External and internal sexual dimorphism in leiognathid fishes: morphological evidence for sex-specific bioluminescent signaling

Fourteen species of leiognathid fishes (Perciformes, Leiognathidae) from the Philippine Islands, Thailand, Japan, Indonesia, and Palau were examined for accessory secondary sexual dimorphism. Thirteen species exhibit either external dimorphism (a clear patch of skin on the flanks of males, a large clear patch of skin on the opercular margins of males, or a flank stripe in males) or internal dimorphism (large light organs in males) or both. Eight of the 14 species (and possibly as many as 11) exhibit both forms of sexual dimorphism. Two species show only internal light organ volume dimorphism, and one species shows neither external nor internal dimorphism. Sexual dimorphism is thus very common in leiognathids. The externally dimorphic skin patches are closely associated with the internally dimorphic light organ system in seven species (and possibly as many as ten), indicating a potential for light emission through the clear patches. A bioluminescent signaling function by males is therefore suggested for the sexual dimorphism in leiognathids, which may play an important role in the schooling behavior as well as in species and sexual recognition of these coastal fishes.     

Evolution of the light organ system in ponyfishes (Teleostei: Leiognathidae)

Members of the leiognathid subfamily Gazzinae, comprising approximately two‐thirds of ponyfish species, are sexually dimorphic with regard to features of the light organ system (LOS). In Gazzinae, the circumesophageal light organ (LO) of males is enlarged and varies in shape compared with similarly sized conspecific females. In association with male species‐specific translucent external patches on the head and flank, these sexually dimorphic LO features are hypothesized to be correlated with species‐specific luminescence displays. Anatomical differences in LO shape, volume, and orientation, and its association with the gas bladder and other internal structures that function in light emission, are compared to observations of luminescence displays for every major lineage within Leiognathidae. We reconstruct the character evolution of both internal and external morphological features of the LOS to investigate the evolution of LO sexual dimorphism and morphology. Both internal and external sexual dimorphism in the ponyfish LOs were recovered as most likely to have evolved in the common ancestor of Leiognathidae, and likelihood‐based correlation analyses indicate that the evolution of internal and external dimorphism in males is statistically correlated. Magnetic resonance imaging technology was applied to examine the unique internal LOs of ponyfishes in situ, which provides a new metric (LO index) for comparison of LO structure across lineages. J. Morphol. 2011. © 2011 Wiley‐Liss, Inc.     

A molecular phylogeny of the Cephinae (Hymenoptera, Cephidae) based on mtDNA COI gene: a test of traditional classification

Cephinae is traditionally divided into three tribes and about 24 genera based on morphology and host utilization. There has been no study testing the monophyly of taxa under a strict phylogenetic criterion. A molecular phylogeny of Cephinae based on a total of 68 sequences of mtDNA COI gene, representing seven genera of Cephinae, is reconstructed to test the traditional limits and relationships of taxa. Monophyly of the traditional tribes is not supported. Monophyly of the genera are largely supported except for Pachycephus. A few host shift events are suggested based on phylogenetic relationships among taxa. These results indicate that a more robust phylogeny is required for a more plausible conclusion. We also report two species of Cephus for the first time from Turkey.     

Molecular phylogeny and systematics of the pseudoxyrhophiine snake genus Liopholidophis (Reptilia, Colubridae): evolution of its exceptional sexual dimorphism and descriptions of new taxa

The pseudoxyrhophiine snake genus Liopholidophis Mocquard, 1904 is endemic to Madagascar and consists of two distinct species groups. We performed molecular phylogenetic analyses using nucleotide sequences of c . 2200 bp from two mitochondrial genes (16S rRNA, cyt b ) and one nuclear gene (c-mos) to test the monophyly of the genus Liopholidophis and to investigate the relationships of the known species and an unidentified Liopholidophis population. Our phylogeny strongly supports the polyphyly of the genus Liopholidophis , confirms the monophyly of both species groups, and reveals that the unidentified population belongs to a new species. We therefore transfer the species of the former L. stumpffi group ( L. epistibes , L. infrasignatus , L. lateralis , L. martae , and L. stumpffi ) to a new genus Bibilava gen. n. and describe Liopholidophis dimorphus sp. n. from the rainforests of Montagne d'Ambre National Park in northern Madagascar. Finally, we compare our phylogeny with previous hypotheses and discuss the evolution of the extreme sexual dimorphism in tail length of Liopholidophis s. str.     

