Phylogenetic systematics of the viviparous halfbeak genera Dermogenys and Nomorhamphus (Teleostei:Hemiramphidae:Zenarchopterinae)

Among the 13 genera and over 100 species of halfbeaks, three genera - Dermogenys, Nomorhamphus and Hemirhamphodon – are internally fertilized and viviparous. These genera belong to a more inclusive clade, the Zenarchopterinae, that also includes Zenarchopterus , inferred to be internally fertilized and to lay fertilized eggs, and the monotypic Tondanichthys , also inferred to be internally fertilized. Whereas the Hemiramphidae are distributed worldwide, internally fertilized halfbeaks are restricted to Southeast Asia. Recent data from histological surveys of the gonads of both males and females as well as embryonic modifications associated with viviparity have been combined here with osteological characters in a phylogenetic analysis. Results indicate overwhelming support for a sister-group relationship between Hemirhamphodon and {Dermogenys + Nomorhamphus). Monophyly of the Dermogenys + Nomorhamphus clade is also well supported. These results confirm earlier suggestions that Dermogenys , as previously defined, is paraphyletic. Within the Dermogenys + Nomorhamphus clade, two monophyletic clades are supported:one comprises ten species including four new species (Dermogenys bruneiensis, Dermogenys robertsi, Dermogenys palawanensis and Dermogenys collettei) and the other comprises 13 species including three undescribed species (Nomorhamphus rossi, Nomorhamphus pinnimaculata and Nomorhamphus manifesta). Diagnoses for the species of Dermogenys and Nomorhamphus , as well as a natural classification for the included species, are presented.     

Factors shaping the evolution of colour patterns in Australian agamid lizards (Agamidae): a comparative study

Identifying general patterns of adaptive coloration in animals can help to elucidate the evolutionary processes that generate them. We examined the evolution of colour patterns in Australian agamid lizards, a morphologically and ecologically diverse group that relies primarily on visual communication. We tested whether certain types of colour (yellow–reds and black) were likely to be used as sexual signals, as indicated by their association with indices of sexual selection, namely, sexual dichromatism and sexual dimorphism in body size and head shape. We then tested whether sexually dichromatic colours are associated with specific patterns (uniform, mottled, striped, blotched, reticulated) or ecological variables such as habitat openness, arboreality, and substrate type. The presence of yellow–red on lateral and ventral body regions and black on ventral body regions was significantly more common in males than females. Lateral yellow–red in males was associated with the total extent of sexual dichromatism and size dimorphism, whereas ventral yellow–red was associated with sexual dichromatism. Both lateral and ventral yellow–red were associated with uniform patterning, suggesting that sexual signals in male agamid lizards may often comprise uniform patches or flushes of yellow–red. Although ventral black coloration was more prevalent on males (i.e. strongly sexually dichromatic), it was not associated with indices of sexual selection, suggesting that, in agamid lizards, yellow–red coloration is more likely to be sexually selected than black. Sexually dichromatic coloration was not strongly associated with any of the ecological variables measured. We found some associations, however, between female dorsal patterns and ecological variables, suggesting that female patterns are influenced by natural selection. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 101–112.     

Fertilization mode, sperm characteristics, mate choice and parental care patterns in Artedius spp. (Cottidae)

The reproductive biology of three species in the genus Artedius ( A. harringtoni, A. lateralis and A. fenestralis ) was studied, including parental behaviour, mate choice, fertilization mode, gamete morphology and behaviour, and gonad morphology to help understand the coevolution of reproductive characteristics with potentially different modes of fertilization. In all three species, males guard multiple clutches at oviposition sites. In laboratory mate‐choice experiments, males appearred to prefer to defend sites containing eggs and were better able to attract mates at these sites. There was evidence that internal gomete association, although more pronounced in A. harringtoni , existed in all three species. Sperm behaviour and morphology and histological analysis of gonads suggested that A. fenestralis and A. lateralis were more likely to engage in typical external fertilization. The adoption of eggs and their effect on female mate‐choice patterns appearred to reduce the importance of certainty of paternity in these species, and probably increased the evolutionary stability of male parental care.     

