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Diverse clades of fishes adapted to feeding on the benthos repeatedly converge on steep craniofacial profiles and shorter, wider heads. But in an incipient radiation, to what extent is this morphological evolution measurable and can we distinguish the relative genetic vs. plastic effects? We use the Trinidadian guppy (Poecilia reticulata) to test the repeatability of adaptation and the alignment of genetic and environmental effects shaping poecilid craniofacial morphology. We compare wild-caught and common garden lab-reared fish to quantify the genetic and plastic components of craniofacial morphology across 4 populations from 2 river drainage systems (n  = 56 total). We first use micro-computed tomography to capture 3D morphology, then place both landmarks and semilandmarks to perform size-corrected 3D morphometrics and quantify shape space. We find a measurable, significant, and repeatable divergence in craniofacial shape between high-predation invertivore and low-predation detritivore populations. As predicted from previous examples of piscine adaptive trophic divergence, we find increases in head slope and craniofacial compression among the benthic detritivore foragers. Furthermore, the effects of environmental plasticity among benthic detritivores produce exaggerated craniofacial morphological change along a parallel axis to genetic morphological adaptation from invertivore ancestors. Overall, many of the major patterns of benthic-limnetic craniofacial evolution appear convergent among disparate groups of teleost fishes. [ABSTRACT FROM AUTHOR]

Coloration facilitates evolutionary investigations in nature because the interaction between genotype, phenotype and environment is relatively accessible. In a landmark set of studies, Endler addressed this complexity by demonstrating that the evolution of male Trinidadian guppy coloration is shaped by the local balance between selection for mate attractiveness versus crypsis. This became a textbook paradigm for how antagonistic selective pressures may determine evolutionary trajectories in nature. However, recent studies have challenged the generality of this paradigm. Here, we respond to these challenges by reviewing five important yet underappreciated factors that contribute to colour pattern evolution: (i) among-population variation in female preference and correlated variation in male coloration, (ii) differences in how predators versus conspecifics view males, (iii) biased assessment of pigmentary versus structural coloration, (iv) the importance of accounting for multi-species predator communities, and (v) the importance of considering the multivariate genetic architecture and multivariate context of selection and how sexual selection encourages polymorphic divergence. We elaborate these issues using two challenging papers. Our purpose is not to criticize but to point out the potential pitfalls in colour research and to emphasize the depth of consideration necessary for testing evolutionary hypotheses using complex multi-trait phenotypes such as guppy colour patterns. [ABSTRACT FROM AUTHOR]

Life‐history phenotypes emerge from clusters of traits that are the product of genes and phenotypic plasticity. If the impact of the environment differs substantially between traits, then life histories might not evolve as a cohesive whole. We quantified the sensitivity of components of the life history to food availability, a key environmental difference in the habitat occupied by contrasting ecotypes, for 36 traits in fast‐ and slow‐reproducing Trinidadian guppies. Our dataset included six putatively independent origins of the slow‐reproducing, derived ecotype. Traits varied substantially in plastic and genetic control. Twelve traits were influenced only by food availability (body lengths, body weights), five only by genetic differentiation (interbirth intervals, offspring sizes), 10 by both (litter sizes, reproductive timing), and nine by neither (fat contents, reproductive allotment). Ecotype‐by‐food interactions were negligible. The response to low food was aligned with the genetic difference between high‐ and low‐food environments, suggesting that plasticity was adaptive. The heterogeneity among traits in environmental sensitivity and genetic differentiation reveals that the components of the life history may not evolve in concert. Ecotypes may instead represent mosaics of trait groups that differ in their rate of evolution. [ABSTRACT FROM AUTHOR]

Dynamic energy budget (DEB) theory provides a model of the intrinsic energetic trade‐offs that shape life histories, in terms of fluxes of energy through biological processes. In these models, life histories are a function of environmental conditions and of fundamental traits of the organism relating to the acquisition, allocation and use of energy.These traits are described by the parameters of the DEB model, which are typically estimated from multiple data sources. DEB parameters have been estimated for over 2,500 species, and recent work has aimed to compare species on the basis of differences in parameter values. However, little is known about intraspecific variation in DEB parameters, and the reliability of comparisons between parameter sets is rarely tested.We estimated DEB parameters for 16 populations of Trinidadian guppy. Our objectives were to determine whether DEB theory could characterise known life‐history differences between populations, and to assess whether differences in parameter values between populations were reliable indicators of biological differences.We found substantial intraspecific variation in parameter values. The models accurately reproduced biological processes, and also predicted differences in resource allocation and metabolic rate consistent with known life‐history differences among these populations.However, parameter estimates varied substantially as an artefact of the amount and types of data used to fit the models. This bias appeared to be systematic, influencing estimates for multiple populations in a comparable fashion. We show that similar patterns of growth and reproduction can be reproduced with very different parameter sets. Our results imply that comparisons of DEB parameters between models fit with different types of data may reflect artefactual rather than biological differences. A free Plain Language Summary can be found within the Supporting Information of this article. [ABSTRACT FROM AUTHOR]

Much of our understanding of natural invasions is retrospective, based on data acquired after invaders become established. As a consequence, we know little about the characteristics of the early population growth and habitat use of the invaders during establishment. Here we report on experimental introductions of guppies into natural streams in which we conducted monthly censuses of each population. Two of the four introductions were in streams with thinned canopies, which mimic a common form of habitat disturbance. We conducted similar censuses of natural populations to characterize natural population densities and generate a null distribution against which we could test a priori hypotheses about the establishment of the experimental invaders. We constructed a pedigree for one population, which enabled us to quantify lifetime reproductive success. Population simulations predict that the nature of the introduced population's life history, in combination with reduced risk of predation in the introduction sites, will result in explosive population growth; however, populations of introduced invaders instead grew to match densities observed in natural streams with intact canopies. Experimental populations in streams with thinned canopies grew to densities that often exceeded those of natural streams with intact canopies. High population densities were associated with the increased use of marginal habitat. Adult females and males that moved into marginal habitat suffered no apparent fitness loss, suggesting lower population densities found there compensated for lower habitat quality. Our results suggest that the ecological setting in which invasions occur plays a role at least comparable in importance to that of the invader's inherent characteristics in shaping early population growth and habitat use. [ABSTRACT FROM AUTHOR]