Phylogeny of Japanese stag beetles (Coleoptera: Lucanidae) inferred from 16S mtrRNA gene sequences, with reference to the evolution of sexual dimorphism of mandibles

As a first step in reconstructing the phylogeny of world stag beetles (Coleoptera: Lucanidae), phylogenetic relationships among the major members of Japanese stag beetles were explored by analyzing a sequence of 1030 nucleotides from the mitochondrial 16S ribosomal RNA (16S rRNA) gene. A total of 20 species and three additional subspecies representing 13 genera were examined to provide basic information on the phylogeny of world Lucanidae. The resultant phylogenetic tree indicates that the family Lucanidae is monophyletic, and contains two major lineages: one consists of the genera Platycerus, Aesalus, Ceruchus, and Nicagus, and the other includes Dorcus, Rhaetulus, Prosopocoilus, Aegus, Neolucanus, Prismognathus, Lucanus, Figulus, and Nigidius. Generic members of the latter lineage are further divided into the following four sublineages: i) Figulus and Nigidius; ii) Prismognathus and Lucanus; iii) Aegus and Neolucanus; and iv) Dorcus, Rhaetulus, and Prosopocoilus. These molecular phylogenetic relationships are used as a basis for a preliminary exploration of the evolution of sexual dimorphism in the shape of the mandible. The results of this investigation suggest that strong sexual dimorphism with well-developed mandibles in males evolved independently at least twice, once in the genus Aegus and once in the ancestor of the Lucanus-Prismognathus and Dorcus-Rhaetulus-Prosopocoilus clades. Alternatively, it is possible that sexual dimorphism of mandibles has undergone secondary loss in the genera Figulus and Nigidius.     

Phylogenetic analyses of the genera Pipra, Lepidothrix and Dixiphia (Pipridae, Passeriformes) using partial cytochrome b and 16S mtDNA genes

The piprids (manakins) are small-bodied Neotropical birds characterized by sexual dimorphism of the plumage and elaborate courtship rituals. The phylogenetic relationships of some of piprid genera, in particular Pipra , are not well defined. Recently, Pipra was divided into three distinct groups, and the recognition of two new genera — Lepidothrix and Dixiphia — was suggested, based on the characteristic of the syringial morphology. In the present study, we analyse the phylogenetic relationships among these genera on the basis of data from the mitochondrial genes rRNA 16S and cytochrome b . Bayesian Inference (BI), Maximum Parsimony (MP) and Maximum Likelihood (ML) methods of analyses produced phylogenetic trees with very similar topologies with regard to the groupings formed by the species representing the three genera. These results indicate that the genera Pipra, Lepidothrix and Dixiphia do not constitute a single monophyletic clade, and support the current hypothesis of polyphyletism for the former representatives of the genus Pipra . However, the topologies presented do not coincide exactly with the phylogeny suggested by the morphological data with regard to the relationship between these three groups and the remaining piprid genera analysed here, emphasizing the need for further studies of this group of birds. The agreement between the molecular data presented here and the biogeographical information available for some of the species analysed appears to confirm the potential of using segments of the mitochondrial genome for the study of phylogeographical patterns in this group of organisms.     

The revision of species-rich genera: a phylogenetic framework for the strategic revision of Pilea (Urticaceae) based on cpDNA, nrDNA, and morphology

The revision of species-rich genera underpins research and supports the sustainable use and monitoring of biological diversity. One fifth to one quarter of the diversity of all seed plant species occurs in such genera, but difficulties with the revision of species-rich genera has resulted in many of them being ignored since the late 1800s. Pilea, with 600-715 species is in need of revision. The only realistic approach is in manageable subunits, which requires confirmation of monophyly and identification of monophyletic subdivisions. Parsimony analyses of trnL-F, ITS, and morphology data were used to test the monophyly of, and explore intrageneric relationships within, Pilea. Analysis of trnL-F data confirms and recovers two morphologically diagnosable monophyletic clades that include all of the taxa within Pilea. Overlaying geographic distribution on a most parsimonious tree indicates a strong association between geography and phylogenetic relatedness. It is suggested that a strategic revision within the framework of morphologically and geographically diagnosable units might enable the revision of the group using an iterative approach. Analysis of the outgroup taxa supports the inclusion of Poikilospermum within the Urticaceae and suggests that the Urticaceae tribes could be placed into two clades that are supported by floral morphology.     