Migration and the evolution of duetting in songbirds

Many groups of animals defend shared resources with coordinated signals. The best-studied of these signals are the vocal duets produced by mated pairs of birds. Duets are believed to be more common among tropical-breeding species, but a comprehensive test of this hypothesis is lacking, and the mechanisms that generate latitudinal patterns in duetting are not known. We used a stratified sample of 372 songbird species to conduct the first broad-scale, phylogenetically explicit analysis of duet evolution. We found that duetting evolves in association with the absence of migration, but not with sexual monochromatism or tropical breeding. We conclude that the evolution of migration exerts a major influence on the evolution of duetting. The perceived association between tropical breeding and duetting may be a by-product of the migration–duetting relationship. Migration reduces the average duration of partnerships, potentially reducing the benefits of cooperative behaviour, including duetting. Ultimately, the evolution of coordinated resource-defence signals in songbirds may be driven by ecological conditions that favour sedentary lifestyles and social stability.     

The evolution of reproductive and genomic diversity in ray-finned fishes: insights from phylogeny and comparative analysis

Collectively, ray-finned fishes (Actinopterygii) display far more diversity in many reproductive and genomic features than any other major vertebrate group. Recent large-scale comparative phylogenetic analyses have begun to reveal the evolutionary patterns and putative causes for much of this diversity. Several such recent studies have offered clues to how different reproductive syndromes evolved in these fishes, as well as possible physiological and genomic triggers. In many cases, repeated independent origins of complex reproductive strategies have been uncovered, probably reflecting convergent selection operating on common suites of underlying genes and hormonal controls. For example, phylogenetic analyses have uncovered multiple origins and predominant transitional pathways in the evolution of alternative male reproductive tactics, modes of parental care and mechanisms of sex determination. They have also shown that sexual selection in these fishes is repeatedly associated with particular reproductive strategies. Collectively, studies on reproductive and genomic diversity across the Actinopterygii illustrate both the strengths and the limitations of comparative phylogenetic approaches on large taxonomic scales.     

A Middle Triassic thoracopterid from China highlights the evolutionary origin of overwater gliding in early ray-finned fishes

Gliding adaptations in thoracopterid flying fishes represent a remarkable case of convergent evolution of overwater gliding strategy with modern exocoetid flying fishes, but the evolutionary origin of this strategy was poorly known in the thoracopterids because of lack of transitional forms. Until recently, all thoracopterids, from the Late Triassic of Austria and Italy and the Middle Triassic of South China, were highly specialized ‘four-winged’ gliders in having wing-like paired fins and an asymmetrical caudal fin with the lower caudal lobe notably larger than the upper lobe. Here, we show that the new genus Wushaichthys and the previously alleged ‘peltopleurid’ Peripeltopleurus, from the Middle Triassic (Ladinian, 235–242 Ma) of South China and near the Ladinian/Anisian boundary of southern Switzerland and northern Italy, respectively, represent the most primitive and oldest known thoracopterids. Wushaichthys, the most basal thoracopterid, shows certain derived features of this group in the skull. Peripeltopleurus shows a condition intermediate between Wushaichthys and Thoracopterus in having a slightly asymmetrical caudal fin but still lacking wing-like paired fins. Phylogenetic studies suggest that the evolution of overwater gliding of thoracopterids was gradual in nature; a four-stage adaption following the ‘cranial specialization–asymmetrical caudal fin–enlarged paired fins–scale reduction’ sequence has been recognized in thoracopterid evolution. Moreover, Wushaichthys and Peripeltopleurus bear hooklets on the anal fin of supposed males, resembling those of modern viviparious teleosts. Early thoracopterids probably had evolved a live-bearing reproductive strategy.     

