Systematic relationships of the palaeogene family Presbyornithidae (Aves: Anseriformes)

The early Tertiary (Paleocene and Eocene) family Presbyornithidae is one of the most completely known group of fossil birds. Essentially all parts of the skeleton are represented in the fossil record, allowing a thorough analysis of the phylogenetic position of the family. Forty-two families of nonpasserine birds representing the orders Ciconiiformes, Anseriformes, Galliformes, Gruiformes and Charadriiformes, were included in a cladistic analysis of 71 skeletal characters. The previously suggested anseriform affinity of the Presbyornithidae was confirmed. Furthermore, the family proved to be closer to the Anatidae than to the Anhimidae or Anseranatidae. The many postcranial similarities with certain charadriiform birds as the Burhinidae, obviously are plesiomorphies. By this observation, a better undestanding of character evolution in nonpasserine skeletal morphology is gained. The often suggested close relationship of anseriform and galliform birds is not confirmed by osteology. Instead, the Anseriformes and the Phoenicopteridae form a monophyletic clade that is the sister to the remaining ciconiiform birds. This result renders the Ciconiiformes sensu Wetmore (1960) polyphyletic.     

A phylogenetic supertree of the fowls (Galloanserae, Aves)

The fowls (Anseriformes and Galliformes) comprise one of the major lineages of birds and occupy almost all biogeographical regions of the world. The group contains the most economically important of all bird species, each with a long history of domestication, and is an ideal model for studying ecological and evolutionary patterns. Yet, despite the relatively large amount of systematic attention fowls have attracted because of their socio‐economic and biological importance, the species‐level relationships within this clade remain controversial. Here we used the supertree method matrix representation with parsimony to generate a robust estimate of species‐level relationships of fowls. The supertree represents one of the most comprehensive estimates for the group to date, including 376 species (83.2% of all species; all 162 Anseriformes and 214 Galliformes) and all but one genera. The supertree was well‐resolved (81.1%) and supported the monophyly of both Anseriformes and Galliformes. The supertree supported the partitioning of Anseriformes into the three traditional families Anhimidae, Anseranatidae, and Anatidae, although it provided relatively poor resolution within Anatidae. For Galliformes, the majority‐rule supertree was largely consistent with the hypothesis of sequential sister‐group relationships between Megapodiidae, Cracidae, and the remaining Galliformes. However, our species‐level supertree indicated that more than 30% of the polytypic genera examined were not monophyletic, suggesting that results from genus‐level comparative studies using the average of the constituent species’ traits should be interpreted with caution until analogous species‐level comparative studies are available. Poorly resolved areas of the supertree reflect gaps or outstanding conflict within the existing phylogenetic database, highlighting areas in need of more study in addition to those species not present on the tree at all due to insufficient information. Even so, our supertree will provide a valuable foundation for understanding the diverse biology of fowls in a robust phylogenetic framework.     

Morphology of the quadrate in the Eocene anseriform Presbyornis and extant galloanserine birds

Despite the notoriety, phylogenetic significance, and large number of available specimens of Presbyornis, its cranial anatomy has never been studied in detail, and its quadrate has been partly misinterpreted. We studied five quadrates of Presbyornis that reveal features hitherto unknown in the anseriforms but otherwise present in galliforms. As a result, we analyzed the variable quadrate characters among all extant galloanserine families and identified synapomorphies and other morphological variation among the major galloanserine clades. In terms of quadrate morphology, Presbyornis is more plesiomorphic than any extant anseriform (including the Anhimidae) and shares ancestral galloanserine characters with the Megapodiidae, the earliest branch of extant galliforms. The quadrate's morphology is inconsistent with the currently accepted anseriform phylogeny that nests Presbyornis within the crown‐group as a close relative of the Anatidae. The presbyornithid quadrates exhibit an unusual variation in the presence of a caudomedial pneumatic foramen, which we interpret as a result of a discontinuous change in the growth path of the pneumatic diverticulum. Another episode of morphogenetic imbalance in the growth path of the pneumatic diverticulum may have accompanied the disappearance of the basiorbital pneumatic foramen (along with the pneumatization of the pterygoid) at the origin of the crown‐group anseriforms. This episode is marked by the striking individual variation in the presence and location of pneumatic foramina in the mandibular part of the quadrate in the Anhimidae. J. Morphol., 2010. © 2009 Wiley‐Liss, Inc.     