Guppies in Trinidad range across aquatic environments with fish communities that vary in risk of predation. These communities are often discrete, separated by waterfalls, with high-predation communities downstream and low-predation communities upstream. This gradient is repeated in many rivers; in each one, we see the same divergence between guppy populations in life history, behavior, morphology, and physiology. We have shown that the agent of selection on the life history, behavior, and physiology in low-predation communities is high population density and the cascade of ecological effects that stems from it. In effect, guppy populations modify their ecosystem and, in so doing, impose selection on themselves and shape their own evolution, which further changes the ecosystem. Evolution unfolds rapidly in this system, which has enabled us to study the dynamics of the process, not just its end points. Those studies enable us to answer some very general questions in ecology and evolutionary biology. [ABSTRACT FROM AUTHOR]

Organisms can change their environment and in doing so change the selection they experience and how they evolve. Population density is one potential mediator of such interactions because high population densities can impact the ecosystem and reduce resource availability. At present, such interactions are best known from theory and laboratory experiments. Here we quantify the importance of such interactions in nature by transplanting guppies from a stream where they co-occur with predators into tributaries that previously lacked both guppies and predators. If guppies evolve solely because of the immediate reduction in mortality rate, the strength of selection and rate of evolution should be greatest at the outset and then decline as the population adapts to its new environment. If indirect effects caused by the increase in guppy population density in the absence of predation prevail, then there should be a lag in guppy evolution because time is required for them to modify their environment. The duration of this lag is predicted to be associated with the environmental modification caused by guppies. We observed a lag in life-history evolution associated with increases in population density and altered ecology. How guppies evolved matched predictions derived from evolutionary theory that incorporates such density effects. [ABSTRACT FROM AUTHOR]

Experimental studies of evolution performed in nature and the associated demonstration of rapid evolution, observable on a time scale of months to years, were an acclaimed novelty in the 1980–1990s. Contemporary evolution is now considered ordinary and is an integrated feature of many areas of research. This shift from extraordinary to ordinary reflects a change in the perception of evolution. It was formerly thought of as a historical process, perceived through the footprints left in the fossil record or living organisms. It is now seen as a contemporary process that acts in real time. Here we review how this shift occurred and its consequences for fields as diverse as wildlife management, conservation biology, and ecosystems ecology. Incorporating contemporary evolution in these fields has caused old questions to be recast, changed the answers, caused new and previously inconceivable questions to be addressed, and inspired the development of new subdisciplines. We argue further that the potential of contemporary evolution has yet to be fulfilled. Incorporating evolutionary dynamics in any research program can provide a better assessment of how and why organisms and communities came to be as they are than is attainable without an explicit treatment of these dynamics. [ABSTRACT FROM AUTHOR]

We describe the occurence of extensive post-fertilization maternal provisioning (matrotrophy) in fish species in the subgenus Pamphorichthys (genus Poecilia : family Poeciliidae) that represents one of two independent origins of this adaptation in this genus. Matrotrophy is accompanied by a reduction in yolk in eggs at fertilization, a thickened follicle throughout development, and externalization and anterior extension of the embryonic pericardial membrane. These features resemble the anatomical adaptations for placentotrophy described in other members of this family and accompany a substantial increase in dry mass of the embryo during development. Species mean values for the increase in embryo mass range from a low of less than two-fold (Pamphorichthys minor) to greater than 50-fold (Pamphorichthys hasemani). Different populations of Pamphorichthys araguaiensis show a range from less than a two-fold to greater than 16-fold increase in dry mass during development. Such substantial differences in matrotrophy among closely related species and within species make Pamphorichthys a promising group with which to study the adaptive value and the genetic basis of matrotrophy. [ABSTRACT FROM AUTHOR]

Annual killifish that are adapted to life in aquatic habitat that dries seasonally have evolved desiccation-resistant eggs capable of undergoing diapause, i.e. developmental arrest, at specific stages during embryology. Although noted for their remarkable ability to live at the land–water interface, species in the genus Rivulus are considered to be non-annual killifish that exhibit typical teleost developmental patterns and no embryonic diapause. Here, we combine a molecular phylogeny with an embryological study in order to demonstrate an independent origin of mid-embryonic diapause within a clade of Rivulus (subgenus Laimosemion) that inhabits small streams or savannah pools. We also observed that some of these species exhibit a short dispersed cell phase separating epiboly and the formation of the embryo axis, which is a feature of development observed only in annual killifish. Lastly, we incubated embryos of Laimosemion species and outgroup taxa in both water and peat moss and observed that on peat moss the embryos of all species are capable of delaying hatching for > 10 days, but when water incubated there are significant differences among species in the duration of this delay before hatching. We hypothesize that the preferred microhabitat of this clade of killifish exposes their embryos to periodic desiccation, creating selection in favour of embryonic diapause. [ABSTRACT FROM AUTHOR]

The fish subfamily Poeciliinae (sensu Parenti, 1981) is widely distributed across the Western Hemisphere and a dominant component of the fish communities of Central America. Poeciliids have figured prominently in previous studies on the roles of dispersal and vicariance in shaping current geographic distributions. Most recently, Hrbek et al. combined a DNA-based phylogeny of the family with geological models to provide a biogeographic perspective that emphasized the role of both vicariance and dispersal. Here we expand on that effort with a database enlarged in the quantity of sequence represented per species, in the number of species included, and in an enlarged and more balanced representation of the order Cyprinodontiformes. We combine a robust timetree based upon multiple fossil calibrations with enhanced biogeographic analyses that include ancestral area reconstructions to provide a detailed biogeographic history of this clade. Key features of our results are that the family originated in South America, but its major diversification dates to a later colonization of Central America. We also resolve additional colonizations among South, Central and North America and the Caribbean and consider how this reconstruction contributes to our understanding of the mechanisms of dispersal. [ABSTRACT FROM AUTHOR]

Speciation is a multifaceted process that involves numerous aspects of the biological sciences and occurs for multiple reasons. Ecology plays a major role, including both abiotic and biotic factors. Whether populations experience similar or divergent ecological environments, they often adapt to local conditions through divergence in biomechanical traits. We investigate the role of biomechanics in speciation using fish predator–prey interactions, a primary driver of fitness for both predators and prey.We highlight specific groups of fishes, or specific species, that have been particularly valuable for understanding these dynamic interactions and offer the best opportunities for future studies that link genetic architecture to biomechanics and reproductive isolation (RI). In addition to emphasizing the key biomechanical techniques that will be instrumental, we also propose that the movement towards linking biomechanics and speciation will include (i) establishing the genetic basis of biomechanical traits, (ii) testing whether similar and divergent selection lead to biomechanical divergence, and (iii) testing whether/how biomechanical traits affect RI. Future investigations that examine speciation through the lens of biomechanics will propel our understanding of this key process. [ABSTRACT FROM AUTHOR]

