Coarse dark patterning functionally constrains adaptive shifts from aposematism to crypsis in strawberry poison frogs.

Ecological specialization often requires tight coevolution of several traits, which may constrain future evolutionary pathways and make species more prone to extinction. Aposematism and crypsis represent two specialized adaptations to avoid predation. We tested whether the combined effects of color and pattern on prey conspicuousness functionally constrain or facilitate shifts between these two adaptations. We combined data from 17 natural populations of strawberry poison frogs, Oophaga pumilio with an experimental approach using digitalized images of frogs and chickens as predators. We show that bright coloration often co-occurs with coarse patterning among the natural populations. Dull green frogs with coarse patterning are rare in nature but in the experiment they were as easily detected as bright red frogs suggesting that this trait combination represents a transient evolutionary state toward aposematism. Hence, a gain of either bright color or coarse patterning leads to conspicuousness, but a transition back to crypsis would be functionally constrained in populations with both bright color and coarse patterning by requiring simultaneous changes in two traits. Thus, populations (or species) signaling aposematism by conspicuous color should be less likely to face an evolutionary dead end and more likely to radiate than populations with both conspicuous color and coarse patterning.

Stimulus salience as an explanation for imperfect mimicry.

The theory of mimicry explains how a mimic species gains advantage by resembling a model species [1-3]. Selection for increased mimic-model similarity should then result in accurate mimicry, yet there are many surprising examples of poor mimicry in the natural world [4-8]. The existence of imperfect mimics remains a major unsolved conundrum. We propose and experimentally test a novel explanation of the phenomenon. We argue that predators perceive prey as having several traits, but that the traits differ in their importance for learning. When predators learn to discriminate prey, high-salience traits overshadow other traits, leaving them under little or no selection for similarity, and allow imperfect mimicry to succeed. We tested this idea experimentally, using blue tits as predators and artificial prey with three prominent traits: color, pattern, and shape. We found that otherwise imperfect color mimics were avoided about as much as perfect mimics, whereas pattern and shape mimics did not gain from their similarity to the model. All traits could separately be perceived and learned by the predators, but the color trait was learned at a higher rate, implying that it had higher salience. We conclude that difference in salience between components of prey appearance is of major importance in explaining imperfect mimicry.

Deflective and intimidating eyespots: a comparative study of eyespot size and position in Junonia butterflies.

Eyespots are conspicuous circular features found on the wings of several lepidopteran insects. Two prominent hypotheses have been put forth explaining their function in an antipredatory role. The deflection hypothesis posits that eyespots enhance survival in direct physical encounters with predators by deflecting attacks away from vital parts of the body, whereas the intimidation hypothesis posits that eyespots are advantageous by scaring away a potential predator before an attack. In the light of these two hypotheses, we investigated the evolution of eyespot size and its interaction with position and number within a phylogenetic context in a group of butterflies belonging to the genus Junonia. We found that larger eyespots tend to be found individually, rather than in serial dispositions. Larger size and conspicuousness make intimidating eyespots more effective, and thus, we suggest that our results support an intimidation function in some species of Junonia with solitary eyespots. Our results also show that smaller eyespots in Junonia are located closer to the wing margin, thus supporting predictions of the deflection hypothesis. The interplay between size, position, and arrangement of eyespots in relation to antipredation and possibly sexual selection, promises to be an interesting field of research in the future. Similarly, further comparative work on the evolution of absolute eyespot size in natural populations of other butterfly groups is needed.

Feature saltation and the evolution of mimicry.

In Batesian mimicry, a harmless prey species imitates the warning coloration of an unpalatable model species. A traditional suggestion is that mimicry evolves in a two-step process, in which a large mutation first achieves approximate similarity to the model, after which smaller changes improve the likeness. However, it is not known which aspects of predator psychology cause the initial mutant to be perceived by predators as being similar to the model, leaving open the question of how the crucial first step of mimicry evolution occurs. Using theoretical evolutionary simulations and reconstruction of examples of mimicry evolution, we show that the evolution of Batesian mimicry can be initiated by a mutation that causes prey to acquire a trait that is used by predators as a feature to categorize potential prey as unsuitable. The theory that species gain entry to mimicry through feature saltation allows us to formulate scenarios of the sequence of events during mimicry evolution and to reconstruct an initial mimetic appearance for important examples of Batesian mimicry. Because feature-based categorization by predators entails a qualitative distinction between nonmimics and passable mimics, the theory can explain the occurrence of imperfect mimicry.

Evolutionary aspects of aggression the importance of sexual selection.

Aggressive behaviors in animals, for example, threat, attack, and defense, are commonly related to competition over resources, competition over mating opportunities, or fights for survival. In this chapter, we focus on aggressive competition over mating opportunities, since this competition explains much of the distribution of weaponry and large body size, but also because this type of competition sheds light on the sex skew in the use of violence in mammals, including humans. Darwin (1871) termed this type of natural selection, where differences in reproductive success are caused by competition over mates, sexual selection. Not all species have a pronounced competition over mates, however. Instead, this aspect of sociality is ultimately determined by ecological factors. In species where competition over mates is rampant, this has evolutionary effects on weaponry and body size such that males commonly bear more vicious weapons and are larger than females. A review of sexual selection in mammals reveals how common aggressive competition over mating opportunities is in this group. Nearly half of all mammal species exhibit male-biased sexual size dimorphism, a pattern that is clearly linked to sexual selection. Sexual selection is also common in primates, where it has left clear historical imprints in body mass differences, in weaponry differences (canines), and also in brain structure differences. However, when comparing humans to our closest living primate relatives, it is clear that the degree of male sexual competition has decreased in the hominid lineage. Nevertheless, our species displays dimorphism, polygyny, and sex-specific use of violence typical of a sexually selected mammal. Understanding the biological background of aggressive behaviors is fundamental to understanding human aggression.