Ecological correlates in the evolution of moult strategies in Western Palearctic passerines

Moult is an important process in the life cycle of birds. Passerines differ widely in the number, seasonality and extension of moult episodes, but the incidence of birds ecology on this variation remains largely uninvestigated. We analysed the patterns of moult in European passerines in relation to their distribution, migration and sexual dichromatism. Longer migrations and southern wintering quarters were characteristic of species with complete moults in summer and an additional moult in winter. The main moult in species with larger seasonal changes in sexual dimorphism tended to be scheduled just before the start of the breeding season, suggesting a link between sexual selection and the timing of moult. These patterns strongly support the importance of migration and dichromatism on the evolution of moult strategies.     

Supertree analyses of the roles of viviparity and habitat in the evolution of atherinomorph fishes

Using supertree phylogenetic reconstructions, we investigate how livebearing and freshwater adaptations may have shaped evolutionary patterns in the Atherinomorpha, a large clade (approximately 1500 extant species) of ray-finned fishes. Based on maximum parsimony reconstructions, livebearing appears to have evolved at least four times independently in this group, and no reversions to the ancestral state of oviparity were evident. With respect to habitat, at least five evolutionary transitions apparently occurred from freshwater to marine environments, at least two transitions in the opposite direction, and no clear ancestral state was identifiable. All viviparous clades exhibited more extant species than their oviparous sister taxa, suggesting that transitions to viviparity may be associated with cladogenetic diversification. Transitions to freshwater were usually, but not invariably associated with increased species richness, but the trend was, overall, not significant among sister clades. Additionally, we investigated whether livebearing and freshwater adaptations are currently associated with elevated risks of extinction as implied by species' presence on the 2004 IUCN Red List. Despite being correlated with decreased brood size, livebearing has not significantly increased extinction risk in the Atherinomorpha. However, freshwater species were significantly more likely than marine species to be listed as endangered.     

Individual variation and the resolution of conflict over parental care in penduline tits

Eurasian penduline tits (Remiz pendulinus) have an unusually diverse breeding system consisting of frequent male and female polygamy, and uniparental care by the male or the female. Intriguingly, 30 to 40 per cent of all nests are deserted by both parents. To understand the evolution of this diverse breeding system and frequent clutch desertion, we use 6 years of field data to derive fitness expectations for males and females depending on whether or not they care for their offspring. The resulting payoff matrix corresponds to an asymmetric Snowdrift Game with two alternative evolutionarily stable strategies (ESSs): female-only and male-only care. This, however, does not explain the polymorphism in care strategies and frequent biparental desertion, because theory predicts that one of the two ESSs should have spread to fixation. Using a bootstrapping approach, we demonstrate that taking account of individual variation in payoffs explains the patterns of care better than a model based on the average population payoff matrix. In particular, a model incorporating differences in male attractiveness closely predicts the observed frequencies of male and female desertion. Our work highlights the need for a new generation of individual-based evolutionary game-theoretic models.     

Sexual conflict over parental care promotes the evolution of sex differences in care and the ability to care

Strong asymmetries in parental care, with one sex providing more care than the other, are widespread across the animal kingdom. At present, two factors are thought to ultimately cause sex differences in care: certainty of parentage and sexual selection. By contrast, we here show that the coevolution of care and the ability to care can result in strong asymmetries in both the ability to care and the level of care, even in the absence of these factors. While the coevolution of care and the ability to care does not predict which sex evolves to care more than the other, once other factors give rise to even the slightest differences in the cost and benefits of care between the sexes (e.g. differences in certainty in parentage), a clear directionality emerges; the sex with the lower cost or higher benefit of care evolves both to be more able to care and to provide much higher levels of care than the other sex. Our findings suggest that the coevolution of levels of care and the ability to care may be a key factor underlying the evolution of sex differences in care.     

