The evolution of flight and echolocation in bats: another leap in the dark

The earliest known complete bats, from the Eocene (49–53 Mya), were already capable of flapping flight and echolocation. In the absence of direct fossil evidence there have been many speculative scenarios advanced to explain the evolution of these behaviours and their distributions in extant bats. Theories assuming chiropteran monophyly have generally presumed the ancestral pre‐bat was nocturnal, arboreal and insectivorous. Following this assumption hypotheses can be divided into the echolocation first, flight first and tandem development hypotheses, all of which assume that flight evolved only once in the lineage. In contrast, the chiropteran diphyly hypothesis suggests that flight evolved twice. Evidence supporting and refuting the different hypotheses are reviewed. It is concluded that there are significant problems attached to all the current models. A novel hypothesis is advanced, which starts from the assumption that bats are monophyletic and the ancestral pre‐bat was arboreal, but diurnal and frugivorous. After the evolution of flight it is suggested that these animals were driven into the nocturnal niche by the evolution of raptorial birds, and different groups evolved either specialised nocturnal vision (megachiropterans) or echolocation (microchiropterans). A block on sensory modality transfer has retained this distribution of perceptual capabilities ever since, despite some Megachiroptera evolving rudimentary echolocation, and the dietary convergence of some Microchiroptera with the Megachiroptera. The new hypothesis overcomes many of the problems identified in previous treatments.     

Intra-individual variation in the vocalized frequency of the Taiwanese leaf-nosed bat, Hipposideros terasensis, influenced by conspecific colony members

We examined the intra-individual variation in resting frequency of the constant-frequency component of the second harmonic of the pulse (F _rest) over 4 years in a laboratory colony of the Taiwanese leaf-nosed bat (Hipposideros terasensis). Patterns of change in F _rest were observed when individuals were added to or removed from the colony so that we investigated whether F _rest was affected by neighboring colony members. F _rest of each bat continually showed a long-term gradual change throughout the year, and all bats in the colony increased or decreased their F _rest in the same direction as a group non-seasonally. The greatest short-term changes were observed when new bats with a relatively low F _rest joined the colony and F _rest of new bats converged with those of the original colony members around 8 –16 days after their introduction. Conversely, a single individual showed sudden short-term decrease in F _rest after its isolation from other colony members. These findings strongly indicate that F _rest is flexible according to the presence of neighboring conspecific bats. We suggest that the audio-vocal feedback for conspecific pulses appears to be involved in the short- or long-term intra-individual variation in F _rest other than factors previously thought such as age or season.     

Analysis of asymmetries in the African fruit bats Eidolon helvum and Rousettus egyptiacus (Mammalia: Megachiroptera) from the islands of the Gulf of Guinea. II. Integration and levels of multivariate fluctuating asymmetry across a geographical range

Using a set of cranial morphometric characters, trends of variation in multivariate fluctuating asymmetry were evaluated and compared in populations of African fruit bats Rousettus egyptiacus and Eidolon helvum from the Gulf of Guinea islands, and the adjacent mainland. Levels of asymmetry were compared across populations and species, and significant differences were found in both comparisons. Differences coincided with species‐specific patterns of morphological and genetic differentiation. Concordance of correlation matrices of asymmetry was also compared. Results were significant; concordance is hypothesized to be a by‐product of developmental processes that produce the ‘fox‐like’ morphology shared by these species. Consistency of asymmetry patterns suggests that the developmental pathway producing it is highly canalized. A prediction of the above hypothesis is that a radical change in the ‘fox‐like’ structural pattern would result in breakage of the asymmetry parameter associated with it.     

Variation in female mate choice within guppy populations: population divergence,multiple ornaments and the maintenance of polymorphism

The evolutionary significance of variation in mate choice behaviour is currently a subject of some debate and considerable empirical study. Here, I review recent work on variation within and among guppy (Poecilia reticulata) populations in female mate choice and mating preferences. Empirical results demonstrate that there is substantial variation within and among populations in female responsiveness and choosiness, and much of this variation is genetic. Evidence for variation in preference functions also exists, but this appears to be more equivocal and the relative importance of genetic variation is less clear cut. In the second half of this review I discuss the potential significance of this variation to three important evolutionary issues: the presence of multiple male ornaments, the maintenance of polymorphism and divergence in mate recognition among populations. Studies of genetic variation in mate choice within populations indicate that females have complex, multivariate preferences that are able to evolve independently to some extent. These findings suggest that the presence of multiple male ornaments may be due to multiple female mating preferences. The extreme polymorphism in male guppy colour patterns demands explanation, yet no single satisfactory explanation has yet emerged. I review several old ideas and a few new ones in order to identify the most promising potential explanations for future empirical testing. Among these are negative frequency dependent selection, environmental heterogeneity coupled with gene flow, and genetic constraints. Last, I review the relative extent of within and among-population variation in mate choice and mating preferences in order to assess why guppies have not speciated despite a history of isolation and divergence. I argue that variation within guppy populations in mate choice and enhanced mating success of new immigrants to a pool are major impediments to population divergence of the magnitude that would be required for speciation to occur.