Despite appearing featureless to our eyes, the open ocean is a highly variable environment for po... more Despite appearing featureless to our eyes, the open ocean is a highly variable environment for polarization-sensitive viewers. Dynamic visual backgrounds coupled with predator encounters from all possible directions make this habitat one of the most challenging for camouflage. We tested open-ocean crypsis in nature by collecting more than 1500 videopolarimetry measurements from live fish from distinct habitats under a variety of viewing conditions. Open-ocean fish species exhibited camouflage that was superior to that of both nearshore fish and mirrorlike surfaces, with significantly higher crypsis at angles associated with predator detection and pursuit. Histological measurements revealed that specific arrangements of reflective guanine platelets in the fish's skin produce angle-dependent polarization modifications for polarocrypsis in the open ocean, suggesting a mechanism for natural selection to shape reflectance properties in this complex environment.
Social interactions can drive distinct gene expression profiles which may vary by social context.... more Social interactions can drive distinct gene expression profiles which may vary by social context. Here we use female sailfin molly fish (Poecilia latipinna) to identify genomic profiles associated with preference behavior in distinct social contexts: male interactions (mate choice) versus female interactions (shoaling partner preference). We measured the behavior of 15 females interacting in a non‐contact environment with either two males or two females for 30 min followed by whole‐brain transcriptomic profiling by RNA sequencing. We profiled females that exhibited high levels of social affiliation and great variation in preference behavior to identify an order of magnitude more differentially expressed genes associated with behavioral variation than by differences in social context. Using a linear model (limma), we took advantage of the individual variation in preference behavior to identify unique gene sets that exhibited distinct correlational patterns of expression with preferen...
Antipredator defenses and warning signals typically evolve in concert. However, the extensive var... more Antipredator defenses and warning signals typically evolve in concert. However, the extensive variation across taxa in both these components of predator deterrence and the relationship between them are poorly understood. Here we test whether there is a predictive relationship between visual conspicuousness and toxicity levels across 10 populations of the color-polymorphic strawberry poison frog, Dendrobates pumilio. Using a mouse-based toxicity assay, we find extreme variation in toxicity between frog populations. This variation is significantly positively correlated with frog coloration brightness, a viewer-independent measure of visual conspicuousness (i.e., total reflectance flux). We also examine conspicuousness from the view of three potential predator taxa, as well as conspecific frogs, using taxon-specific visual detection models and three natural background substrates. We find very strong positive relationships between frog toxicity and conspicuousness for bird-specific perceptual models. Weaker but still positive correlations are found for crab and D. pumilio conspecific visual perception, while frog coloration as viewed by snakes is not related to toxicity. These results suggest that poison frog colors can be honest signals of prey unpalatability to predators and that birds in particular may exert selection on aposematic signal design.
Proceedings of the National Academy of Sciences, 2009
It is commonly assumed that natural selection imposed by predators is the prevailing force drivin... more It is commonly assumed that natural selection imposed by predators is the prevailing force driving the evolution of aposematic traits. Here, we demonstrate that aposematic signals are shaped by sexual selection as well. We evaluated sexual selection for coloration brightness in populations of the poison frog Oophaga [Dendrobates] pumilio in Panama's Bocas del Toro archipelago. We assessed female preferences for brighter males by manipulating the perceived brightness of spectrally matched males in two-way choice experiments. We found strong female preferences for bright males in two island populations and weaker or ambiguous preferences in females from mainland populations. Spectral reflectance measurements, coupled with an O. pumilio -specific visual processing model, showed that O. pumilio coloration was significantly brighter in island than in mainland morphs. In one of the island populations (Isla Solarte), males were significantly more brightly colored than females. Taken to...