Hypothetical stages in the evolution of modern ducks and flamingos

A re-evaluation of the evolutionary stages in modern ducks and flamingos is permitted by recent discoveries of large concentrations of the Eocene avian fossil Presbyornis pervetus from Wyoming.Presbyornis was originally described as a new family, genus and species of recurvirostrid shorebird. Recent discoveries of nearly complete elements (including the skull) of this species show it to be a mosaic, combining characters primarily of the modem avian orders Charadrii-formes (shorebirds), ducks and allies (Anseriformes), and living flamingos (Phoenicopteriformes). Thus, living flamingos are mosaics, combining characters primarily of the shorebirds and the ducks and allies. Presbyornis is probably a temporal relict (like the anthracosaur Seymouria), structurally close to the ancestry of both modern ducks and modern flamingos, and was derived from shorebird ancestry.     

Australia's oldest Anseriform fossil: a quadrate from the Early Eocene Tingamarra Fauna

Abstract: A partial quadrate (essentially the otic part) from the nonmarine, earliest Eocene (54.6 Ma) Tingamarra Local Fauna in Queensland, Australia, has been identified as the oldest Australian anseriform fossil. The Tingamarra quadrate shows a combination of plesiomorphic anseriform characters with a unique synapomorphic character complex of the Anhimidae (screamers), which today are endemic to South America. In concert with the basal position of the Anhimidae among the crown‐group anseriforms, this set of characters suggests a stem group of the Anhimidae, raising a possibility of the Transantarctic migration of stem anhimids to South America. The quadrate morphology supports palaeognathous rather than recently claimed anhimid relationships of the Dromornithidae and identifies Sylviornis as an anseriform rather than a galliform.     

More taxa,more characters: the hoatzin problem is still unresolved

The apparently rapid and ancient diversification of many avian orders complicates the resolution of their relationships using molecular data. Recent studies based on complete mitochondrial DNA (mtDNA) sequences or shorter lengths of nuclear sequence have helped corroborate the basic structure of the avian tree (e.g., a basal split between Paleognathae and Neognathae) but have made relatively little progress in resolving relationships among the many orders within Neoaves. We explored the potential of a moderately sized mtDNA data set ( approximately 5000 bp for each of 41 taxa), supplemented with data from a nuclear intron ( approximately 700 bp per taxon), to resolve relationships among avian orders. Our sampling of taxa addresses two issues: (1). the sister relationship and monophyly, respectively, of Anseriformes and Galliformes and (2). relationships of the enigmatic hoatzin Opisthocomus hoazin. Our analyses support a basal split between Galloanserae and Neoaves within Neognathae and monophyly of both Galliformes and Anseriformes. Within Galliformes, megapodes and then cracids branch basally. Within Anseriformes, mitochondrial data support a screamer (Anhimidae) plus magpie goose (Anseranatidae) clade. This result, however, may be an artifact of divergent base composition in one of the two anatids we sampled. With deletion of the latter taxon, Anseranas is sister to anatids as in traditional arrangements and recent morphological studies. Although our data provide limited resolution of relationships within Neoaves, we find no support for a sister relationship between either cuckoos (Cuculiformes) or turacos (Musophagiformes) and hoatzin. Both mitochondrial and nuclear data are consistent with a relationship between hoatzin and doves (Columbiformes), although this result is weakly supported. We also show that mtDNA sequences reported in another recent study included pervasive errors that biased the analysis towards finding a sister relationship between hoatzin and turacos.     

The Complete Mitochondrial Genome of Aix galericulata and Tadorna ferruginea: Bearings on Their Phylogenetic Position in the Anseriformes

