J. Jordan Price

The evolution of animal communication

Communication is a central part of social behaviour, often directly affect-ing an individual’s fitness. As such, the study of animal communication hasbecome a central focus of animal behaviour researchers. It is a field that inte-grates a broad and diverse range of biological disciplines, from neurobiologyand biomechanics to evolution and psychology, and over the years it has at-tracted a broad range of researchers as well. The contributors to this SpecialIssue of

RECONSTRUCTING THE EVOLUTION OF COMPLEX BIRD SONG IN THE OROPENDOLAS

Abstract The elaborate songs of songbirds are frequent models for investigating the evolution of animal signals. However, few previous studies have attempted to reconstruct historical changes in song evolution using a phylogenetic comparative approach. In particular, no comparative studies of bird song have used a large number of vocal characters and a well-supported, independently derived phylogeny. We identified 32 features in the complex vocal displays of male oropendolas (genera Psarocolius, Gymnostinops, and Ocyalus) that are relatively invariant within taxa and mapped these characters onto a robust molecular phylogeny of the group. Our analysis revealed that many aspects of oropendola song are surprisingly evolutionarily conservative and thus are potentially useful characters for reconstructing historical patterns. Of the characters that varied among taxa, nearly two thirds (19 of 29) showed no evidence of evolutionary convergence or reversal when mapped onto the tree, which was reflected in a high overall consistency index (CI = 0.78) and retention index (RI = 0.88). Some reconstructed patterns provided evidence of selection on these signals. For example, rapid divergence of the songs of the Montezuma oropendola, Gymnostinops montezuma, from those of closely related taxa suggests the recent influence of strong sexual selection. In general, our results provide insights into the mode of vocal evolution in songbirds and suggest that complex vocalizations can provide information about phylogeny. Based on this evidence, we use song characters to estimate the phylogenetic affinities of three oropendola taxa for which molecular data are not yet available.

Losses of female song with changes from tropical to temperate breeding in the New World blackbirds

Birds in which both sexes produce complex songs are thought to be more common in the tropics than in temperate areas, where typically only males sing. Yet the role of phylogeny in this apparent relationship between female song and latitude has never been examined. Here, we reconstruct evolutionary changes in female song and breeding latitude in the New World blackbirds (Icteridae), a family with both temperate and tropical representatives. We provide strong evidence that members of this group have moved repeatedly from tropical to temperate breeding ranges and, furthermore, that these range shifts were associated with losses of female song more often than expected by chance. This historical perspective suggests that male-biased song production in many temperate species is the result not of sexual selection for complex song in males but of selection against such songs in females. Our results provide new insights into the differences we see today between tropical and temperate songbirds, and suggest that the role of sexual selection in the evolution of bird song might not be as simple as we think.

Recognition of family-specific calls in stripe-backed wrens

Males of the cooperatively breeding stripe-backed wren Campylorhynchus nuchalis, learn repertoires of stereotyped calls (termed WAY calls) from older male relatives. As a result, these vocalizations are normally specific to patrilineal family groups but are sometimes shared by male relatives in different groups. To determine whether or not this species can recognize the calls of different family groups, I performed playback experiments with individual call types recorded from males of known social and genealogical relationships. Subjects discriminated between the calls of unrelated neighbouring groups and unfamiliar groups, and they discriminated both of these from calls of their own groups. However, subjects failed to distinguish calls of males in other groups from calls of their own groups when these males were members of the same patriline. These results indicate that stripe-backed wrens can discriminate between repertoires of these calls that match or differ from their own. Consequently, they can recognize members of their patriline, not just members of their immediate group. These vocalizations probably provide a useful mechanism for recognizing group membership in this species and might also provide a mechanism for recognizing unfamiliar relatives in other groups. Copyright 1999 The Association for the Study of Animal Behaviour.

SONG AND PLUMAGE EVOLUTION IN THE NEW WORLD ORIOLES (ICTERUS) SHOW SIMILAR LABILITY AND CONVERGENCE IN PATTERNS

Abstract Both song and color patterns in birds are thought to evolve rapidly and exhibit high levels of homoplasy, yet few previous studies have compared the evolution of these traits systematically using the same taxa. Here we reconstruct the evolution of song in the New World orioles (Icterus) and compare patterns of vocal evolution to previously reconstructed patterns of change in plumage evolution in this clade. Individual vocal characters exhibit high levels of homoplasy, reflected in a low overall consistency index (CI = 0.27) and retention index (RI = 0.35). Levels of lability in song are comparable to those found for oriole plumage patterns using the same taxa (CI = 0.31, RI = 0.63), but are strikingly dissimilar to the conservative patterns of change seen in the songs of oropendolas (Psarocolius, Ocyalus; CI = 0.82, RI = 0.87), a group closely related to the orioles. Oriole song is also similar to oriole plumage in exhibiting repeated convergence in overall patterns, with some distantly related taxa sounding remarkably similar. Thus, both song and plumage in orioles show repeated convergence in individual elements and in overall patterns across the clade, suggesting that both of these character classes are highly labile between taxa yet highly conserved within the genus. Our results provide new insights into the tempo and mode of evolution in sexually selected traits within and across clades.