The evolution and maintenance of colour polymorphisms within species has emerged as an important ... more The evolution and maintenance of colour polymorphisms within species has emerged as an important focus of attention within evolutionary biology (Gray & McKinnon, 2007). Research on such polymorphisms has enhanced our understanding of the evolutionary forces that maintain phenotypic variation in nature (Magurran,
Aposematic signals may be subject to conflicting selective pressures from predators and conspecif... more Aposematic signals may be subject to conflicting selective pressures from predators and conspecifics. We studied female preferences for different components of aposematic coloration in the polymorphic poison frog Oophaga pumilio across several phenotypically distinct populations. This frog shows striking diversity in color and pattern between geographically isolated populations in western Panama. Results indicate that male dorsal color is the most important determiner of female preferences. We did not find consistent evidence for effects of other signal components, such as spotting pattern or ventral color. Females in two populations showed assortative preferences mediated by male dorsal coloration. In a third population we found incomplete color-assortative preference behavior, with females exhibiting strong discrimination toward one novel color but not another. These results hint at a possible interaction between sexual and natural selection: female tolerance of unfamiliar coloration patterns could facilitate the establishment of novel phenotypes that are favored by other selective pressures (e.g., predator biases). Furthermore, our study suggests that specific components of the aposematic signal (i.e., dorsal color, ventral color, and spotting pattern) are affected differently by natural and sexual selection.
Circularly polarized light is rare in the terrestrial environment, and cuticular reflections from... more Circularly polarized light is rare in the terrestrial environment, and cuticular reflections from scarab beetles are one of the few natural sources. Chrysina gloriosa LeConte 1854, a scarab beetle found in montane juniper forests of the extreme southwestern United States and northern Mexico, are camouflaged in juniper foliage; however, when viewed with right circularly polarizing filters, the beetles exhibit a stark black contrast. Given the polarization-specific changes in the appearance of C. gloriosa, we hypothesized that C. gloriosa can detect circularly polarized light. We tested for phototactic response and differential flight orientation of C. gloriosa toward different light stimuli. Chrysina gloriosa exhibited (a) positive phototaxis, (b) differential flight orientation between linear and circularly polarized light stimuli of equal intensities, and (c) discrimination between circularly polarized and unpolarized lights of different intensities consistent with a model of circular polarization sensitivity based on a quarter-wave plate. These results demonstrate that C. gloriosa beetles respond differentially to circularly polarized light. In contrast, Chrysina woodi Horn 1885, a close relative with reduced circularly polarized reflection, exhibited no phototactic discrimination between linear and circularly polarized light. Circularly polarized sensitivity may allow C. gloriosa to perceive and communicate with conspecifics that remain cryptic to predators, reducing indirect costs of communication.
The relationship between an individual's cognitive abilities and other behavioural attributes... more The relationship between an individual's cognitive abilities and other behavioural attributes is complex, yet critical to understanding how individual differences in cognition arise. Here we use western mosquitofish, Gambusia affinis, to investigate the relationship between individual associative learning performance in numerical discrimination tests and independent measures of activity, exploration, anxiety and sociability. We found extensive and highly repeatable inter-individual variation in learning performance (r = 0.89; ICC = 0.89). Males and females exhibited similar learning performance, yet differed in sociability, activity and their relationship between learning and anxiety/exploration tendencies. Sex-specific multivariate behaviour scores successfully predicted variation in individual learning performance, whereas combined sex analyses did not. Female multivariate behaviour scores significantly predict learning performance across females (ρ = 0.80, p = 0.005) with hig...
Journal of the Royal Society, Interface / the Royal Society, Jan 6, 2015
Predator evasion in the open ocean is difficult because there are no objects to hide behind. The ... more Predator evasion in the open ocean is difficult because there are no objects to hide behind. The silvery surface of fish plays an important role in open water camouflage. Various models have been proposed to account for the broadband reflectance by the fish skin that involve one-dimensional variations in the arrangement of guanine crystal reflectors, yet the three-dimensional organization of these guanine platelets have not been well characterized. Here, we report the three-dimensional organization and the optical properties of integumentary guanine platelets in a silvery marine fish, the lookdown (Selene vomer). Our structural analysis and computational modelling show that stacks of guanine platelets with random yaw angles in the fish skin produce broadband reflectance via colour mixing. Optical axes of the guanine platelets and the collagen layer are aligned closely and provide bulk birefringence properties that influence the polarization reflectance by the skin. These data demons...