Evolution of alternative sex-determining mechanisms in teleost fishes

We compiled information from the literature on the taxonomic distributions in extant teleost fishes of alternative sex-determination systems: male-heterogametic (XY) gonochorism, female-heterogametic (ZW) gonochorism, hermaphroditism, unisexuality, and environmental dependency. Then, using recently published molecular phylogenies based on whole-genomic or partial mitochondrial DNA sequences, we inferred the histories and evolutionary transitions between these reproductive modes by employing maximum parsimony and maximum likelihood methods of phylogenetic character mapping. Across a broad teleost phylogeny involving 25 taxonomic orders, a highly patchy distribution of different sex-determination mechanisms was uncovered, implying numerous transitions between alternative modes, but this heterogeneity also precluded definitive statements about ancestral states for most clades. Closer inspection of family-level and genus-level phylogenies within each of four orders further bolstered the conclusion that shifts in sex-determining modes are evolutionarily frequent and involve a variety of distinct ancestral-descendant pathways. For possible reasons discussed herein, the evolutionary lability of sex-determining modes in fishes contrasts strikingly with the evolutionary conservatism of sex determination within both mammals and birds. © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 83–93.     

Effects of parental care on resource allocation into immune defense and buccal microbiota in mouthbrooding cichlid fishes*

Sexual dimorphism is founded upon a resource allocation trade‐off between investments in reproduction versus other life‐history traits including the immune system. In species with conventional parental care roles, theory predicts that males maximize their lifetime reproductive success by allocating resources toward sexual selection, while females achieve this through prolonging their lifespan. Here, we examine the interrelation between sexual dimorphism and parental care strategies in closely related maternal and biparental mouthbrooding cichlid fishes from East African Lake Tanganyika. We measured cellular immune parameters, examined the relative expression of 28 immune system and life history‐related candidate genes and analyzed the microbiota composition in the buccal cavity. According to our predictions, maternal mouthbrooders are more sexually dimorphic in immune parameters than biparental mouthbrooders, which has possibly arisen through a differential resource allocation into parental care versus secondary sexual traits. Biparental mouthbrooders, on the other hand, which share the costs of parental care, feature an upregulated adaptive immune response and stronger antiviral properties, while their inflammation response is reduced. Overall, our results suggest a differential resource allocation trade‐off between the two modes of parental investment.     

Sexual size dimorphism is not associated with the evolution of parental care in frogs

Sex differences in parental care are thought to arise from differential selection on the sexes. Sexual dimorphism, including sexual size dimorphism (SSD), is often used as a proxy for sexual selection on males. Some studies have found an association between male‐biased SSD (i.e., males larger than females) and the loss of paternal care. While the relationship between sexual selection on males and parental care evolution has been studied extensively, the relationship between female‐biased SSD (i.e., females larger than males) and the evolution of parental care has received very little attention. Thus, we have little knowledge of whether female‐biased SSD coevolves with parental care. In species displaying female‐biased SSD, we might expect dimorphism to be associated with the evolution of paternal care or perhaps the loss of maternal care. Here, drawing on data for 99 extant frog species, we use comparative methods to evaluate how parental care and female‐biased SSD have evolved over time. Generally, we find no significant correlation between the evolution of parental care and female‐biased SSD in frogs. This suggests that differential selection on body size between the sexes is unlikely to have driven the evolution of parental care in these clades and questions whether we should expect sexual dimorphism to exhibit a general relationship with the evolution of sex differences in parental care.     