Aim: The theory of island biogeography provides a predictive framework relating species richness to island size and distance from the mainland. However, the theory as originally formulated does not necessarily scale to large islands and continental landmasses that are capable of generating species through in situ speciation (rather than entirely by colonization), nor does it necessarily account for how human introduction of species alters traditional biogeographical patterns. Here, we examine the ecological (colonization and extinction), evolutionary (in situ speciation) and human-mediated (deliberate introductions) determinants of species richness in a taxonomic group that has undergone a radiation on Caribbean islands: live-bearing fishes of the family Poeciliidae. Location: The Caribbean. Methods: We created a database of both native and introduced poeciliid species occurrence on Caribbean islands through literature review, and estimated the number of colonizations versus speciation events on each island using a molecular phylogeny. Linear regression and other statistical tests were used to explore species-area and species-isolation relationships. Results: Species richness on small islands results entirely from colonization and does not significantly increase with island area, whereas on larger islands species richness increases dramatically as a function of area due primarily to in situ speciation. Poeciliid fishes have been introduced widely, both as a byproduct of their popularity in the aquarium hobby and as a means of mosquito control. We show that such establishments have occurred disproportionately on islands depauperate in native species, and that introduced species richness is positively correlated with economic interconnectedness (shipping traffic) and human population size. Main conclusions: On large Caribbean islands in situ speciation has elevated the number of poeciliid species beyond that predicted from ecological processes alone. Introduced species significantly alter biogeographical patterns. [ABSTRACT FROM AUTHOR]

The livebearing Trinidadian guppy ( Poecilia reticulata) produces bigger offspring in populations exposed to low predation and produces smaller, more numerous offspring in populations subject to high predation (HP)., Like most fishes, guppies respond to predator attacks with a fast-start escape response. From the scaling of teleost fast-start performance, we predict that larger guppy neonates should exhibit faster, more effective escape responses than smaller neonates. Increasing performance with increasing size could be due simply to size, or to both size and the acquisition of more mature body forms, as is seen in larval-stage fishes., We find no difference in external body proportions among guppy offspring varying in size from 5·1 mm to 7·1 mm at birth, suggesting offspring are born morphologically mature. However, based on the degree of skeletal ossification, as a proxy for internal maturity, we find that guppies are still maturing rapidly around the time of birth. The smallest neonates from the very HP Caroni confluence lack ossification of key skeletal elements that are present in their larger low predation counterparts., In guppies from the Aripo/Caroni drainage, we show that neonatal escape performance covaries with responsiveness and increases with size along a gradient of HP offspring, but is lower in the largest low predation offspring. The scaling of escape performance in the HP populations performing maximally (scaling with length as L2·76) exceeds predictions from size alone (scaling exponent of L1·0 indicates performance increases linearly with body size). This suggests that guppy neonates, like rapidly developing larval fishes, vary substantially in morphological maturity among populations., The finding that neonatal guppy offspring covary in both size and maturity at birth means that being smaller also means being less mature, which amplifies the negative escape performance effects of being born small. Despite the negative consequences of being born both small and immature, HP environments select heavily for high fecundity, and thus small offspring size. We find selection favours female life-history traits over offspring escape performance. [ABSTRACT FROM AUTHOR]

There are many theoretical and empirical studies explaining variation in offspring sex ratio but relatively few that explain variation in adult sex ratio. Adult sex ratios are important because biased sex ratios can be a driver of sexual selection and will reduce effective population size, affecting population persistence and shapes how populations respond to natural selection. Previous work on guppies ( Poecilia reticulata) gives mixed results, usually showing a female-biased adult sex ratio. However, a detailed analysis showed that this bias varied dramatically throughout a year and with no consistent sex bias. We used a mark-recapture approach to examine the origin and consistency of female-biased sex ratio in four replicated introductions. We show that female-biased sex ratio arises predictably and is a consequence of higher male mortality and longer female life spans with little effect of offspring sex ratio. Inconsistencies with previous studies are likely due to sampling methods and sampling design, which should be less of an issue with mark-recapture techniques. Together with other long-term mark-recapture studies, our study suggests that bias in offspring sex ratio rarely contributes to adult sex ratio in vertebrates. Rather, sex differences in adult survival rates and longevity determine vertebrate adult sex ratio. [ABSTRACT FROM AUTHOR]

An eco-evolutionary feedback loop is defined as the reciprocal impacts of ecology on evolutionary dynamics and evolution on ecological dynamics on contemporary timescales. We experimentally tested for an eco-evolutionary feedback loop in the green peach aphid, Myzus persicac, by manipulating initial densities and evolution. We found strong evidence that initial aphid density alters the rate and direction of evolution, as measured by changes in genotype frequencies through time. We also found that evolution of aphids within only 16 days, or approximately three generations, alters the rate of population growth and predicts density compared to nonevolving controls. The impact of evolution on population dynamics also depended on density. In one evolution treatment, evolution accelerated population growth by up to 10.3% at high initial density or reduced it by up to 6.4% at low initial density. The impact of evolution on population growth was as strong as or stronger than that caused by a threefold change in intraspecific density. We found that, taken together, ecological condition, here intraspecific density, alters evolutionary dynamics, which in turn alter concurrent population growth rate (ecological dynamics) in an eco-evolutionary feedback loop. Our results suggest that ignoring evolution in studies predicting population dynamics might lead us to overor underestimate population density and that we cannot predict the evolutionary c.utcome within aphid populations without considering population size. [ABSTRACT FROM AUTHOR]

An introduction is presented in which the author discusses various reports within the issue on topics including the interaction between evolution and ecology, evolution changes ecology, and conceptual unification of the life sciences.