It has been 25 years since the formalization of the Sensory Drive hypothesis was published in the... more It has been 25 years since the formalization of the Sensory Drive hypothesis was published in the American Naturalist (1992). Since then, there has been an explosion of research identifying its utility in contributing to our understanding of inter-and intra-specific variation in sensory systems and signaling properties. The main tenet of Sensory Drive is that environmental characteristics will influence the evolutionary trajectory of both sensory (detecting capabilities) and signaling (detectable features and behaviors) traits in predictable directions. We review the accumulating evidence in 154 studies addressing these questions and categorized their approach in terms of testing for environmental influence on sensory tuning, signal characteristics, or both. For the subset of studies that examined sensory tuning, there was greater support for Sensory Drive processes shaping visual than auditory tuning, and it was more prevalent in aquatic than terrestrial habitats. Terrestrial habitats and visual traits were the prevalent habitat and sensory modality in the 104 studies showing support for environmental influence on signaling properties. An additional 19 studies that found no supporting evidence for environmental influence on signaling traits were all based in terrestrial ecosystems and almost exclusively involved auditory signals. Only 29 studies examined the complete coevolutionary process between sensory and signaling traits and were dominated by fish visual communication. We discuss biophysical factors that may contribute to the visual and aquatic bias for Sensory Drive evidence, as well as biotic factors that may contribute to the lack of Sensory Drive processes in terrestrial acoustic signaling systems.
Batesian mimicry-resemblance of a toxic model by an edible mimic-depends on deceiving predators. ... more Batesian mimicry-resemblance of a toxic model by an edible mimic-depends on deceiving predators. Mimetic advantage is considered to be dependent on frequency because an increase in mimic abundance leads to breakdown of the warning signal. Where multiple toxic species are available, batesian polymorphism is predicted-that is, mimics diversify to match sympatric models. Despite the prevalence of batesian mimicry in nature,
Despite appearing featureless to our eyes, the open ocean is a highly variable environment for po... more Despite appearing featureless to our eyes, the open ocean is a highly variable environment for polarization-sensitive viewers. Dynamic visual backgrounds coupled with predator encounters from all possible directions make this habitat one of the most challenging for camouflage. We tested open-ocean crypsis in nature by collecting more than 1500 videopolarimetry measurements from live fish from distinct habitats under a variety of viewing conditions. Open-ocean fish species exhibited camouflage that was superior to that of both nearshore fish and mirrorlike surfaces, with significantly higher crypsis at angles associated with predator detection and pursuit. Histological measurements revealed that specific arrangements of reflective guanine platelets in the fish's skin produce angle-dependent polarization modifications for polarocrypsis in the open ocean, suggesting a mechanism for natural selection to shape reflectance properties in this complex environment.
Keywords: female mate preference female size male courtship behaviour poeciliid sexual experience... more Keywords: female mate preference female size male courtship behaviour poeciliid sexual experience Xiphophorus Building a complete model of animal behaviour requires knowing both the internal and external factors that influence behaviour. As recent genomic and neural studies begin to establish the northern swordtail, Xiphophorus nigrensis, as a model for proximate explorations of female mate preference, it is important to recognize the complexity of other factors that can drive variation in female preference. In this study, we determined how different experiential and social factors correlate with interindividual variation in female mate preference: female body size, sexual experience and male behavioural displays. We found a significant positive correlation between female size and female preference for large males, as well as a significant relationship between male activity and female preference. We also demonstrate that copulating at least once may be sufficient to increase preference for large males. Our identification of these relationships emphasizes the multiple and diverse types of mechanisms that can influence variation in female mate preference and encourage their consideration in future studies when trying to assess why and how females choose males. Ó
Trinidad guppies (Poecilia reticulata) are distributed along an environmental gradient in caroten... more Trinidad guppies (Poecilia reticulata) are distributed along an environmental gradient in carotenoid avail- ability that limits the carotenoid content of the orange spots of males. The amount of synthetic red pteridines (dro- sopterins) in the orange spots covaries with the carotenoid content, such that the ratio of the two types of pigments is roughly conserved across streams. Carotenoids and drosopterins
Proceedings of The National Academy of Sciences, 2006
Many animals advertise their chemical defense to predators with conspicuous coloration and unpala... more Many animals advertise their chemical defense to predators with conspicuous coloration and unpalatability, but little is known about the information in these signal elements. To effectively avoid predation, is it more advantageous to invest in increased conspicuousness or greater noxiousness, or to allocate equally to both signal modalities? Using natural variation among poison frog species measured with spectral reflectance and
Despite appearing featureless to our eyes, the open ocean is a highly variable environment for po... more Despite appearing featureless to our eyes, the open ocean is a highly variable environment for polarization-sensitive viewers. Dynamic visual backgrounds coupled with predator encounters from all possible directions make this habitat one of the most challenging for camouflage. We tested open-ocean crypsis in nature by collecting more than 1500 videopolarimetry measurements from live fish from distinct habitats under a variety of viewing conditions. Open-ocean fish species exhibited camouflage that was superior to that of both nearshore fish and mirrorlike surfaces, with significantly higher crypsis at angles associated with predator detection and pursuit. Histological measurements revealed that specific arrangements of reflective guanine platelets in the fish's skin produce angle-dependent polarization modifications for polarocrypsis in the open ocean, suggesting a mechanism for natural selection to shape reflectance properties in this complex environment.