Aix galericulata and Tadorna ferruginea are two Anatidae species representing different taxonomic groups of Anseriformes. We used a PCR-based method to determine the complete mtDNAs of both species, and estimated phylogenetic trees based on the complete mtDNA alignment of these and 14 other Anseriforme species, to clarify Anseriform phylogenetics. Phylogenetic trees were also estimated using a multiple sequence alignment of three mitochondrial genes (Cyt b, ND2, and COI) from 68 typical species in GenBank, to further clarify the phylogenetic relationships of several groups among the Anseriformes. The new mtDNAs are circular molecules, 16,651 bp (Aix galericulata) and 16,639 bp (Tadorna ferruginea) in length, containing the 37 typical genes, with an identical gene order and arrangement as those of other Anseriformes. Comparing the protein-coding genes among the mtDNAs of 16 Anseriforme species, ATG is generally the start codon, TAA is the most frequent stop codon, one of three, TAA, TAG, and T-, commonly observed. All tRNAs could be folded into canonical cloverleaf secondary structures except for tRNASer (AGY) and tRNALeu (CUN), which are missing the "DHU" arm.Phylogenetic relationships demonstrate that Aix galericula and Tadorna ferruginea are in the same group, the Tadorninae lineage, based on our analyses of complete mtDNAs and combined gene data. Molecular phylogenetic analysis suggests the 68 species of Anseriform birds be divided into three families: Anhimidae, Anatidae, and Anseranatidae. The results suggest Anatidae birds be divided into five subfamilies: Anatinae, Tadorninae, Anserinae, Oxyurinae, and Dendrocygninae. Oxyurinae and Dendrocygninae should not belong to Anserinae, but rather represent independent subfamilies. The Anatinae includes species from the tribes Mergini, Somaterini, Anatini, and Aythyini. The Anserinae includes species from the tribes Anserini and Cygnini.     

Phylogenetic analysis of pelecaniformes (aves) based on osteological data: implications for waterbird phylogeny and fossil calibration studies

Background

Debate regarding the monophyly and relationships of the avian order Pelecaniformes represents a classic example of discord between morphological and molecular estimates of phylogeny. This lack of consensus hampers interpretation of the group''s fossil record, which has major implications for understanding patterns of character evolution (e.g., the evolution of wing-propelled diving) and temporal diversification (e.g., the origins of modern families). Relationships of the Pelecaniformes were inferred through parsimony analyses of an osteological dataset encompassing 59 taxa and 464 characters. The relationships of the Plotopteridae, an extinct family of wing-propelled divers, and several other fossil pelecaniforms (Limnofregata, Prophaethon, Lithoptila, ?Borvocarbo stoeffelensis) were also assessed. The antiquity of these taxa and their purported status as stem members of extant families makes them valuable for studies of higher-level avian diversification.

Methodology/Principal Findings

Pelecaniform monophyly is not recovered, with Phaethontidae recovered as distantly related to all other pelecaniforms, which are supported as a monophyletic Steganopodes. Some anatomical partitions of the dataset possess different phylogenetic signals, and partitioned analyses reveal that these discrepancies are localized outside of Steganopodes, and primarily due to a few labile taxa. The Plotopteridae are recovered as the sister taxon to Phalacrocoracoidea, and the relationships of other fossil pelecaniforms representing key calibration points are well supported, including Limnofregata (sister taxon to Fregatidae), Prophaethon and Lithoptila (successive sister taxa to Phaethontidae), and ?Borvocarbo stoeffelensis (sister taxon to Phalacrocoracidae). These relationships are invariant when ‘backbone’ constraints based on recent avian phylogenies are imposed.

Conclusions/Significance

Relationships of extant pelecaniforms inferred from morphology are more congruent with molecular phylogenies than previously assumed, though notable conflicts remain. The phylogenetic position of the Plotopteridae implies that wing-propelled diving evolved independently in plotopterids and penguins, representing a remarkable case of convergent evolution. Despite robust support for the placement of fossil taxa representing key calibration points, the successive outgroup relationships of several “stem fossil + crown family” clades are variable and poorly supported across recent studies of avian phylogeny. Thus, the impact these fossils have on inferred patterns of temporal diversification depends heavily on the resolution of deep nodes in avian phylogeny.     

