Reinterpreting recapitulation: systematics of needlefishes and their allies (Teleostei: Beloniformes)

Abstract. As needlefishes (Belonidae) grow, their jaws pass through a "halfbeak" stage that resembles the adult jaw condition of the closely related family of halfbeaks (Hemiramphidae). Based on this pattern, some authors have suggested that halfbeaks are "developmentally arrested" or paedomorphic needlefish derivatives, whereas others have supported the notion that needlefishes are descended from halfbeak-like ancestors and that needlefish ontogeny thereby recapitulates phylogeny. To test these ideas and to better understand evolutionary changes in jaw ontogeny, phylogenetic relationships among genera of needlefishes, sauries (Scomberesocidae), halfbeaks, and flyingfishes (Exocoetidae) were assessed using mitochondrial (cytochrome b and 16S), nuclear (Tmo-4C4), and morphological characters. The resultant tree provides several novel taxonomic findings: (1) flyingfishes appear to be nested within halfbeaks; (2) sauries appear to be nested within needlefishes; and (3) the Indo-West Pacific freshwater halfbeaks appear to be most closely related to the needlefish/saury clade. The structure of the tree falsifies the idea that halfbeaks are paedomorphic needlefishes. Instead, halfbeaks are basal relative to needlefishes, fitting the pattern predicted by the hypothesis of recapitulation. I discuss limitations to phylogenetic perspectives on recapitulation based on discrete character data by comparing aspects of von Baerian and Haeckelian views of the relation between ontogeny and phylogeny.     

Historical biogeography and molecular systematics of the Indo-Pacific genus Dascyllus (Teleostei: Pomacentridae)

The phylogeographical and systematic relationships among species in the tropical marine fish genus Dascyllus were inferred using mitochondrial DNA (mtDNA) sequence data. Although our results were generally consistent with previously published phylogenies based on both morphological and mitochondrial data, our broad taxonomic and geographical sampling design revealed novel insights into the phylogenetic history of Dascyllus that had escaped previous notice. These results include: (a) the polyphyletic nature of D. reticulatus mtDNAs, representing two divergent and geographically separated lineages, one shared with D. flavicaudus and the second forming the sister lineage of D. carneus; (b) the paraphyly of D. trimaculatus relative to the closely related D. abisella; and (c) phylogeographical structure within the widespread taxa D. aruanus and D. trimaculatus. Application of a molecular clock permits us to posit a causative role for tectonic and oceanic changes regarding some Dascyllus speciation events. Finally, we mapped body size and the presence or absence of protogynous sex change on the mtDNA tree, and tested published hypotheses regarding determinants of the evolution of mating system and protogyny in the genus. Our data rejected a model based on body size but not one based on phylogenetic inertia. The ability to change sex arose once in the ancestor to the entire genus, and was lost once in the ancestor of the D. trimaculatus complex. For taxa that are as geographically widespread as many Indo-Pacific genera, this study highlights the importance of adequate geographical sampling when inferring patterns of species diversification and life history evolution.     

Phylogenetic systematics and historical biogeography of the Neotropical electric fish Gymnotus (Teleostei: Gymnotidae)