The biomass of living organisms hosts oniy a small portion of the elemental abundance at the surface of the Earth, yet biology plays a defining role in the composition and stability of the biosphere by acting on sensitive geochemical feedbacks controlling global element cycles. This type of influence is evident in a class of evolutionary innovations that have a profoundly disproportionate effect on the biosphere, referred to here as evolutionary innovation biospheric feedbacks (EIBFs). A particular biological innovation need not be complex, rather its influence is amplified by its effect on geochemical feedbacks controlling elemental cycling. The lead-up to the metazoan radiation (~585 million years ago) provides an example of such an EIBF. While commonly attributed to an increase in free oxygen concentration, the reason for this step increase in O2 almost 2 billion years after the advent of oxygenic photosynthesis is traced to a seemingly unrelated evolutionary innovation resulting in a critical by-product of the first soils: secondary clay minerals. Detrital clay minerals deposited in continental margin sediments sequester organic carbon compounds and thus prevent consumption ofatmospheric oxygen produced during photosynthesis. The transition from the abiotic to biotic land surface at the end of the Precambrian shifted biogeochemical cycling to this terrestrial-dominated modern mode that enabled sufficient oxygenation of the biosphere to trigger the metazoan radiation. [ABSTRACT FROM AUTHOR]

In prior research, we found the way guppy life histories evolve in response to living in environments with a high or low risk of predation is consistent with life-history theory that assumes no density dependence. We later found that guppies from high-predation environments experience higher mortality rates than those from low-predation environments, but the increased risk was evenly distributed across all age/size classes. Life-history theory that assumes density-independent population growth predicts that life histories will not evolve under such circumstances, yet we have shown with field introduction experiments that they do evolve. However, theory that incorporates density regulation predicts this pattern of mortality can result in the patterns of life-history evolution we had observed. Here we report on density manipulation experiments performed in populations of guppies from low-predation environments to ask whether natural populations normally experience density regulation and, if so, to characterize the short-term demographic changes that underlie density regulation. Our experiments reveal that these populations are density regulated. Decreased density resulted in higher juvenile growth, decreased juvenile mortality rates, and increased reproductive investment by adult females. Increased density causes reduced offspring size, decreased fat storage by adult females, and increased adult mortality. [ABSTRACT FROM AUTHOR]

Ecology Letters (2011) 14: 1084-1092 Abstract Rapid evolution challenges the assumption that evolution is too slow to impact short-term ecological dynamics. This insight motivates the study of ' Eco-Evolutionary Dynamics' or how evolution and ecological processes reciprocally interact on short time scales. We tested how rapid evolution impacts concurrent population dynamics using an aphid ( Myzus persicae) and an undomesticated host ( Hirschfeldia incana) in replicated wild populations. We manipulated evolvability by creating non-evolving (single clone) and potentially evolving (two-clone) aphid populations that contained genetic variation in intrinsic growth rate. We observed significant evolution in two-clone populations whether or not they were exposed to predators and competitors. Evolving populations grew up to 42% faster and attained up to 67% higher density, compared with non-evolving control populations but only in treatments exposed to competitors and predators. Increased density also correlates with relative fitness of competing clones suggesting a full eco-evolutionary dynamic cycle defined as reciprocal interactions between evolution and density. [ABSTRACT FROM AUTHOR]

Summary 1. The ability to adjust the allocation of energy to maintenance, growth and reproduction in response to fluctuations in resource availability, in a way that enhances fitness, is thought to depend on the mode of maternal provisioning. 2. We manipulated food availability in the matrotrophic, livebearing fish Phalloptychus januarius (Poeciliidae) to examine patterns of allocation under fluctuating resource conditions. 3. We observed an asynchrony in the adjustment of offspring traits in response to changes in food availability. A reduction in food availability caused an immediate reduction in allocation of energy to offspring size and lipid content at birth, but a delayed reduction in offspring number (fecundity). Similarly, an increase in food availability caused an immediate increase in offspring size and lipid content and a delayed increase in fecundity. This asynchrony is thought to be inherent to matrotrophy, limiting a female's ability to attain an optimal fitness in fluctuating resource environments, regardless of whether food availability changes from high to low, or low to high. 4. We found no evidence for embryo abortion under low food conditions. All developing offspring were retained, yet were smaller at birth. Furthermore, although females carried large fat reserves, these were rapidly depleted during low food conditions and were not sufficient to fully buffer gestating females or their developing offspring against the detrimental effects of reduced food availability. 5. Our study shows that matrotrophy is likely to be a maladaptive strategy in environments that are characterized by fluctuations in resource availability. It further suggests that matrotrophy is most likely to evolve in high and stable resource environments. [ABSTRACT FROM AUTHOR]

Life-history theory predicts that increased predation on juvenile age/size-classes favors delayed maturation and decreased reproductive investment. Although this theory has received correlative support, experimental tests in nature are rare. In 1976 and 1981, guppies ( Poecilia reticulata) were transplanted into localities that previously only contained a killifish, Rivulus hartii. This situation presents an opportunity to experimentally test this life-history prediction because guppies prey upon young Rivulus. We evaluated the response to selection in Rivulus by measuring phenotypic and genotypic divergence between introduction and upstream 'control' localities that lack guppies. Contrary to expectations, Rivulus from the introduction sites evolved earlier maturation and increased reproductive investment within 25 years. Such evolutionary changes parallel previous investigations on natural communities of Rivulus, but do not comply with predictions of age/size-specific theory. Guppies also caused reduced densities and increased growth rates of Rivulus, which are hypothesized indirect effects of predation. Additional life-history theories show that changes in density and growth can interact with predator-induced mortality to alter the predicted trajectory of evolution. We discuss how these latter frameworks improve the fit between theory and evolution in Rivulus. [ABSTRACT FROM AUTHOR]

Early theories of life-history evolution predict that increased predation on young/small individuals selects for delayed maturation and decreased reproductive effort, but such theory only considers changes in mortality. Predators reduce prey abundance and increase food to survivors. Theory that incorporates such indirect effects yields different predictions. Trinidadian killifish, Rivulus hartii, inhabit communities with and without guppies. Guppies prey on young Rivulus and Rivulus densities decline and growth rates increase when guppies are present. Prior work showed that Rivulus phenotypes from communities with guppies matured earlier and had higher fecundity, consistent with theories that incorporate indirect effects. Here we examined the genetic basis of these differences by rearing 2nd generation, laboratory-born Rivulus from sites with and without guppies under two food levels that match natural differences in growth. Many locality × food interactions were significant, often reversing the relationship between communities. Such interactions imply that there are fitness trade-offs associated with adaptation to high or low resource environments. On high food, Rivulus from localities with guppies matured earlier, produced many small eggs, and exhibited increased reproductive investment; these differences reversed on low food. Our results suggest that indirect effects mold Rivulus evolution and thereby highlight connections between community processes and evolutionary change. [ABSTRACT FROM AUTHOR]

