, which is usually attributed to sexual selection. However, the effect of ecological factors on genitalia divergence could also be important, especially so in animals with nonretractable genitalia because of their stronger interaction with the surrounding environment in comparison with animals with retractable genitalia. Here, we examine the potential of a pervasive ecological factor (predation) to influence the length and allometry of the male genitalia in guppies. We sampled guppies from pairs of low‐predation (LP) and high‐predation (HP) populations in seven rivers in Trinidad, and measured their body and gonopodium length. A key finding was that HP adult males do not have consistently longer gonopodia than do LP adult males, as had been described in previous work. However, we did find such divergence for juvenile males: HP juveniles have longer gonopodia than do LP juveniles. We therefore suggest that an evolutionary trend toward the development of longer gonopodia in HP males (as seen in the juveniles) is erased after maturity owing to the higher mortality of mature males with longer gonopodia. Beyond these generalities, gonopodium length and gonopodium allometry were remarkably variable among populations even within a predation regime, thus indicating strong context dependence to their development/evolution. Our findings highlight the complex dynamics of genitalia evolution in Trinidadian guppies.     

Cross-generational effects of sexual harassment on female fitness in the guppy

Sexual harassment is a common outcome of sexual conflict over mating rate. A large number of studies have identified several direct costs to females of sexual harassment including energy expenditure and reduced foraging ability. However, the fitness consequences of sexual harassment for descendants have rarely been investigated. Here, we manipulated the level of sexual harassment and mating rate in two groups of female guppies, Poecilia reticulata, a live-bearing fish in which sexual conflict over mating rate is particularly pronounced. Each female was allowed to interact with three males for one day (low sexual harassment, LSH) or for eight days (high sexual harassment, HSH) during each breeding cycle throughout their life. Female lifetime fecundity did not differ between the groups, but we found a strong effect on offspring fitness. HSH females produced (1) daughters with smaller bodies and (2) sons with shorter gonopodia, which were less attractive to females and less successful in coercive matings than their LSH counterparts. Although these results may be influenced by the indirect effects of sex ratio differences between treatments, they suggest that sexual harassment and elevated mating rate can have negative cross-generational fitness effects and more profound evolutionary consequences than currently thought.     

Insights from social transmission of information in rodents

Direct exposure to stimuli in their environment is not the only way that animals learn about important information. Individuals can infer fear from a social context through observation. Like humans, rats are very social animals, and may learn to infer information about their environment through their interactions with conspecifics. Here, we first review different models for social transmission of information in rodents. Second, we examine different modes of communication that are important to social learning. Then, we cover the different proximate factors that are thought to modulate the social transmission of information. Next, we identify social and environmental conditions that impact social learning, and finally, we conclude by revisiting social transmission through the lens of the Tinbergen framework.     

Social enhancement can create adaptive,arbitrary and maladaptive cultural traditions

Many animals are known to learn socially, i.e. they are able to acquire new behaviours by using information from other individuals. Researchers distinguish between a number of different social-learning mechanisms such as imitation and social enhancement. Social enhancement is a simple form of social learning that is among the most widespread in animals. However, unlike imitation, it is debated whether social enhancement can create cultural traditions. Based on a recent study on capuchin monkeys, we developed an agent-based model to test the hypotheses that (i) social enhancement can create and maintain stable traditions and (ii) social enhancement can create cultural conformity. Our results supported both hypotheses. A key factor that led to the creation of cultural conformity and traditions was the repeated interaction of individual reinforcement and social enhancement learning. This result emphasizes that the emergence of cultural conformity does not necessarily require cognitively complex mechanisms such as ‘copying the majority’ or group norms. In addition, we observed that social enhancement can create learning dynamics similar to a ‘copy when uncertain’ learning strategy. Results from additional analyses also point to situations that should favour the evolution of learning mechanisms more sophisticated than social enhancement.