Phylogeny and evolution of dorsal pattern in the Mexican endemic lizard genus Barisia (Anguidae: Gerrhonotinae)

The phylogeny of the Mexican lizard genus Barisia was assessed using an 878 bp fragment of the mtDNA ND4 gene and a section of associated tRNA genes, as well as 16 external morphological characters. The terminal taxa comprised the currently recognized members of Barisia, including the four subspecies of the polytypic Barisia imbricata and individuals from different populations of the widespread B. i. imbricata and Barisia i. ciliaris, although for Barisia levicollis only morphology could be examined. The ‘step‐matrix frequency’ and the ‘step‐matrix gap‐weighting’ coding approaches were employed simultaneously for the morphological data set, and three different scaling methods were evaluated for the last approach. Maximum parsimony (MP) analyses were performed for the separate and combined data sets and Bayesian analysis was also performed for the mtDNA sequence data. The hypothesis obtained from the simultaneous MP analysis strongly supports the monophyly of Barisia, but the ‘exclusivity’ of B. imbricata as well as of B. i. imbricata and B. i. ciliaris were not recovered. Moreover, inclusion of the morphological data showed B. levicollis nested within a clade together with the taxa assigned to B. i. ciliaris. This, together with the genetic distances and geographic concordance among the haplotypes examined, confirms that B. imbricata represents several species, although the actual species limits in this composite taxon are still unclear. Applying previously published rates of molecular evolution to the mtDNA data gives ages of divergence similar to the times proposed for some Pleistocene–Miocene geological and climatic phenomena that occurred in the Mexican territory. Variation of the dorsal pattern within Barisia was mapped onto the simultaneous morphological and molecular phylogeny, indicating that the two main states present in the taxa assigned to B. imbricata, an adult dorsal pattern present in females and absent in males and the absence of any pattern in both sexes, occur each in separate lineages. This suggests a possible scenario, where sexual dichromatism within Barisia has been repeatedly lost in different lineages.     

Phylogenetic perspectives on diversification and character evolution in the species‐rich genus Erysimum (Erysimeae; Brassicaceae) based on a densely sampled ITS approach

Erysimum includes 150–350 species distributed in the Northern Hemisphere, with Eurasia being the centre of greatest diversity. It is well known for its taxonomic complexity as a result of overlapping morphological characters. We present the first densely sampled phylogenetic analysis of Erysimum using internal transcribed spacer (ITS) DNA sequences from c. 85% of the species (117 for the first time), representing the full range of morphological variation and geographical distribution. We used several approaches to reconstruct phylogenetic relationships, dating of diversification and patterns of evolution of morphological characters in the genus. Ancestral‐state reconstructions of four morphological diagnostic characters were performed using maximum parsimony, maximum likelihood and Bayesian methods. Our phylogenetic framework strongly supports the monophyly of Erysimum and recovers some well‐supported clades that are geographically, rather than morphologically, correlated. Our study confirms the placement of Erysimum in lineage I and reveals two Malcolmia spp. (M. maritima and M. orsiniana) as its sister taxa. The results suggest that the biennial duration and caespitose habit (vs. annual or perennial duration and herbaceous or woody habit) and large, yellow, glabrous (vs. small, non‐yellow, pubescent) petals are ancestral in Erysimum. The ancestral‐state reconstruction results show that annual vs. perennial and woody vs. herbaceous features have been independently derived several times. The dating analyses suggest an early radiation of Erysimum during the late Pliocene or early Pleistocene. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 497–522.     

Evolution of egg dummies in Tanganyikan cichlid fishes: the roles of parental care and sexual selection

Sexual selection has been suggested to be an important driver of speciation in cichlid fishes of the Great Lakes of Africa, and the presence of male egg dummies is proposed to have played a key role. Here, we investigate how mouthbrooding and egg dummies have evolved in Tanganyikan cichlids, the lineage which seeded the other African radiations, with a special emphasis on the egg dummies. Using modern phylogenetic comparative analyses and a phylogeny including 86% of the 200 described species, we provide formal evidence demonstrating correlated evolution between mouthbrooding and egg dummies in Tanganyikan cichlids. These results concur with existing evidence, suggesting that egg dummies have evolved through sensory exploitation. We also demonstrate that there is a strong evolutionary correlation between the presence of egg dummies and both pre‐ and post‐copulatory sexual selection. Moreover, egg dummy evolution was contingent on the intensity of pre‐ and post‐copulatory sexual selection in Tanganyikan cichlids. In sum, our results provide evidence supporting the hypothesis of egg dummies evolving through sensory exploitation and highlight the role of sexual selection in favouring the evolution and maintenance of this trait.     