Stature and sexual stature dimorphism in Sweden, from the 10th to the end of the 20th century.

Mean stature in a population has been observed to vary with living conditions. If, and how, this affects sexual dimorphism in stature is not fully understood. We analyzed stature data from Swedish populations from the 10th to the end of the 20th century to investigate if male stature is more plastic than female stature in response to environmental changes. Further, we examined if there, as a consequence of this, exists an allometric relationship between male and female stature that is not caused by genetic factors, coupling greater stature with greater dimorphism. We found no significant change in stature from the 10th century to the 17th century, but a clear increase in both male and female stature during the 20th century, most likely because of improved living conditions. Regression analyses revealed no consistent change in sexual stature dimorphism over time for any of the time periods, including the 20th century. Further, we found no significant allometric relationship between male and female stature, and could consequently not identify any significant relationship between stature and stature dimorphism. Thus, contrary to previous suggestions, the regressions did not provide support for the assertion that male stature is more sensitive to environmental changes than female stature, nor that stature dimorphism increases with increasing stature.

Aposematism and crypsis combined as a result of distance dependence: functional versatility of the colour pattern in the swallowtail butterfly larva.

The idea that an aposematic prey combines crypsis at a distance with conspicuousness close up was tested in an experiment using human subjects. We estimated detectability of the aposematic larva of the swallowtail butterfly, Papilio machaon, in two habitats, by presenting, on a touch screen, photographs taken at four different distances and measuring the time elapsed to discovery. The detectability of larvae in these images was compared with images that were manipulated, using existing colours either to increase or decrease conspicuousness. Detection time increased with distance for all colourations. However, at the closest distance, detection time was longer for the larvae manipulated to be more cryptic than for the natural and more conspicuous forms. This indicates that the natural colouration is not maximally cryptic at a short distance. Further, smaller increments in distance were needed to increase detection time for the natural than for the conspicuous larva. This indicates that the natural colouration is not maximally conspicuous at longer distances. Taken together, we present the first empirical support for the idea that some colour patterns may combine warning colouration at a close range with crypsis at a longer range. The implications of this result for the evolution of aposematism are discussed.

Constrained camouflage facilitates the evolution of conspicuous warning coloration.

The initial evolution of aposematic and mimetic antipredator signals is thought to be paradoxical because such coloration is expected to increase the risk of predation before reaching a stage when predators associate it effectively with a defense. We propose, however, that constraints associated with the alternative strategy, cryptic coloration, may facilitate the evolution of antipredator signals and thus provide a solution for the apparent paradox. We tested this hypothesis first using an evolutionary simulation to study the effect of a constraint due to habitat heterogeneity, and second using a phylogenetic comparison of the Lepidoptera to investigate the effect of a constraint due to prey motility. In the evolutionary simulation, antipredator warning coloration had an increased probability to invade the prey population when the evolution of camouflage was constrained by visual difference between microhabitats. The comparative study was done between day-active lepidopteran taxa, in which camouflage is constrained by motility, and night-active taxa, which rest during the day and are thus able to rely on camouflage. We compared each of seven phylogenetically independent day-active groups with a closely related nocturnal group and found that antipredator signals have evolved at least once in all the diurnal groups but in none of their nocturnal matches. Both studies lend support to our idea that constraints on crypsis may favor the evolution of antipredator warning signals.

Phylogenetic reconstruction of parental-care systems in the ancestors of birds.

Due to the controversy surrounding incipient avian parental care, ancestral parental care systems were reconstructed in a phylogeny including major extant amniote lineages. Using two different resolutions for the basal avian branches, transitions between the states no care, female care, biparental care and male care were inferred for the most basal branches of the tree. Uniparental female care was inferred for the lineage to birds and crocodiles. Using a phylogeny where ratites and tinamous branch off early and an ordered character-state assumption, a transition to biparental care was inferred for the ancestor of birds. This ancestor could be any organism along the lineage leading from the crocodile-bird split up to modern birds, not necessarily the original bird. We discuss the support for alternative avian phylogenies and the homology in parental care between crocodiles and birds. We suggest that the phylogenetic pattern should be used as a starting point for a more detailed analysis of parental care systems in birds and their relatives.

A critique of the naturalistic fallacy thesis.

If the prescriptive "ought" is separated from the factual "is," an intellectual analysis of the real world is by definition without normative value. The naturalistic fallacy thesis -- maintaining that normative and descriptive spheres must remain separated -- is often presented in a weak sense that seems reasonable. However, only in a strong sense -- by strictly separating facts and values -- are fallacy accusations supported. We claim that this naturalistic fallacy thesis is unsound and that normative statements instead should be based on rational understanding as found in the Darwinian and social sciences. The Cartesian compromise should be abandoned, since only naturalism can provide a cogent framework for better understanding and support ethics with a solid foundation. Many people nurture values based on tradition, whim, subgroup identification etc., and they demand respect for those values. However, we can demand respect for values only when they have a rational foundation. The common belief in the thesis of naturalistic fallacy is an anti-intellectual device that shields values from rational inquiry.