Changes in age/size-specific mortality, due to such factors as predation, have potent evolutionary consequences. However, interactions with predators commonly impact prey growth rates and food availability and such indirect effects may also influence evolutionary change. We evaluated life-history differences in Trinidadian killifish, Rivulus hartii, across a gradient in predation. Rivulus are located in (1) “high predation” sites with large piscivores, (2) “ Rivulus/guppy” sites with guppies, and (3) “ Rivulus-only” sites with just Rivulus. Rivulus suffer higher mortality with large predators, and guppies may prey upon small/young Rivulus in Rivulus/guppy environments. In turn, population densities decline while growth rates increase in both localities compared to Rivulus-only sites. To explore how the direct and indirect effects of predators and guppies influence trait diversification in Rivulus, we examined life-history phenotypes across five rivers. High predation phenotypes exhibited a smaller size at reproduction, a greater number of eggs that were smaller, and increased reproductive allotment. Such changes are consistent with a direct response to predation. Rivulus from Rivulus/guppy sites were intermediate; they exhibited a smaller size at reproduction, increased fecundity, smaller eggs, and larger reproductive allotment than Rivulus-only fish. These changes are consistent with models that incorporate the impacts of growth and resources. [ABSTRACT FROM AUTHOR]

Numerous studies of wild populations have shown that phenotypic traits can change adaptively on short timescales, but very few studies have considered coincident changes in major fitness components. We here examine adaptive changes in life-history traits and survival rates for wild guppies introduced into new environments. Female life-history traits in the derived (Damier River) populations diverged from the ancestral (Yarra River) population, as a result of adaptation to predation regime (high vs. low) and other aspects of the local river. Moreover, some components of the derived Damier populations, particularly juveniles, now show higher survival in the Damier than do contemporary representatives from the ancestral Yarra population. These results suggest that adaptive change can improve survival rates after fewer than 10 years (fewer than 30 guppy generations) in a new environment. [ABSTRACT FROM AUTHOR]

Evolutionary biologists have long sought to understand the relationship between microevolution (adaptation), which can be observed both in nature and in the laboratory, and macroevolution (speciation and the origin of the divisions of the taxonomic hierarchy above the species level, and the development of complex organs), which cannot be witnessed because it occurs over intervals that far exceed the human lifespan. The connection between these processes is also a major source of conflict between science and religious belief. Biologists often forget that Charles Darwin offered a way of resolving this issue, and his proposal is ripe for re-evaluation in the light of recent research. [ABSTRACT FROM AUTHOR]

The evolution of exaggerated sexual ornamentation is classically thought to proceed as a compromise between opposing vectors of sexual and natural selection. In colour-based ornamentation, as exhibited by guppies ( Poecilia reticulata), heightened trait expression may be beneficial in promoting attractiveness, but costly in terms of predation. Opportunities to reconcile this compromise will exist if there are differences between conspecifics and predators in their sensory systems; in such situations guppies should evolve to exploit the ways in which their ornamentation would appear maximally conspicuous to conspecifics. In the present study, we addressed this hypothesis via a study of geographic variation employing the most sophisticated colour analysis yet attempted for Trinidadian guppies. We made two paired contrasts, one between two Aripo populations that vary in the presence of the potential predator Aequidens pulcher, and another between Quare and Marianne populations that vary in exposure to a predatory prawn, Macrobrachium crenulatum. We predicted that, if ornamentation is constrained by the presence of either predator, then guppy conspicuousness should change most markedly across each of the two paired populations as viewed by that predator. Although disparity analysis of entire colour patterns indicated significant differences in both contrasts, this prediction was most clearly supported for the Marianne/Quare contrast. Marianne fish, which co-exist with prawns, exhibited larger black spots coupled with less extensive, less bright flank iridescence. The brightness reductions are notable because, as the only potential guppy predator with a dedicated ultraviolet (UV) photoreceptor, prawns may detect passing male guppies via their UV-bright blues, violets and ‘UV/oranges’. We discuss our findings in light of the additional insights that might be obtained by combining spectral assessments and visual modeling with more traditional methods of colour pattern appraisal. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95, 734–747. [ABSTRACT FROM AUTHOR]

The ecological impacts of the indirect effects of predators are well established, but the evolutionary consequences are unknown. Predators often decrease prey density, which indirectly increases the resources available to surviving prey. This ecological effect could provide a link to evolution because it is generally assumed that resource availability influences life history evolution. Yet, predictions from theory that consider food availability are inconsistent, and evidence for an important role of resources in shaping life history evolution is absent. We compared life history traits in a Trinidadian killifish, Rivulus barth, from fish communities that differ in predation intensity; predators are associated with lower population density and faster growth rates. To determine whether the indirect effects of predators influence evolutionary change, we reared second-generation-born fish under two food levels that approximated natural differences in resources between communities. Rivulus from sites with predators are younger and smaller at maturity. They have increased reproductive investment and produce many small eggs and smaller hatchlings. Such divergence is predicted as a direct effect of predation. We also found significant interactions between predator community and food level for age and size at maturity, fecundity, and egg size. These interactions, whereby the differences between communities were more pronounced at high-food levels, argue that evolution in Rivulus has been influenced by the indirect effects of predators mediated through resource availability. Rivulus from sites with predators better exploit the higher resources in those habitats. Therefore, both direct and indirect effects of predators have evolutionary consequences. [ABSTRACT FROM AUTHOR]

Guppies ( Poecilia reticulata) in Trinidadian streams are found with a diversity of predators in the lower reaches of streams, but few predators in the headwaters. These differences have caused the adaptive evolution of guppy behaviour, morphology, male colouration and life history. Waterfalls often serve as barriers to the upstream distribution of predators and/or guppies. Such discontinuities make it possible to treat streams like giant test tubes by introducing guppies or predators to small segments of streams from which they were previously excluded. Such experiments enable us to document how fast evolution can occur and the fine spatial scales over which adaptation is possible. They also demonstrate that the role predators play in structuring this ecosystem resembles many others studied from a more purely ecological perspective; in these streams, as elsewhere, predators depress the numbers of individuals in prey species which in turn reduces the effects of the prey species on other trophic levels and hence the structure of the ecosystem. A focus on predators is important in conservation biology because predators are often the organisms that are most susceptible to local extinction. Their selective loss occurs because large predators have been deliberately exterminated and/or are more susceptible to environmental disturbances. Furthermore, we will argue that predator re-introductions might be destabilizing if, in the absence of predators, their prey have evolved in a fashion that makes them highly susceptible to predation, even after time intervals as short as 50–100 years. A better understanding of the evolutionary impacts of top predators will be critical goal for the policy and practice of large carnivore restoration in the future. [ABSTRACT FROM AUTHOR]