RECONSTRUCTING THE EVOLUTION OF SEXUAL DICHROMATISM: CURRENT COLOR DIVERSITY DOES NOT REFLECT PAST RATES OF MALE AND FEMALE CHANGE

Males of sexually dimorphic species often appear more divergent among taxa than do females, so it is often assumed that evolutionary changes have occurred primarily in males. Yet, sexual dimorphisms can result from historical changes in either or both of the sexes, and few previous studies have investigated such patterns using phylogenetic methods. Here, we describe the evolution of male and female plumage colors in the grackles and allies (Icteridae), a songbird clade with a broad range in levels of sexual dichromatism. Using a model of avian perceptual color space, we calculated color distances within and among taxa on a molecular phylogeny. Our results show that female plumage colors have changed more dramatically than male colors in the evolutionary past, yet male colors are significantly more divergent among species today. Historical increases in dichromatism have involved changes in both sexes, whereas decreases in dichromatism have nearly always involved females evolving rapidly to look like males. Dichromatism is also associated with mating system in this group, with monogamous taxa tending to exhibit relatively low levels of sexual dichromatism. Our findings suggest that, despite appearances, female plumage evolution plays a more prominent role in sexual dichromatism than is generally assumed.

Sex-Role Reversal in Song? Females Sing More Frequently Than Males in the Streak-Backed Oriole

Abstract Birds in which both sexes produce complex song are more common in the tropics than in the temperate north, where typically only males sing. Yet surprisingly little is known about female song characteristics in most tropical species. Here we present a comparison of female and male singing behaviors in the Streak-backed Oriole (Icterus pustulatus), a tropical songbird in which both sexes perform solo songs. Females sing much more frequently than males and produce songs with similar acoustic complexity. Rates of singing by both sexes were higher during breeding than postbreeding while the rates of most other vocalizations did not change, suggesting that song plays an important role in breeding. To our knowledge, this is the first reported species in which females regularly sing at higher rates than males; however, few studies have examined female song in other sexually monomorphic or weakly dimorphic species, so such patterns might not be unique.

Plumage Evolution in the Oropendolas and Caciques: Different Divergence Rates in Polygynous and Monogamous Taxa

Avian plumage colors are frequently used in studies of sexual selection, yet surprisingly little is known about how these traits evolve under different mating systems. We compared historical rates of divergence in male color patterns among the oropendolas and caciques (genera Cacicus, Gymnostinops, Ocyalus, and Psarocolius), a group with both polygynous and monogamous representatives. Reconstructing the evolution of individual color patches on a molecular phylogeny showed that overall color patterns have changed much more rapidly in oropendolas, which comprise two groups that evolved polygyny independently, than in caciques, which are predominantly monogamous. None of these taxa are notably sexually dichromatic, however, suggesting that higher rates of plumage evolution occurred in both sexes rather than just males. Despite high rates of change, color patterns show few examples of convergence among taxa, similar to the lack of homoplasy in male song among oropendolas but in a stark contrast to the repeated convergence in both plumage and song patterns found in a closely related, monogamous clade, the New World orioles (Icterus). Our results support previous suggestions that display traits evolve more rapidly and with less homoplasy in polygynous mating systems, and we provide surprising evidence that these patterns may occur in both sexes.

Song evolution in Maluridae: influences of natural and sexual selection on acoustic structure

Abstract Many factors may influence the evolution of acoustic signals, including sexual selection, morphological constraints and environmental variation. These factors can play simultaneous and interacting roles in determining signal phenotypes. Here, we assess the evolution of song features in the Maluridae, a passerine family with significant variation among taxa in levels of sperm competition, morphological features and breeding habitats ranging from arid grasslands in Australia to tropical rainforests in New Guinea. We used phylogenetic comparative methods and a robust molecular phylogeny to compare song characteristics with a variety of other measures, including testes mass, body-size and latitude. Several aspects of the temporal and frequency structure of song were associated with relative testes mass, suggesting that sexual selection may influence some song characteristics in this family. The lowest frequencies of song were strongly predicted by body-size, indicating that morphological constraints have also likely influenced acoustic phenotypes. Song versatility, reflecting the diversity of note types in a song, was positively correlated with latitude, suggesting that complexity may increase in association with more temperate or variable environments. Variation in song structure across the family appears to reflect a complex interaction between natural and sexual selection.

A ROBUST PHYLOGENY OF THE OROPENDOLAS: POLYPHYLY REVEALED BY MITOCHONDRIAL SEQUENCE DATA

Abstract We present a robust, fully resolved phylogeny for the oropendolas that will serve as a basis for comparative studies in this group. We sequenced 2,011 base pairs (bp) of the mitochondrial cytochrome-b and ND2 genes from 22 individuals to reconstruct relationships between recognized species and subspecies and to assess variation within polytypic taxa. A single phylogenetic tree was produced despite the use of a wide range of weighting schemes and phylogenetic reconstruction methods. Our data provide strong evidence that oropendolas are polyphyletic, with two distinct groups within a larger clade of oropendolas and caciques. We confirm the monophyly of recognized species, but indicate that some within-species relationships do not conform to recognized subspecies limits. Our findings thus demonstrate the importance of including multiple exemplars from each taxon of interest. The two genes provided complimentary and equally effective phylogenetic information for comparisons within the oropendolas, but exhibited lower resolution in comparisons above the species level.