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THE TRINIDADIAN GUPPY Fig. 1 Female guppies hiding among plants. Guppies (Poecilia reticulata) are small, freshwater fish, native to the island nation of Trinidad and Tobago and to the adjacent South American mainland. Although guppies have been introduced to tropical regions and pet stores worldwide, they exist in a ‘natural laboratory’ within the streams of Trinidad. Guppies have become a model system in Behavior, Ecology, and Evolution due to their wealth of easily observable behaviors and their ability to rapidly evolve in response to changing environmental pressures. In guppies, the females are larger and drab in color, while the males are smaller and brightly colored. Guppies are unusual among fish because they are livebearers. This means that they give birth to live young, rather than laying eggs like most fish. The time from birth to sexual maturity in guppies is about 3 months, and females can give birth to 3-30 babies. In fish, a group of sibling babies is referred to as a brood. Many of the differences in behavior and morphology seen in distinct populations of guppies are associated with differences in predation pressure. In downstream “highpredation” environments guppies co-occur with many predators. These predators are prevented from moving upstream by waterfall barriers, resulting in “low-predation” populations at higher elevations. Highpredation guppies have repeatedly colonized and adapted to upstream low-predation environments resulting in parallel changes in life-history traits, morphology, and behavior (see Table 1 at the end of this packet). Thus, this evolutionary scenario has been naturally replicated many times and is unique because it allows us to better understand why certain characteristics are favorable in different environments and how these characteristics evolve. In essence, we have access to a natural experiment that has been replicated many times. To understand how and why differences among populations evolve, we have to think in terms of natural selection (also called survival of the fittest), an idea first proposed by Charles Darwin (1809-1882). The central idea behind this theory is that animals in the wild must overcome the challenges to survive and reproduce in changing environments. Only those individuals that best match their environment survive and pass on their genes to the next generation. As this process repeats itself, advantageous characteristics become more common and undesirable characteristics become less common. Over time this persistence and spread of advantageous traits allows populations to evolve and become adapted (well-matched) to their environments. The theory of natural selection is a central principle in evolution and biology in general. Fig. 2 A male (top right) and female (bottom left) guppy. Pet traders have taken advantage of guppies’ ability to rapidly evolve by selecting for very elaborate colors and tail shapes. Selective breeding or artificial selection is similar to natural selection, except that it is the breeders who decide which traits are desirable, rather than the natural Guppy Behavior Fig. 3 Typical high-predation (top) and lowpredation (bottom) sites. environment. Selective breeding has factored heavily into many aspects of human society (domestic animals, agricultural products, etc.). The fancy guppies we find in pet stores originated from wild guppies, but many years of selective breeding have changed both their appearance and their behavior. Studies have demonstrated that fancy guppies have lost many of the innate fear responses seen in wild guppies, making them beautiful but unlikely to survive in the wild. Fig. 4 The pike cichlid, Crenicichla frenata, a common and voracious guppy predator. Anti-predator behavior Shoaling behavior is a grouping behavior used as an anti-predator tactic by many fish species. It is less coordinated than schooling behavior, but serves the same purpose of creating safety in numbers. Highpredation guppies shoal more than lowpredation guppies, and guppies from both populations increase shoaling behavior if they detect a predator in their immediate environment. Nearly all fish display a special escape response known as a ‘fast-start’ escape response. Guppies can perform this behavior so quickly that it is only visible when recording with a high-speed camera. When watching with the naked eye we see the guppy move from one position to another without any apparent movement in between. Other predator evasion behaviors displayed by guppies include freezing and/or hiding to make themselves less visible to predators. Courtship behavior The mating display of the male guppy is the ‘sigmoid display’, in which a male will position himself perpendicularly in front of a female, assume the S-shaped posture that gives the display its name, and quiver his body. This display is basically a dance to show off and get the female’s attention. The problem is that sigmoid displays also make males more visible to predators. Similarly, females tend to prefer brighter males, but bright coloration also makes the males more conspicuous to predators. In addition, studies have found that females prefer rare males, males that are daring (i.e. that they have observed in the presence of a predator), and males that are preferred by other females (i.e. that they have observed mating with other females). Together, these factors create a trade-off between getting mates and avoiding predators, something that makes sexual selection important in guppies. Sexual selection is a special case of natural selection that is focused on differences between individuals in their ability to acquire mates and produce offspring. Sexual selection is an important extension of natural selection because it helps explain the presence of traits that seem maladaptive – such as long tail feathers in birds and bright colors in guppies. In other words, traits that are bad for predator avoidance can still become common if they are good for mating success. In fact, some of these traits are also thought to be indicators of good genes. For example, bright colors in guppies are an indicator of good health (males become dull when they are sick), and a bright male guppy that survives to reproduce despite being more visible may be particularly good at predator avoidance. Foraging behavior As with most animals, finding food takes up much of guppies’ lives. As with mating behavior, there is a trade-off between foraging and avoiding predators. Guppies looking for food might be distracted and not as aware of predators. Being in a group can help, since there are more eyes to look for predators, but being in a group also increases competition for food and mates. Because guppies have to balance these different factors, they will alter their behaviors depending on how hungry they are. Hungrier guppies take more risks than guppies that have recently eaten. Fig. 5 A male guppy tries to get the attention of a female. Fig. 6 Our favorite guppy predator. He is a pike cichlid named Pickles who lives in our lab. High-predation Low-predation body size smaller bigger body shape streamlined robust # of babies many few size of babies smaller larger time to maturity short long # sigmoids few many # sneaky attempts many few amount of color less more shoaling often and close not often, less close time foraging less more