Superfetation, the ability of females to simultaneously carry more than one brood at different developmental stages, is an unusual reproductive strategy that has independently evolved several times in the livebearing fish family Poeciliidae. Why this strategy has evolved remains uncertain. One hypothesis is that superfetation is a response to selective pressures that constrain the physical space within a female in which her offspring can develop. This hypothesis is reasonable, because superfetation should reduce the total volume needed to house developing embryos – that is, fewer large, fully developed embryos will be held by a superfetating female (with several broods at different developmental stages) than a non-superfetating female (where all embryos reach a fully developed stage at the same time). In this study, we explore this ‘morphological constraint’ hypothesis of superfetation by examining the livebearing fish, Poeciliopsis turrubarensis. We found that populations vary markedly in degree of superfetation, with individuals carrying from two to four distinct broods across different geographic areas. These populations also occupy a range of habitat types: some populations occur in slow moving coastal rivers near the ocean, while other populations occur far inland in fast moving waters that drain steep mountain environments. In comparing populations from these two types of environments, we find a strong association between stream habitat type and the degree of superfetation within populations. Fish from inland populations have higher levels of superfetation than their coastal counterparts. In addition, geometric morphometric analysis revealed that inland populations are also more fusiform than fish from coastal locations. Combined, these two lines of evidence support the ‘morphological constraint’ hypothesis, and suggest that the life history strategy of superfetation could be driven by environmental pressures that favor a more streamlined phenotype. [ABSTRACT FROM AUTHOR]

The colour pattern of male guppies ( Poecilia reticulata) is thought to evolve as a compromise between sexual selection (favouring conspicuousness) and natural selection (favouring crypsis). Underpinning this classic explanation is the observation that guppies living with dangerous fish predators are less colourful than guppies living without these predators. However, high fish-predation sites are generally farther downstream than low fish-predation sites, and so may also differ in physical habitat features related to stream size, as well as in the abundance of predatory prawns ( Macrobrachium crenulatum). The goal of our study was to disentangle the effects of fish predation on colour evolution from the potential effects of physical habitat features and predation by prawns. We collected 20 male guppies from each of 29 sites in two Trinidadian rivers. We then quantified the colour pattern of these fish; each spot was measured for size and assigned to a colour category. For each site, we determined the fish predation regime and quantified stream size, water colour, canopy openness, and prawn abundance. We then used regressions to assess the relative importance of these factors in explaining variation in guppy colour. Supporting previous work, the presence of predatory fishes was the most important explanatory variable for many components of colour pattern. Physical habitat features explained some of the remaining variation, but in inconsistent ways between the two rivers. The abundance of predatory prawns also explained variation in male colour. Our results suggest that predatory fishes impose the strongest selection on the colour pattern of male guppies but that other factors are also important. [ABSTRACT FROM AUTHOR]

Numerous studies have documented evolution by natural selectionatural selection in natural populations, but few are genuine selection experiments that are designed and then executed in nature. We will focus on these few cases to illustrate what can be learned from field selection experiments alone or field and laboratory selection experiments together that cannot be learned from laboratory experiments alone. Both types of study allow us to evaluate cause and effect relationships because a planned experiment can be accompanied by a more direct evaluation of the factors that cause evolution. A unique benefit of field experiments is that they give us the opportunity to measure the rate and magnitude of selection in nature. We have found that this rate is far greater than one might imagine based on observations of the fossil record. A combination of field and laboratory selection experiments has revealed the importance of population size and structure in shaping the genetics of adaptation. For example, laboratory selection experiments on insecticide resistance tend to attain resistance though polygenic inheritance. The evolution of insecticide resistance in nature often eventually yields to single genes of large effect that are rare but, once they arise, represent a higher fitness solution to resistance and spread among populations. Finally, field studies enable us to test evolutionary theory in a context in which all of the tradeoffs associated with a trait are realized; in the laboratory, organisms may be shielded from the fitness tradeoffs associated with the evolution of a trait. For example, we have compared the patterns of senescence in guppies from high and low mortality rate environments in the laboratory and in the field. In the laboratory, guppies from high predation environments had delayed senescence relative to those from low predation environments. In the field the apparent relationship is the opposite. One hypothesis for this difference is that a tradeoff associated with the evolution of the high predation life history is a decrease in the investment in the immune system. Such a sacrifice would be evident in nature where there is exposure to disease and parasites but less so in the laboratory, which is relatively disease and parasite free. [ABSTRACT FROM AUTHOR]

Life history theory predicts that high adult mortality rates select for earlier maturity and increased reproduction. If such evolution occurs in response to the commercial exploitation of natural fish populations, then the correlated reduction in body size would reduce the yield of the fishery. Earlier maturity and reduced body size are seen in commercially exploited populations. Here, we compare the life histories of natural populations of guppies (Poecilia reticulata) from Trinidad that live in either high- or low-predation environments, which serve as surrogates for the presence or absence of commercial fishing. We can quantify mortality rate and life history variables, including age and size at maturity, in the laboratory and in nature. We have manipulated mortality rates in nature and measured the rate of evolution. High mortality selects for earlier maturity at a smaller size, as observed in commercial fisheries and as predicted by theory. Furthermore, the nature and magnitude of predator-induced mortality are comparable to those caused by commercial fishing. The rate of evolution in guppies predicts similar evolution in commercial fisheries on a time scale of decades. These attributes support arguments that humans, like predators, have acted as an agent of selection when exploiting populations of fish. [ABSTRACT FROM AUTHOR]

Classical theories for the evolution of senescence predict that organisms that experience low mortality rates attributable to external factors, such as disease or predation, will evolve a later onset of senescence. Here we use patterns of senescence in guppies derived from natural populations that differ in mortality risk to evaluate the generality of these predictions. We have previously found that populations experiencing higher mortality rates evolve earlier maturity and invest more in reproduction, as predicted by evolutionary theory. We report here that these same populations do not have an earlier onset of senescence with respect to either mortality or reproduction but do with respect to swimming performance, which assesses neuromuscular function. This mosaic pattern of senescence challenges the generality of the association between decreased extrinsic mortality and delayed senescence and invites consideration of more derived theories for the evolution of senescence. [ABSTRACT FROM AUTHOR]