Phylogenetic eigenvectors and nonstationarity in the evolution of theropod dinosaur skulls

Despite the long‐standing interest in nonstationarity of both phenotypic evolution and diversification rates, only recently have methods been developed to study this property. Here, we propose a methodological expansion of the phylogenetic signal‐representation (PSR) curve based on phylogenetic eigenvectors to test for nonstationarity. The PSR curve is built by plotting the coefficients of determination R² from phylogenetic eigenvector regression (PVR) models increasing the number of phylogenetic eigenvectors against the accumulated eigenvalues. The PSR curve is linear under a stationary model of trait evolution (i.e. the Brownian motion model). Here we describe the distribution of shifts in the models R² and used a randomization procedure to compare observed and simulated shifts along the PSR curve, which allowed detecting nonstationarity in trait evolution. As an applied example, we show that the main evolutionary pattern of variation in the theropod dinosaur skull was nonstationary, with a significant shift in evolutionary rates in derived oviraptorosaurs, an aberrant group of mostly toothless, crested, birdlike theropods. This result is also supported by a recently proposed Bayesian‐based method (AUTEUR). A significant deviation between Ceratosaurus and Limusaurus terminal branches was also detected. We purport that our new approach is a valuable tool for evolutionary biologists, owing to its simplicity, flexibility and comprehensiveness.     

Ecological constraint and the evolution of sexual dichromatism in darters

It is not known how environmental pressures and sexual selection interact to influence the evolution of extravagant male traits. Sexual and natural selection are often viewed as antagonistic forces shaping the evolution of visual signals, where conspicuousness is favored by sexual selection and crypsis is favored by natural selection. Although typically investigated independently, the interaction between natural and sexual selection remains poorly understood. Here, we investigate whether sexual dichromatism evolves stochastically, independent from, or in concert with habitat use in darters, a species‐rich lineage of North American freshwater fish. We find the evolution of sexual dichromatism is coupled to habitat use in darter species. Comparative analyses reveal that mid‐water darter lineages exhibit a narrow distribution of dichromatism trait space surrounding a low optimum, suggesting a constraint imposed on the evolution of dichromatism, potentially through predator‐mediated selection. Alternatively, the transition to benthic habitats coincides with greater variability in the levels of dichromatism that surround a higher optimum, likely due to relaxation of the predator‐mediated selection and heterogeneous microhabitat dependent selection regimes. These results suggest a complex interaction of sexual selection with potentially two mechanisms of natural selection, predation and sensory drive, that influence the evolution of diverse male nuptial coloration in darters.     

Faithful females receive more help: the extent of male parental care during incubation in relation to extra‐pair paternity in songbirds

Parental care provided by males occurs in a diverse array of animals and there are large differences among species in its extent compared with female care. However, social and ecological factors responsible for interspecific differences in male's share of parental duties remain unclear. Genetic fidelity of females has been long considered important. Theory predicts that females should receive more help from their mates in raising the offspring in species with high genetic fidelity. Using avian incubation behaviour as a model system, we confirmed this prediction. The extent of male's help during incubation increased with decreasing rate of extra‐pair paternity across species (22 species of socially monogamous songbirds from 13 families; male's share of incubation ranged from 6% to 58%), even after accounting for covariates, biases in species selection and intraspecific variability. Moreover, this result was not sensitive to two different phylogenies and branch length estimates. We suggest that our findings support the notion, backed by theory, that genetic fidelity is an important factor in the evolution of male parental care. We offer several behavioural scenarios for the coevolution between male's share of parental duties and the genetic mating system.