Phylogenetic interrelationships of the Neotropical electric fish genus Gymnotus are documented from comparative study of phenotypic data. A data matrix was compiled of 113 phenotypic characters for 40 taxa, including 31 recognized Gymnotus species, six allopatric populations of G. carapo, two allopatric populations of G. coropinae, and three gymno‐tiform outgroups. MP analysis yielded 15 trees of equal length, the strict consensus of which is presented as a working hypothesis of Gymnotus interrelationships. Diagnoses are presented for 26 clades, including three species groups; the G. cylindricus group with two species restricted to Middle America, the G. pantherinus group with 12 species in South America, and the G. carapo group with 16 species in South America. The basal division of Gymnotus is between clades endemic to Middle and South America. Both the G. pantherinus and G. carapo groups include trans‐Andean sister‐taxon pairs, suggesting a minimum date for the origins of these groups in the late Middle Miocene (c. 12 Ma.). The geographically widespread species G. carapo is paraphyletic. Analysis of character state evolution shows characters of external morphology are more phylogenetically plastic and provide more phylogenetic information in recent branches than do characters of internal morphology, which themselves provide the more information in deeper branches. Nine regional species assemblages of Gymnotus are recognized, none of which is monophyletic. There are at least two independent origins of Gymnotus species in sediment rich, high conductivity, perennially hypoxic whitewater floodplains (varzea´) derived from an ancestral condition of being restricted to low conductivity non‐floodplain (terra firme) black and clearwater rivers and streams. These phylogenetic, biogeographic and ecological patterns suggest a lengthy and complex history involving numerous instances of speciation, extinction, migration and coexistence in sympatry. Evolution in Gymnotus has been a continent‐wide phenomenon; i.e. Amazonian species richness is not a consequence of strictly Amazonian processes. These patterns are similar to those of other highly diverse groups of Neotropical fishes and do not resemble those of monophyletic, rapidly generated species flocks.     

线粒体DNA序列特点与昆虫系统学研究

昆虫线粒体DNA是昆虫分子系统学研究中应用最为广泛的遗传物质之一。线粒体DNA具有进化速率较核DNA快 ,遗传过程不发生基因重组、倒位、易位等突变 ,并且遵守严格的母系遗传方式等特点。本文概述了mtDNA中的rRNA、tRNA、蛋白编码基因和非编码区的一般属性 ,分析了它们在昆虫分子系统学研究中的应用价值 ,以及应用DNA序列数据来推导分类阶 (单 )元的系统发育关系时 ,基因或DNA片段选择的重要性     

蝗虫分子系统学研究进展

本文从遗传多样性、近缘种鉴别以及分子进化和系统发育重建 3个方面综述了蝗虫分子系统学的研究进展。     

Phylogenetic study of the Characinae (Teleostei: Characiformes: Characidae)

The Characinae is a subunit of the Characidae of special significance in including Charax, the type genus of the family and the order Characiformes. Twelve genera and 79 species have been traditionally assigned to the Characinae, but the subfamily still lacks a phylogenetic diagnosis. Herein, a data matrix including 150 morphological characters and 64 taxa (35 species representing all genera of the Characinae and 29 included in other lineages within the Characiformes) was submitted to two cladistic analyses that differ in the inclusion/exclusion of Priocharax due to the difficulty of coding most of the character states in the miniature species of this genus. Both analyses resulted in a non‐monophyletic Characinae and this subfamily is herein restricted to only seven of the original 12 genera forming the clade (Phenacogaster((Charax Roeboides)(Acanthocharax(Cynopotamus(Acestrocephalus Galeocharax))))), which is supported by ten non‐ambiguous synapomorphies and is more closely related to other genera of the Characidae than those traditionally placed in the subfamily. A second clade includes the members of the tribe Heterocharacini (Lonchogenys(Heterocharax Hoplocharax)) as the sister‐group of Gnathocharax, supported by seven non‐ambiguous synapomorphies. This clade is more closely related to a taxon formed by Roestes and Gilbertolus based on seven non‐ambiguous synapomorphies. Results do not corroborate a close relationship between Roestes–Gilbertolus and the Cynodontinae. Inclusion of the genus Priocharax suggests that it is related more closely to the Heterocharacini, but the profound modifications in its anatomy possibly related to ontogenetic truncations obscure a better understanding of its relationships. A new classification of the Characinae and the Heterocharacinae is proposed. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 165 , 809–915.     