Phylogenetic study of heliconiine butterflies based on morphology and restriction analysis of ribosomal RNA genes

Relationships of ten heliconiine butterflies (genera Dryas and Heliconius , family Nymphalidae) were elucidated by phylogenetic analysis of characters based on ribosomal DNA restriction site variation and morphology. Agraulis vanillae , also a heliconiine, was used as the outgroup species. Although neither the morphological nor the molecular data unambiguously resolve relationships among the heliconiines, a combined analysis of both data sets results in a tree that is similar to traditional systematic arrangements and previous views of radiation in the group. Both pupal-mating and nonpupal-mating species group as clades in the combined analysis. However, the restriction site data alone do not support the monophyly of the pupal-mating clade, and the morphological data alone do not support the monophyly of the non-pupal-mating clade. Furthermore, relationships of H. melpomene, H. cydno and the silvaniform species depart from traditional arrangements based on morphology and reproductive compatibility experiments. All trees support the independent evolution of similar wing patterns of species previously suggested to be members of mimicry complexes. Several mimicry complexes appear to have a member in each of the two major monophyletic groups (pupal-mating and non-pupal-mating clades).     

Phylogeny of North American fireflies (Coleoptera: Lampyridae): implications for the evolution of light signals

Representatives of the beetle family Lampyridae ("fireflies", "lightningbugs") are well known for their use of light signals for species recognition during mate search. However, not all species in this family use light for mate attraction, but use chemical signals instead. The lampyrids have a worldwide distribution with more than 2000 described species, but very little is known about their phylogenetic relationships. Within North America, some lampyrids use pheromones as the major mating signal whereas others use visual signals such as extended glows or short light flashes. Here, we use a phylogenetic approach to illuminate the relationships of North American lampyrids and the evolution of their mating signals. Specifically, to establish the first phylogeny of all North American lampyrid genera, we sequenced nuclear (18S) and mitochondrial (16S and COI) genes to investigate the phylogenetic relationships of 26 species from 16 North American (NA) genera and one species from the genus Pterotus that was removed recently from the Lampyridae. To test the monophyly of the NA firefly fauna we sequenced the same genes from three European lampyrids and three Asian lampyrids, and included all available Genbank data (27 additional Asian lampyrids and a former lampyrid from Asia, Rhagophthalmus). Our results show that the North American lampyrids are not monophyletic. Different subgroups are closely related to species from Europe, Asia and tropical America, respectively. The present classification of fireflies into subfamilies and tribes is not, for the most part, supported by our phylogenetic analysis. Two former lampyrid genera, Pterotus and Rhagophthalmus, which have recently been removed from this family, are in fact nested within the Lampyridae. Further, we found that the use of light as a sexual signal may have originated one or four times among lampyrids, followed by nine or four losses, respectively. Short flashes originated at least twice and possibly three times independently among our study taxa. The use of short flashes as a mating signal was replaced at least once by the use of long glows, and light signals as mating signals were lost at least three times in our study group and replaced by pheromones as the main signal mode.     

The specificity of symbiosis: Pony fish and luminescent bacteria

Luminescent bacteria isolated from light organs of seven different species (3 genera) of fishes of the family Leiognathidae were subjected to taxonomic analysis. Of the 733 isolated all but seven were identified as Photobacterium leiognathi; the others are considered to be either chance contaminants of the sampling procedure or transients within the organ. In most fish, the luminous organ appeared to contain a single predominating strain of P. leiognathi with small numbers of one to three other strains of the same species, differing by only one or two characters.     

New insights into the systematics and molecular phylogeny of the Malagasy snake genus Liopholidophis suggest at least one rapid reversal of extreme sexual dimorphism in tail length

The pseudoxyrhophiine snake genus Liopholidophis Mocquard, 1904 (family Lamprophiidae) is endemic to Madagascar and according to its present definition comprises six medium-sized, terrestrial and diurnal snake species, most of which are characterised by an unusual and extreme sexual dimorphism in tail length. We performed molecular phylogenetic analyses using nucleotide sequences of three mitochondrial genes (16S rRNA, cytochrome b and cytochrome oxidase I) and one nuclear gene (c-mos) for all described and two additional species newly described herein. The two new species are very small sized (total length: 234–312.5 mm), have comparatively short tails and a reduced number of dorsal scale rows (15 at midbody), the lowest value among all non-scolecophidian snakes of Madagascar. Both species are secretive or rare, and they have a reddish belly in life that fades in preservative. In terms of colouration and morphology, they are most similar to each other and furthermore to Liopholidophis rhadinaea. Together with this species and L. dimorphus, they form a well-supported clade. Liopholidophis baderi sp. nov. from central eastern Madagascar is characterised by 149–158 ventrals and 71–77 subcaudals, whereas the similar L. oligolepis sp. nov. from the northeast has even fewer ventrals (137) and subcaudals (54). The phylogenetic tree suggests that the tail length dimorphism in the genus Liopholidophis has evolved in a complex pattern including at least one reversal. The phylogenetic position of the two new dwarf species indicates that both the absence of extreme sexual dimorphism in tail length and their body size reduction are derived and probably correlated features. Also the close phylogenetic relationships between the long-tailed L. sexlineatus and the similar but relatively short-tailed L. varius demonstrate that dimorphism in tail length can be strongly mitigated in short evolutionary time periods.     