The empirical study of natural selection reveals that adaptations often involve trade-offs between competing functions. Because natural selection acts on whole organisms rather than isolated traits, adaptive evolution may be constrained by the interaction between traits that are functionally integrated. Yet, few attempts have been made to characterize how and when such constraints are manifested or whether they limit the adaptive divergence of populations. Here we examine the consequences of adaptive life-history evolution on locomotor performance in the live-bearing guppy. In response to increased predation from piscivorous fish, Trinidadian guppies evolve an increased allocation of resources toward reproduction. These populations are also under strong selection for rapid fast-start swimming performance to evade predators. Because embryo development increases a female's wet mass as she approaches parturition, an increased investment in reproductive allocation should impede fast-start performance. We find evidence for adaptive but constrained evolution of fast-start swimming performance in laboratory trials conducted on second-generation lab-reared fish. Female guppies from high-predation localities attain a faster acceleration and velocity and travel a greater distance during fast-start swimming trials. However, velocity and distance traveled decline more rapidly over the course of pregnancy in these same females, thus reducing the magnitude of divergence in swimming performance between high- and low-predation populations. This functional trade-off between reproduction and swimming performance reveals how different aspects of the phenotype are integrated and highlights the complexity of adaptation at the whole-organism level. [ABSTRACT FROM AUTHOR]

We evaluated differences in larval habitats and life history of three species of spadefoot toads, then compared their life histories in a common garden study. Our field work defined the selective regime encountered by each species. Our Great Basin spadefoot (S pea intermontana) bred asynchronously in permanent streams and springs where there was no risk of larval mortality due to drying. The water chemistry remained fairly stable throughout the larval period. The western spadefoot toad, Sp. hammondii, bred fairly synchronously following heavy spring rains in temporary pools that remained filled an average of 81 d. Fifteen % of the breeding pools dried completely on or before the day the first larvae metamorphosed. The desert spadefoot toad, Scaphiopus couchii, bred synchronously after heavy summer showers in very short duration pools; 62% of the breeding pools dried completely on or before the day the first larvae metamorphosed. The concentration of ammonium nitrogen and CaCO3 increased markedly as the Sp. hammondii and S. couchii pools dried. S. couchii attained metamorphosis at a much earlier age and smaller size than the other two species. S. couchii also showed little variation in the age at metamorphosis but considerable variation in the size at metamorphosis, while the other two species varied in both age and size. The results identify some variables that could serve as cues of pool drying and demonstrate an association between breeding pool duration, breeding synchrony, development rate, and larval development. Our laboratory study yields information about the genetic basis of the differences in development and controlled comparisons of phenotypic plasticity. We manipulated food supply to study the plastic response of age and size at metamorphosis and hence construct the reaction norm for these variables as a function of growth rate. The growth rates ranged from below to above those observed in natural populations. As in the field, in the lab S. couchii attained metamorphosis at an earlier age and smaller size than the other two species. All three species had a similarly shaped reaction norm for size(y-axis) and age (x-axis) at metamorphosis, which was a concave upward curve. A consequence of this shape is that age at metamorphosis changes more readily at low levels of food availability and size at metamorphosis changes more readily at high levels of food availability. If we restrict our observations to just those growth rates that are seen in nature, then S. couchii has almost no variation in the age at metamorphosis but considerable variation in size at metamorphosis, while the other two species vary in both age and size at metamorphosis. All three species increased in size at metamorphosis with increased food levels. Our comparative reaction norm approach thus demonstrates that S. couchii has adapted to ephemeral environments by shifting its growth rate reaction norm so that age at metamorphosis is uniformly fast and is not associated with growth rate. The realized variation is concentrated in size rather than age at metamorphosis. [ABSTRACT FROM AUTHOR]

Studies the guppies' responses to social environment through phenotypic plasticity in the guppies' life history traits. Contribution of induced phenotypic variation to the guppies' life history differences in Rivulus hartii and Chrenicichla alta communities; Evolution of the guppies' patterns of plasticity due to the manipulation of the social environment.

Studies the importance of predation and life histories in the demography of guppy populations. Observation of fewer small immature guppies coexisting with Rivulus hartii than those living with larger predators; Attribution of variation in adult sex ratio to the mortality and individual growth rates.

Estimates the life-history patterns of guppies in natural populations. Comparative mortality rates of guppies; Assumption that predation is the primary cause of the increased mortality rates of guppies; Mechanism that accounts for the evolution of life-history patterns in guppies.

Reports on the rate of evolution in natural populations of guppies (Poecilia reticulata). Rate of evolution of age and size under natural selection similar to those under artificial selection; Rate of evolution of male and female traits; Implications.

Documents a genetic basis for convergent life-history evolution in guppies from high and low predation sites on the north slope of the Northern Range Mountains of Trinidad. Controlled food availability; Estimation of reproductive efforts; Predator effects; Reasons for convergence.

Compares guppies from high-predation sites (Crenichichla localities) with those from low-predation sites (Rivulus localities) found on the south slope of the Northern Range Mountains of Trinidad. Characterization of the life-history of phenotype; Multivariate comparison of female life-history traits.

The evolution of complex organs is a source of controversy because they require the contributions of many adaptations to function properly. We argue that placentas are complex, that they have evolved multiple times in Poeciliopsis, and that there are closely related sister taxa that have either no placentas or intermediate stages in the evolution of a placenta. Furthermore, placentas can evolve in 750,000 years or less, on the same time scale as suggested by theoretical calculations for the evolution of complex eyes. Independent origins of such complexity, accompanied by sister taxa that either lack or have intermediate stages in the evolution of the trait, present an opportunity to study the evolution of novelty and complexity from a comparative, evolutionary perspective. [ABSTRACT FROM AUTHOR]

The placenta is a complex life‐history trait that is ubiquitous across the tree of life. Theory proposes that the placenta evolves in response to high performance‐demanding conditions by shifting maternal investment from pre‐ to post‐fertilisation, thereby reducing a female's reproductive burden during pregnancy. We test this hypothesis by studying populations of the fish species Poeciliopsis retropinna in Costa Rica. We found substantial variation in the degree of placentation among natural populations associated with predation risk: females from high predation populations had significantly higher degrees of placentation compared to low predation females, while number, size and quality of offspring at birth remained unaffected. Moreover, a higher degree of placentation correlated with a lower reproductive burden and hence likely an improved swimming performance during pregnancy. Our study advances an adaptive explanation for why the placenta evolves by arguing that an increased degree of placentation offers a selective advantage in high predation environments. [ABSTRACT FROM AUTHOR]