Turbellaria Lecithoepitheliata: morphology,systematics, phylogeny

A review of the literature on the world fauna of lecithoepitheliata and Prolecithophora and extensive observations by the author on species of the Prorhynchidae, including cosmopolitan species as well as some endemic to Lake Baikal, and on endemic species of Baicalarctiinae (Prolecithophora) show that the Prorhynchidae and Gnosonesimidae are properly classified in the Neoophora. The morphological similarity of the Prorhynchidae and Prolecithophora proves these taxa are closely related. A hypothesis relating the Lecithoepitheliata Prorhynchidae to primitive prolecithophorans appears to be the only tenable hypothesis on the phylogeny of Lecithoepitheliata. The question of whether the Lecithoepitheliata is monophyletic still needs investigation; more evidence on the phylogeny of the Gnosonesimidae is needed.     

Diversity,phylogeny and biogeography of Systomus (Teleostei,Cyprinidae) in Sri Lanka

The South and South-East Asian freshwater fish genus Systomus (Cyprinidae) comprises 17 valid species. Six nominal species, including three endemics, have been reported from Sri Lanka, a continental island separated from India by a shallow-shelf sea. The species diversity of Systomus on the island has until now not been assessed; neither has an evaluation been made of their phylogenetic history. Here, based on an analysis of the nuclear recombination activating protein 1 (rag1), and mitochondrial cytochrome c oxidase subunit 1 (cox1) and cytochrome b (cytb) gene markers, and a morphological examination of 143 specimens from 49 locations in Sri Lanka, we reassess the diversity of Systomus on the island and analyse patterns of their evolution and biogeography. Divergence-time estimates, based on a substitution rate calibration, date the basal split between Systomus and its sister group, the Afrotropical small barbs, to 30.0 Ma (95% highest posterior density: 25.4–35.2 Ma). The species of Systomus belong to two distinct clades. The first includes the Sri Lankan endemics S. asoka, S. martenstyni and S. pleurotaenia, which comprise an insular diversification following the immigration of a common ancestor during the Oligocene. The second, which includes the remaining species of Indian, Sri Lankan and South-East Asian Systomus, has a crown age dating to the Late Miocene. Morphological and molecular species delimitation analyses failed to validate the two nominal species, S. spilurus and S. timbiri, previously reported from Sri Lanka: both are considered synonyms of S. sarana, as are the nomina S. chryseus, S. chrysopoma, S. laticeps, S. rufus, S. pinnauratus and S. subnasutus. Four genetically and geographically discrete lineages of S. sarana occur in the island, and three in India. Molecular species delimitation analysis suggests these all belong to a single species, S. sarana. The genetically distinct Sri Lankan populations of S. sarana result from Plio-Pleistocene dispersal or vicariance events between India and Sri Lanka—as a result of emergence and inundation of the now submerged isthmus connecting the two landmasses—as well as autochthonous insular diversification.     

Osteology and ontogeny of the wrymouths,genus Cryptacanthodes (Cottiformes: Zoarcoidei: Cryptacanthodidae)

The four species included in the family Cryptacanthodidae are eel‐like, burrowing fishes distributed in the cold‐temperate coastal waters of the North Pacific and the western North Atlantic. This study describes the osteology and aspects of the ontogenetic skeletal development of two species, Cryptacanthodes maculatus from the western North Atlantic and C. aleutensis from the eastern North Pacific. We discuss the relationships of Cryptacanthodidae among other zoarcoid families. The Cryptacanthodidae have been previously included in the Stichaeidae, but removed and classified as a separate family based on the skull, pectoral radial, and cephalic lateral‐line morphology. Our observations (similarities in gill arch and pectoral girdle morphology; specifically, a thin sheet‐like flange of bone from the posterior margin of the supracleithrum) suggest a close relationship to at least some of the members of the family Stichaeidae. J. Morphol. 276:185–208, 2015. © 2014 Wiley Periodicals, Inc.     