SPECIES PHYLOGENY OF COSMARIUM AND STAURASTRUM (DESMIDIACEAE) BASED ON RBC L SEQUENCES

Cosmarium and Staurastrum are the two most diverse genera of placoderm desmids (Family Desmidiaceae), with approximately 1100 and 800 species, respectively. Phylogenetic analysis of relationships of species has been extremely difficult. In a monograph of North American placoderm desmids, Prescott et al. described early phylogenetic work that concluded Staurastrum to be polyphyletic and certainly polymorphic. Likewise, Cosmarium has also been viewed as polyphyletic, and a number of workers have proposed splitting these genera. The classical view of West and West grouped species within each genus into two divisions and 6–8 sections based on wall features and semicell shape. We sequenced rbc L from 18 species of Cosmarium (2 divisions, 7 sections) and 12 species of Staurastrum (2 divisions and 7 sections) and performed a phylogenetic analysis (parsimony, maximum likelihood, bootstrap) using other placoderm desmids and Zygnematales as outgroups. The results exhibit little support for the monophyly of sections or divisions of the two genera. Furthermore, although there is support for the monophyly of clades within each genus, there is also support for a separate clade containing species from both genera.     

A molecular phylogeny of Planorboidea (Gastropoda, Pulmonata): insights from enhanced taxon sampling

Planorbid gastropods are the most diverse group of limnic pulmonates, with both discoidal and highspired taxa. Phylogenetic relationships among these genera are confused and controversial. In particular, the monophyly of the limpet‐like taxa (traditionally Ancylidae) is disputed. Even recent molecular studies have concluded that substantially more work is necessary to solve the remaining issues concerning intergeneric phylogenetic relationships and higher taxa systematics. Planorbid snails are of great significance for humans as several members of this group are intermediate hosts of blood flukes (schistosomes) causing a chronic disease, schistosomiasis. We used the two independent molecular markers COI and 18S (concatenated dataset of 2837 nucleotide bp) to infer phylogenetic relationships of 26 genera (27 species) of Planorboidea, represented mostly by type species from mainly topotypical populations. With the majority of the taxa discussed not having been studied previously, this study attempted to test several hypotheses on planorbid phylogenetic relationships using Bayesian inference techniques. The monophyly of Planorboidea (= ‘Ancyloplanorbidae’) is strongly suggested on the basis of our extensive molecular analysis. Besides a distinct Burnupia clade, two major clades were recovered that correspond to family level taxa (traditional Bulinidae and Planorbidae). Considerable rearrangements of suprageneric taxa are evident from the phylogeny inferred. Therefore, the only clades recognized by current classifications and supported by our analysis are Planorbini and Segmentinini. The present study found that Ancylidae as traditionally understood, i.e. covering most freshwater limpet gastropods, is paraphyletic, as the genera of Burnupia and Protancylus have been shown to lie phylogenetically outside the Ancylini. Chromosome numbers and levels of polyploidy are discussed in the light of the new phylogeny. An earlier theory of shell shape evolution, i.e. that of patelliform taxa being most advanced, was not supported by this study; a limpet‐shaped taxon is most basal within Planorboidea. Although many taxa still remain to be studied, our results will hopefully contribute towards a better understanding of this very important group of freshwater organisms. Some taxonomic implications are discussed.     