Adaptive behavioral plasticity can play a beneficial role when a population becomes established in a novel environment if environmental cues allow the expression of appropriate behavior. Further, plasticity itself can evolve over time in a new environment causing changes in the way or degree to which animals respond to environmental cues. Colonization events provide an opportunity to investigate such relationships between behavioral plasticity and adaptation to new environments. Here, we investigated the evolution of behavior and behavioral plasticity during colonization of a new environment, by testing if female mate‐choice behavior diverged in Trinidadian guppies 2–3 years (~6–9 generations) after being introduced to four locations with reduced predation risk. We collected wild‐caught fish from the source and introduced populations, and we reared out second‐generation females in the laboratory with and without predator cues to examine their plastic responses to a bright and dull male. We found introduced females were less responsive to males when reared without predator cues, but both introduced and source females were similarly responsive when reared with predator cues. Thus, the parallel evolution of behavior across multiple populations in the low‐predation environment was only observed in the treatment mimicking the introduction environment. Such results are consistent with theory predicting that the evolution of plasticity is a by‐product of differential selection across environments. [ABSTRACT FROM AUTHOR]

Maternal effects often provide a mechanism for adaptive transgenerational phenotypic plasticity. The maternal phenotype can profoundly influence the potential for such environmentally induced adjustments of the offspring phenotype, causing correlations between offspring and maternal traits. Here, we study potential effects of the maternal phenotype on offspring provisioning prior to and during gestation in the matrotrophic live‐bearing fish species Poeciliopsis retropinna. Specifically, we examine how maternal traits such as body fat, lean mass, and length relate to pre‐ (i.e., allocation to the egg prior to fertilization) and post‐fertilization (i.e., allocation to the embryo during pregnancy) maternal provisioning and how this ultimately affects offspring size and body composition at birth. We show that pre‐ and post‐fertilization maternal provisioning is associated with maternal length and body fat, but not with maternal lean mass. Maternal length is proportionally associated with egg mass at fertilization and offspring mass at birth, notably without changing the ratio of pre‐ to post‐fertilization maternal provisioning. This ratio, referred to as the matrotrophy index (MI), is often used to quantify the level of matrotrophy. By contrast, the proportion of maternal body fat is positively associated with post‐fertilization, but not pre‐fertilization, maternal provisioning and consequently is strongly positively correlated with the MI. We furthermore found that the composition of embryos changes throughout pregnancy. Females invest first in embryo lean mass, and then allocate fat reserves to embryos very late in pregnancy. We argue that this delay in fat allocation may be adaptive, because it delays an unnecessary high reproductive burden to the mother during earlier stages of pregnancy, potentially leading to a more slender body shape and improved locomotor performance. In conclusion, our study suggests that (a) offspring size at birth is a plastic trait that is predicted by both maternal length and body fat, and (b) the MI is a plastic trait that is predicted solely by the proportion of maternal body fat. It herewith provides new insights into the potential maternal causes and consequences of embryo provisioning during pregnancy in matrotrophic live‐bearing species. Maternal effects are particularly important, frequently having widespread consequences for offspring phenotype and fitness. Here, we studied potential effects of the maternal phenotype on offspring provisioning prior to and during gestation in the matrotrophic live‐bearing fish species Poeciliopsis retropinna. We show that maternal traits strongly correlate with embryo size and body composition throughout pregnancy. Though, different maternal traits are differently related to pre‐ and post‐fertilization maternal provisioning. We furthermore found that the composition of embryos changes during pregnancy independently of the maternal phenotype. [ABSTRACT FROM AUTHOR]

Examining the association between trait variation and development is crucial for understanding the evolution of phenotypic differences. Male guppy ornamental caudal fin coloration is one trait that shows a striking degree of variation within and between guppy populations. Males initially have no caudal fin coloration, then gradually develop it as they reach sexual maturity. For males, there is a trade‐off between female preference for caudal fin coloration and increased visibility to predators. This trade‐off may reach unique endpoints in males from different predation regimes. Caudal fin coloration includes black melanin, orange/yellow pteridines or carotenoids, and shimmering iridescence. This study examined the phenotypic trajectory and genetics associated with color development. We found that black coloration always developed first, followed by orange/yellow, then iridescence. The ordering and timing of color appearance was the same regardless of predation regime. The increased expression of melanin synthesis genes correlated well with the visual appearance of black coloration, but there was no correlation between carotenoids or pteridine synthesis gene expression and the appearance of orange/yellow. The lack of orange/yellow coloration in earlier male caudal fin developmental stages may be due to reduced expression of genes underlying the development of orange/yellow xanthophores. [ABSTRACT FROM AUTHOR]

The question of whether populations evolve predictably and consistently under similar selective regimes is fundamental to understanding how adaptation proceeds in the wild. We address this question with a replicated evolution experiment focused upon male sexual coloration in guppies (Poecilia reticulata). Fish were transplanted from a single high predation population in the Guanapo River to four replicate, guppy‐free low predation headwater streams. Two streams had their canopies thinned to adjust the setting under which male coloration is displayed and perceived. We assessed evolutionary divergence using second‐generation lab‐bred offspring of fish sampled four to six years following translocation. A prior experiment of the same design, performed in an adjacent drainage, resulted in the evolution of more extensive orange, black, and iridescent markings. We however found evidence for expansion only in structural coloration (iridescent blue/green), no change in orange, and a reduction in black. This response amplifies earlier findings for Guanapo fish, revealing that trajectories of color elaboration differ among drainages. We also found that color phenotypes evolved more greatly at the thinned‐canopy sites. Our findings support the predictability of sexual trait evolution in the wild, and underscore the importance of signaling conditions and ornamental starting points in shaping adaptive trajectories. [ABSTRACT FROM AUTHOR]

The article explores the concept of phenotypic convergence, where similar traits evolve independently in different lineages due to shared selective pressures. It focuses on the evolution of the placenta in closely related fish species (Poeciliidae) as a case study. It further reveals two distinct types of placental structures and examines their genetic and morphological differences.