Phylogenetic relationships of the Sparidae (Teleostei: Percoidei) and implications for convergent trophic evolution

A phylogenetic analysis of the majority of sparid genera and representatives of the sparoid families Centracanthidae, Lethrinidae and Nemipteridae is presented using 87 predominately osteological characters. The Sparidae constitute a monophyletic grouping, with the inclusion of the centracanthid Spicara smaris , which nests deep within the ingroup. The phylogeny was then used to investigate agreement with the most recent molecular study, taxonomic stability of subfamilial classification and the evolution of feeding strategies. Results show that the incongruence between morphological and molecular data appears largely to be an artifact of errors in rooting. However, there appears to be real and substantial conflict between the molecular tree and the morphological data, which is not attributable to the different positions of the least congruent taxa. The data support the molecular hypothesis that none of the subfamilial classification, based on dentition and trophic specialization, is monophyletic, and should be rejected pending further taxonomic revision. The phylogeny supports multiple independent origins of trophic types and it is suggested that the evolutionary plasticity of the oral teeth of sparids has been fundamental to the adaptive radiation of this family compared to their closest allies. ©2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 76 , 269–301.     

Phylogenetic relationships among atheriniform fishes (Teleostei: Atherinomorpha)

We present evidence from adult and larval morphology for the monophyly and relationships of Atheriniformes, using other atherinomorphs, mugilids and acanthomorph fishes as outgroups. Atheriniformes is diagnosed by ten characters (larval: short preanal length, single mid-dorsal row of melanophores; adult: vomerine ventral face concave, long Al muscle tendon to lacrimal, two anterior infraorbital bones, pelvic-rib ligament, pelvic medial plate not extended to anterior end, and second dorsal-fin spine flexible). We recognize six families within the order, the hierarchical relationships among which are: (Atherinopsidae (Notocheiridae (Melanotaeniidae (Atherionidae (Phallostethidae, Atherinidae))))). Other major conclusions include: (1) Atherinopsidae (Menidiinae, Atherinopsinae) is diagnosed by 20 characters (e.g. ethmomaxillary ligament attached to palatine dorsal process, ventral postcleithrum with two dorsal rami); (2) Melanotaeniidae (Bedotiinae (Melanotaeniinae (Telmatherinini, Pseudomugilini))) is diagnosed by six characters (e.g. absence of second dorsal-fin spine, sexual dimorphism in body colour and median-fin development, greater body depth); (3) Dentatherina is in Phallostethidae; (4) Atherinidae (Atherinomorinae (Craterocephalinae, Atherininae)) is diagnosed by three characters (lacrimal notch, ventral postcleithrum between first and second pleural ribs, pelvic ventral spine); (5) Atherinidae and Phallostethidae form the Atherinoidea clade diagnosed by seven characters (e.g. interopercle dorsal process absent, dorsal wings of urohyal absent, ventral postcleithrum laminar, pelvic medial plate extended to anterior end, presence of anal plate). Bedotia, Rhodes , and melanotaeniines are shown to be derived within atheriniforms rather than the plesiomorphic sister groups to a paraphyletic 'atherinoid' group. We also demonstrate that groups traditionally placed in Atherinidae (Menidiinae, Atherininae, Atherioninae, etc.) comprise a paraphyletic assemblage.     

Phylogenetic relationships of subgenera of the genus Exorista Meigen,with a revision of the Japanese species (Diptera: Tachinidae)

Phylogenetic relationships of the subgenera of Exorista Meigen (Diptera: Tachinidae) are inferred from morphological data. Our results show that the genus Exorista is not monophyletic and that members of the subgenus Spixomyia Crosskey are divided into two clades. Each subgenus is redefined based on male and female morphological features. The Japanese species of Exorista are revised and classified into five subgenera: Adenia Robineau‐Desvoidy, Exorista Meigen, Podotachina Brauer and Bergenstamm, Ptilotachina Brauer and Bergenstamm, and Spixomyia Crosskey. Thirteen species are recognized, including two newly recorded species, Exorista (Adenia) cuneata Herting and Exorista (Spixomyia) lepis Chao. Exorista cantans Mesnil is transferred to the subgenus Podotachina from Spixomyia.     