Social interactions can drive distinct gene expression profiles which may vary by social context.... more Social interactions can drive distinct gene expression profiles which may vary by social context. Here we use female sailfin molly fish (Poecilia latipinna) to identify genomic profiles associated with preference behavior in distinct social contexts: male interactions (mate choice) versus female interactions (shoaling partner preference). We measured the behavior of 15 females interacting in a non‐contact environment with either two males or two females for 30 min followed by whole‐brain transcriptomic profiling by RNA sequencing. We profiled females that exhibited high levels of social affiliation and great variation in preference behavior to identify an order of magnitude more differentially expressed genes associated with behavioral variation than by differences in social context. Using a linear model (limma), we took advantage of the individual variation in preference behavior to identify unique gene sets that exhibited distinct correlational patterns of expression with preferen...
Antipredator defenses and warning signals typically evolve in concert. However, the extensive var... more Antipredator defenses and warning signals typically evolve in concert. However, the extensive variation across taxa in both these components of predator deterrence and the relationship between them are poorly understood. Here we test whether there is a predictive relationship between visual conspicuousness and toxicity levels across 10 populations of the color-polymorphic strawberry poison frog, Dendrobates pumilio. Using a mouse-based toxicity assay, we find extreme variation in toxicity between frog populations. This variation is significantly positively correlated with frog coloration brightness, a viewer-independent measure of visual conspicuousness (i.e., total reflectance flux). We also examine conspicuousness from the view of three potential predator taxa, as well as conspecific frogs, using taxon-specific visual detection models and three natural background substrates. We find very strong positive relationships between frog toxicity and conspicuousness for bird-specific perceptual models. Weaker but still positive correlations are found for crab and D. pumilio conspecific visual perception, while frog coloration as viewed by snakes is not related to toxicity. These results suggest that poison frog colors can be honest signals of prey unpalatability to predators and that birds in particular may exert selection on aposematic signal design.
Proceedings of the National Academy of Sciences, 2009
It is commonly assumed that natural selection imposed by predators is the prevailing force drivin... more It is commonly assumed that natural selection imposed by predators is the prevailing force driving the evolution of aposematic traits. Here, we demonstrate that aposematic signals are shaped by sexual selection as well. We evaluated sexual selection for coloration brightness in populations of the poison frog Oophaga [Dendrobates] pumilio in Panama's Bocas del Toro archipelago. We assessed female preferences for brighter males by manipulating the perceived brightness of spectrally matched males in two-way choice experiments. We found strong female preferences for bright males in two island populations and weaker or ambiguous preferences in females from mainland populations. Spectral reflectance measurements, coupled with an O. pumilio -specific visual processing model, showed that O. pumilio coloration was significantly brighter in island than in mainland morphs. In one of the island populations (Isla Solarte), males were significantly more brightly colored than females. Taken to...