Phylogenetic evidence for loss of sound production and a shift in sexual recognition signals in Hamadryas butterflies (Nymphalidae: Biblidinae)

The neotropical butterfly genus Hamadryas Hübner comprises 20 species that exhibit an intriguing variation in their natural history traits. Although revised in 1983, no phylogenetic hypothesis was presented: the first phylogenetic hypothesis is estimated here based on 93 characters and including species from the three other genera in the tribe Ageroniini. The phylogeny is used to test the monophyly of the genus, establish the sister group of Hamadryas and identify its apomorphies. The tree allows the inference of patterns of character change in sound production and sexual dimorphism. Implied weights show that Hamadryas is monophyletic and corroborate Ectima Doubleday as a sister genus. Previously suggested subgenera for Hamadryas were non‐monophyletic, with the exception of the laodamia clade, supported by the presence of a complete sterigma. Sound production is inferred to be a derived condition in Hamadryas that has been lost in the laodamia clade. This, plus the presence of androconial organs and sexual dimorphism in the laodamia clade, suggests a shift in sexual recognition signalling. Furthermore, the phylogeny indicates that the colour pattern of males in the laodamia clade is novel, supporting a Darwinian origin of sexual dimorphism.     

Sexual dimorphism in the craniofacial growth of the guinea pig (Cavia porcellus)

Variation between the sexes during ontogeny is frequently overlooked in discussions of the phylogenetic patterns of adult sexual dimorphism. Different growth trajectories can produce identical degrees and direction of adult dimorphism and the possibility exists that similarities in adults may be the result of differing growth patterns, suggesting independent evolutionary pathways among species to the seemingly identical adult morphology. We quantified the sexual dimorphism in craniofacial skeletal growth of Cavia porcellus, the guinea pig, using longitudinally collected radiographs. Guinea pigs have male-biased sexual dimorphism in size and in growth parameters, despite literature reports to the contrary. These results, analyzed with equivalent data for five species of rodents, and two outgroups representing similarly sized mammals, a rabbit and a marsupial, indicate that some aspects of sexual differences in growth follow phylogenetic lines, while others are a function of whether the species has male- or female-biased dimorphism.     

Molecular systematics of the deep-sea hydrothermal vent endemic Brachyuran family Bythograeidae: a comparison of three Bayesian species tree methods

Brachyuran crabs of the family Bythograeidae are endemic to deep-sea hydrothermal vents and represent one of the most successful groups of macroinvertebrates that have colonized this extreme environment. Occurring worldwide, the family includes six genera (Allograea, Austinograea, Bythograea, Cyanagraea, Gandalfus, and Segonzacia) and fourteen formally described species. To investigate their evolutionary relationships, we conducted Maximum Likelihood and Bayesian molecular phylogenetic analyses, based on DNA sequences from fragments of three mitochondrial genes (16S rDNA, Cytochrome oxidase I, and Cytochrome b) and three nuclear genes (28S rDNA, the sodium-potassium ATPase a-subunit 'NaK', and Histone H3A). We employed traditional concatenated (i.e., supermatrix) phylogenetic methods, as well as three recently developed Bayesian multilocus methods aimed at inferring species trees from potentially discordant gene trees. We found strong support for two main clades within Bythograeidae: one comprising the members of the genus Bythograea; and the other comprising the remaining genera. Relationships within each of these two clades were partially resolved. We compare our results with an earlier hypothesis on the phylogenetic relationships among bythograeid genera based on morphology. We also discuss the biogeography of the family in the light of our results. Our species tree analyses reveal differences in how each of the three methods weighs conflicting phylogenetic signal from different gene partitions and how limits on the number of outgroup taxa may affect the results.     

Anatomy and phylogenetic analysis of the neotropical callichthyid catfishes (Ostariophysi, Siluriformes)

Based mainly on morphological characters, the phylogenetic relationships among genera and some species groups of the neotropical family Callichthyidae were examined. A study of the osteology of a generalized callichthyid, Callichthys callichthys (Linnaeus), with detailed comparisons among representatives of the remaining genera in the family, is presented and used as a basis for the phylogenetic analysis. A single most parsimonious tree supported the monophyly of the family Callichthyidae based on 28 derived features and the division of the family in the subfamilies Corydoradinae and Callichthyinae. In the subfamily Corydoradinae, the genus Aspidoras is the sister-group of the clade formed by Corydoras plus Brochis. Five derived features support the monophyly of this clade and four support the monophyly of Brochis. No characters, however, were found to support the genus Corydoras. In the subfamily Callichthyinae, Dianema and Hopbstemum are sister-taxa. Megalechis represents the sister-group of Dianema plus Hoplosternum and Lepthoplosternum represents the sister-group to Megalechis plus Dianema plus Hopbstemum. Finally, Callichthys is considered the least derived member of the subfamily, and is hypothesized as the sister-group of the remaining species. A key to all callichthyid genera is provided.