Molecular phylogeny of the Forcipulatacea (Asteroidea: Echinodermata): systematics and biogeography

We present a comprehensively sampled three‐gene phylogeny of the monophyletic Forcipulatacea, one of three major lineages within the crown‐group Asteroidea. We present substantially more Southern Hemisphere and deep‐sea taxa than were sampled in previous molecular studies of this group. Morphologically distinct groups, such as the Brisingida and the Zoroasteridae, are upheld as monophyletic. Brisingida is supported as the derived sister group to the Asteriidae (restricted), rather than as a basal taxon. The Asteriidae is paraphyletic, and is broken up into the Stichasteridae and four primary asteriid clades: (1) a highly diverse boreal clade, containing members from the Arctic and sub‐Arctic in the Northern Hemisphere; (2) the genus Sclerasterias; (3) and (4) two sister clades that contain asteriids from the Antarctic and pantropical regions. The Stichasteridae, which was regarded as a synonym of the Asteriidae, is resurrected by our results, and represents the most diverse Southern Hemisphere forcipulatacean clade (although two deep‐sea stichasterid genera occur in the Northern Hemisphere). The Labidiasteridae is artificial, and should be synonymized into the Heliasteridae. The Pedicellasteridae is paraphyletic, with three separate clades containing pedicellasterid taxa emerging among the basal Forcipulatacea. Fossils and timing estimates from species‐level phylogeographic studies are consistent with prior phylogenetic hypotheses for the Forcipulatacea, suggesting diversification of basal taxa in the early Mesozoic, with some evidence for more widely distributed ranges from Cretacous taxa. Our analysis suggests a hypothesis of an older fauna present in the Antarctic during the Eocene, which was succeeded by a modern Antarctic fauna that is represented by the recently derived Antarctic Asteriidae and other forcipulatacean lineages. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162 , 646–660.     

Phylogenetic systematics of the nymphaeales

A cladistic analysis was applied to reveal the phylogenetic relationships among the Nymphaeales. Seventeen out of twenty three characters in gross morphology, anatomy and palynology were analyzed, for their evolutionary polarities. From the results of the present analysis, the phylogenetic status of each genus and their relationships were clarified: 1)Nelumbo is a distinct taxon and is presumed to have originated from an ancestral stock of the Nymphaeales; 2)Ceratophyllum has a close phylogenetic relationship withCabomba; and 3) in the Nymphaeaceaesensu stricto, Nuphar and the remaining genral constitute a monophyletic group. A conclusion obtained from the present analysis was that the following three families should be recognized in the Nymphaeales; Nelumbonaceae Nymphaeaceae, and Ceratophyllaceae. The generaBrasenia andCabomba are traditionally classified in the Nymphaeaceae or in the independent family Cabombaceae. However, they should be included in the family Ceratophyllaceae.     

Phylogenetic utility of the first two introns of the S7 ribosomal protein gene in African electric fishes (Mormyroidea: Teleostei) and congruence with other molecular markers

A series of recent molecular systematic studies of the African electric fishes (Mormyroidea) have challenged many aspects of their traditional taxonomy and precladistic hypotheses of their phylogeny. However, poor resolution of some interrelationships within the subfamily Mormyrinae in these studies highlights the need for additional data and analyses. Here we evaluate the phylogenetic information content of nucleotide sequences from the first two introns of the low‐copy nuclear S7 ribosomal protein gene in 40 mormyroid species. Alignment of S7 sequences from 38 taxa within the subfamily Mormyrinae is non‐problematic, but these are difficult to align with sequences of Petrocephalus bovei (Petrocephalinae) and Gymnarchus niloticus (Gymnarchidae), which we exclude from our analysis. There are no significant differences in base frequencies among these sequences and base compositional bias is low. Maximum parsimony (MP) analysis on the S7 dataset, designating Myomyrus macrops as the outgroup, generates a phylogenetic hypothesis for these taxa with a low level of homoplasy (RI = 0.87). We examine agreement between the S7 data with previously published mitochondrial (12S/16S, cytochrome b) and nuclear (rag 2) datasets for the same taxa by means of incongruence length difference tests and partitioned Bremer support (decay) analysis. While we find significant agreement between the S7 dataset and the others, MP analysis of the S7 data alone and in combination with the other datasets indicates two novel relationships within the Mormyrinae: (1) Mormyrus is the sister group to Brienomyrus brachyistius and Isichthys henryi, and (2) Hippopotamyrus pictus is the sister group of a clade, previously recovered, containing Marcusenius senegalensis. S7 data provide additional support for a number of clades recovered in the earlier molecular studies, some of which conflict with current mormyrid taxonomy. Inferred indels and a single inversion in the S7 fragment provide supplemental character support for many of these relationships. These phylogenetic results strengthen recent hypotheses concerning the evolution of electric organ structure in these fishes. The evolutionary characteristics of this nuclear marker and its phylogenetic utility in this group suggests that it could be widely useful for systematic studies at the subfamilial level in teleost fishes. © 2003 The Linnean Society of London. Biological Journal of the Linnean Society, 2003, 78 , 273–292.     