The evolution and maintenance of colour polymorphisms within species has emerged as an important ... more The evolution and maintenance of colour polymorphisms within species has emerged as an important focus of attention within evolutionary biology (Gray & McKinnon, 2007). Research on such polymorphisms has enhanced our understanding of the evolutionary forces that maintain phenotypic variation in nature (Magurran,
Aposematic signals may be subject to conflicting selective pressures from predators and conspecif... more Aposematic signals may be subject to conflicting selective pressures from predators and conspecifics. We studied female preferences for different components of aposematic coloration in the polymorphic poison frog Oophaga pumilio across several phenotypically distinct populations. This frog shows striking diversity in color and pattern between geographically isolated populations in western Panama. Results indicate that male dorsal color is the most important determiner of female preferences. We did not find consistent evidence for effects of other signal components, such as spotting pattern or ventral color. Females in two populations showed assortative preferences mediated by male dorsal coloration. In a third population we found incomplete color-assortative preference behavior, with females exhibiting strong discrimination toward one novel color but not another. These results hint at a possible interaction between sexual and natural selection: female tolerance of unfamiliar coloration patterns could facilitate the establishment of novel phenotypes that are favored by other selective pressures (e.g., predator biases). Furthermore, our study suggests that specific components of the aposematic signal (i.e., dorsal color, ventral color, and spotting pattern) are affected differently by natural and sexual selection.
Circularly polarized light is rare in the terrestrial environment, and cuticular reflections from... more Circularly polarized light is rare in the terrestrial environment, and cuticular reflections from scarab beetles are one of the few natural sources. Chrysina gloriosa LeConte 1854, a scarab beetle found in montane juniper forests of the extreme southwestern United States and northern Mexico, are camouflaged in juniper foliage; however, when viewed with right circularly polarizing filters, the beetles exhibit a stark black contrast. Given the polarization-specific changes in the appearance of C. gloriosa, we hypothesized that C. gloriosa can detect circularly polarized light. We tested for phototactic response and differential flight orientation of C. gloriosa toward different light stimuli. Chrysina gloriosa exhibited (a) positive phototaxis, (b) differential flight orientation between linear and circularly polarized light stimuli of equal intensities, and (c) discrimination between circularly polarized and unpolarized lights of different intensities consistent with a model of circular polarization sensitivity based on a quarter-wave plate. These results demonstrate that C. gloriosa beetles respond differentially to circularly polarized light. In contrast, Chrysina woodi Horn 1885, a close relative with reduced circularly polarized reflection, exhibited no phototactic discrimination between linear and circularly polarized light. Circularly polarized sensitivity may allow C. gloriosa to perceive and communicate with conspecifics that remain cryptic to predators, reducing indirect costs of communication.
The relationship between an individual's cognitive abilities and other behavioural attributes... more The relationship between an individual's cognitive abilities and other behavioural attributes is complex, yet critical to understanding how individual differences in cognition arise. Here we use western mosquitofish, Gambusia affinis, to investigate the relationship between individual associative learning performance in numerical discrimination tests and independent measures of activity, exploration, anxiety and sociability. We found extensive and highly repeatable inter-individual variation in learning performance (r = 0.89; ICC = 0.89). Males and females exhibited similar learning performance, yet differed in sociability, activity and their relationship between learning and anxiety/exploration tendencies. Sex-specific multivariate behaviour scores successfully predicted variation in individual learning performance, whereas combined sex analyses did not. Female multivariate behaviour scores significantly predict learning performance across females (ρ = 0.80, p = 0.005) with hig...
Journal of the Royal Society, Interface / the Royal Society, Jan 6, 2015
Predator evasion in the open ocean is difficult because there are no objects to hide behind. The ... more Predator evasion in the open ocean is difficult because there are no objects to hide behind. The silvery surface of fish plays an important role in open water camouflage. Various models have been proposed to account for the broadband reflectance by the fish skin that involve one-dimensional variations in the arrangement of guanine crystal reflectors, yet the three-dimensional organization of these guanine platelets have not been well characterized. Here, we report the three-dimensional organization and the optical properties of integumentary guanine platelets in a silvery marine fish, the lookdown (Selene vomer). Our structural analysis and computational modelling show that stacks of guanine platelets with random yaw angles in the fish skin produce broadband reflectance via colour mixing. Optical axes of the guanine platelets and the collagen layer are aligned closely and provide bulk birefringence properties that influence the polarization reflectance by the skin. These data demons...
It has been 25 years since the formalization of the Sensory Drive hypothesis was published in the... more It has been 25 years since the formalization of the Sensory Drive hypothesis was published in the American Naturalist (1992). Since then, there has been an explosion of research identifying its utility in contributing to our understanding of inter-and intra-specific variation in sensory systems and signaling properties. The main tenet of Sensory Drive is that environmental characteristics will influence the evolutionary trajectory of both sensory (detecting capabilities) and signaling (detectable features and behaviors) traits in predictable directions. We review the accumulating evidence in 154 studies addressing these questions and categorized their approach in terms of testing for environmental influence on sensory tuning, signal characteristics, or both. For the subset of studies that examined sensory tuning, there was greater support for Sensory Drive processes shaping visual than auditory tuning, and it was more prevalent in aquatic than terrestrial habitats. Terrestrial habitats and visual traits were the prevalent habitat and sensory modality in the 104 studies showing support for environmental influence on signaling properties. An additional 19 studies that found no supporting evidence for environmental influence on signaling traits were all based in terrestrial ecosystems and almost exclusively involved auditory signals. Only 29 studies examined the complete coevolutionary process between sensory and signaling traits and were dominated by fish visual communication. We discuss biophysical factors that may contribute to the visual and aquatic bias for Sensory Drive evidence, as well as biotic factors that may contribute to the lack of Sensory Drive processes in terrestrial acoustic signaling systems.
Batesian mimicry-resemblance of a toxic model by an edible mimic-depends on deceiving predators. ... more Batesian mimicry-resemblance of a toxic model by an edible mimic-depends on deceiving predators. Mimetic advantage is considered to be dependent on frequency because an increase in mimic abundance leads to breakdown of the warning signal. Where multiple toxic species are available, batesian polymorphism is predicted-that is, mimics diversify to match sympatric models. Despite the prevalence of batesian mimicry in nature,
Despite appearing featureless to our eyes, the open ocean is a highly variable environment for po... more Despite appearing featureless to our eyes, the open ocean is a highly variable environment for polarization-sensitive viewers. Dynamic visual backgrounds coupled with predator encounters from all possible directions make this habitat one of the most challenging for camouflage. We tested open-ocean crypsis in nature by collecting more than 1500 videopolarimetry measurements from live fish from distinct habitats under a variety of viewing conditions. Open-ocean fish species exhibited camouflage that was superior to that of both nearshore fish and mirrorlike surfaces, with significantly higher crypsis at angles associated with predator detection and pursuit. Histological measurements revealed that specific arrangements of reflective guanine platelets in the fish's skin produce angle-dependent polarization modifications for polarocrypsis in the open ocean, suggesting a mechanism for natural selection to shape reflectance properties in this complex environment.
Keywords: female mate preference female size male courtship behaviour poeciliid sexual experience... more Keywords: female mate preference female size male courtship behaviour poeciliid sexual experience Xiphophorus Building a complete model of animal behaviour requires knowing both the internal and external factors that influence behaviour. As recent genomic and neural studies begin to establish the northern swordtail, Xiphophorus nigrensis, as a model for proximate explorations of female mate preference, it is important to recognize the complexity of other factors that can drive variation in female preference. In this study, we determined how different experiential and social factors correlate with interindividual variation in female mate preference: female body size, sexual experience and male behavioural displays. We found a significant positive correlation between female size and female preference for large males, as well as a significant relationship between male activity and female preference. We also demonstrate that copulating at least once may be sufficient to increase preference for large males. Our identification of these relationships emphasizes the multiple and diverse types of mechanisms that can influence variation in female mate preference and encourage their consideration in future studies when trying to assess why and how females choose males. Ó
Trinidad guppies (Poecilia reticulata) are distributed along an environmental gradient in caroten... more Trinidad guppies (Poecilia reticulata) are distributed along an environmental gradient in carotenoid avail- ability that limits the carotenoid content of the orange spots of males. The amount of synthetic red pteridines (dro- sopterins) in the orange spots covaries with the carotenoid content, such that the ratio of the two types of pigments is roughly conserved across streams. Carotenoids and drosopterins
Proceedings of The National Academy of Sciences, 2006
Many animals advertise their chemical defense to predators with conspicuous coloration and unpala... more Many animals advertise their chemical defense to predators with conspicuous coloration and unpalatability, but little is known about the information in these signal elements. To effectively avoid predation, is it more advantageous to invest in increased conspicuousness or greater noxiousness, or to allocate equally to both signal modalities? Using natural variation among poison frog species measured with spectral reflectance and
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Papers by Molly Cummings