Phylogenetic systematics of Erythronium (Liliaceae): morphological and molecular analyses

For a nearly complete set of species of Erythronium (Liliaceae), we examined two plastid loci (the rps16 intron and the 5′ trnK intron, excluding the matK exon), one nuclear locus (nrITS) and morphology to evaluate species relationships and that of Erythronium to Amana, the putatively most closely allied genus. A matrix of morphological characters was developed through observation of around 900 living and herbarium specimens; evolution of these was examined using character optimization on the combined (total‐evidence) tree. Parsimony methods were used to examine the morphological and molecular data sets produced, both separately and in combination, with Bayesian methods also used on the molecular data sets. These established that the genus is probably sister to Amana (although most analyses placed Amana inside Erythronium) and that Tulipa is sister to the pair of Erythronium and Amana. Within Erythronium, there are three strongly supported geographically distinct clades: (1) Eurasian and (2) eastern and (3) western North American. Separation of species in these three clades is less clear, particularly among the western North American taxa. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, ●●, ●●–●●.     

Phylogenetic systematics and historical biogeography of the Neotropical electric fish Sternopygus (Teleostei: Gymnotiformes)

Interrelationships among 10 extant species of the Neotropical electric fish Sternopygus are inferred from phylogenetic analysis of 66 morphological characters, including features of pigmentation, body proportions, meristics and osteology. A total of 287 lots containing 677 specimens were examined. The important findings of this study are: (1) S. branco is the most basal species unique among congeners in being restricted to whitewater rivers in the Central Amazon Basin, (2) S. sp. ‘cau’ from the Rio Caura of Venezuela is the sister taxon to (S. obtusirostris + S. astrabes), (3) S. castroi is a junior synonym of S. astrabes, (4) S. macrurus is the sister taxon to (S. arenatus + S. xingu + S. aequilabiatus species group) and (5) S. arenatus is the sister taxon to (S. xingu + S. aequilabiatus species group). A key to the adults of Sternopygus species is provided. Several features of S. astrabes previously thought to be plesiomorphic are now considered derived, including: short body cavity, paedomorphic cranial osteology, and the habitat restriction to terra firme streams. Sternopygus species assemblages in the Pacific (trans‐Andean) and Atlantic (cis‐Andean) slopes of northwestern South America are not monophyletic and do not result exclusively from local or regional radiations. The clade composed of S. macrurus, S. arenatus, S. xingu and the S. aequilabiatus species group is inferred to predate the Middle Miocene uplift of the Eastern Cordillera (c. 11.8–12.2 Ma). As currently recognized S. macrurus is the most widely distributed and most eurytopic gymnotiform species, inhabiting all hydrogeographical regions of tropical South America and most lowland aquatic habitats. Other Sternopygus species have much more restricted geographic and ecological distributions. Perceptions of phylogenetic patterns in Sternopygus are shown to be highly sensitive to taxon sampling.     

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.