New nuclear evidence for the oldest divergence among neognath birds: the phylogenetic utility of ZENK (i)

To date, there is little consensus concerning the phylogenetic relationships among neognath orders, which include all extant birds except ratites and tinamous. Different data sets, both molecular and morphologic, have yielded radically different and often unresolved ordinal topologies, especially within the neoaves clade. This lack of resolution and ongoing conflict indicates a need for additional phylogenetic characters to be applied to the question of higher-level avian phylogeny. In this study, sequences of a single-copy nuclear gene, ZENK, were used to reconstruct an ordinal-level phylogeny of neognath birds. Strong support was indicated for the oldest divergence within Neognathae; the chicken- and duck-like birds formed a clade that was sister to all other modern birds. In addition, many families of traditional taxonomic orders clustered together in the ZENK tree, indicating the gene's general phylogenetic reliability. However, within the neoaves clade, there was little support for relationships among orders, which is a result similar to all other recent molecular studies of higher-level avian phylogeny. This similarity among studies suggests the possibility of a rapid radiation of the major neoaves lineages. Despite the ongoing lack of neoaves resolution, ZENK's sequence divergence and base composition patterns indicate its general utility as a new phylogenetic marker for higher-level avian systematics.     

Phylogenetic relationships and biogeography of the Ipsiura cuckoo wasps (Hymenoptera: Chrysididae)

Phylogenetic studies addressing relationships among chrysidid wasps have been limited. There are no hypotheses proposed for the Neotropical lineages of Chrysidini other than the classic cladogram published in the 1990s by Kimsey and Bohart. Herein we present a cladistic analysis based on 64 morphological characters coded for 54 species of Chrysidini, 32 of them being Ipsiura and 22 representing Caenochrysis, Chrysis, Exochrysis, Gaullea, Neochrysis, and Pleurochrysis. The species of Ipsiura were recovered as monophyletic and as the sister clade of Neochrysis in all most parsimonious trees. We discuss the high plasticity of some morphological characters as evidenced by their high homoplasy in the phylogenetic results, and we clarify the main morphological changes inferred on the phylogenetic tree for this genus. The effects of the inferred homoplasy were evaluated under an implied weighting cladistic analysis, and from a probabilistic perspective with Bayesian inference. Those alternative strategies did not alter the general conclusions about the monophyly of Ipsiura or the generic relationships in Chrysidini (changes were noticed in the species‐level relationships within certain parts of Ipsiura, where low branch support was common across all approaches). Among the species groups proposed by Linsenmaier (1985), only the marginalis group was recovered as monophyletic. We also evaluated the convoluted biogeographic history of the group. The resulting historical reconstructions indicate a complicated scenario of diversification of these wasps in the Neotropics, and a close association with forested biomes is discussed.     

Molecules, morphology and fossils: a comprehensive approach to odonate phylogeny and the evolution of the odonate wing

We undertook a comprehensive morphological and molecular phylogenetic analysis of dragonfly phylogeny, examining both extant and fossil lineages in simultaneous analyses. The legitimacy of higher‐level family groups and the phylogenetic relationship between families were tested. Thirteen families were supported as monophyletic (Aeshnidae, Calopterygidae, Chlorocyphidae, Euphaeidae, Gomphidae, Isostictidae, Lestidae, Libellulidae, Petaluridae, Platystictidae, Polythoridae, Pseudostigmatidae and Synthemistidae) and eight as non‐monophyletic (Amphipterygidae, Coenagrionidae, Corduliidae, Megapodagrionidae, Protoneuridae and Synlestidae), although Perilestidae and Platycnemididae were recovered as monophyletic under Bayesian analyses. Nine families were represented by one species, thus monophyly was not tested (Epiophlebiidae, Austropetaliidae, Chlorogomphidae, Cordulegastridae, Macromiidae, Chorismagrionidae, Diphlebiidae, Lestoideidae and Pseudolestidae). Epiprocta and Zygoptera were recovered as monophyletic. Ditaxinerua is supported as the sister lineage to Odonata, Epiophlebiidae and the lestid‐like damselflies are sister to the Epiprocta and Zygoptera, respectively. Austropetaliidae + Aeshnidae is the sister lineage to the remaining Anisoptera. Tarsophlebia's placement as sister to Epiprocta or as sister to Epiprocta + Zygoptera was not resolved. Refinements are made to the current classification. Fossil taxa did not seem to provide signals crucial to recovering a robust phylogeny, but were critical to understanding the evolution of key morphological features associated with flight. Characters associated with wing structure were optimized revealing two wing character complexes: the pterostigma–nodal brace complex and the costal wing base & costal–ScP junction complex. In turn, these two complexes appear to be associated; the pterostigma–nodal brace complex allowing for further modification of the wing characters comprised within the costal wing base & costal–ScP junction complex leading the modern odonate wing. © The Willi Hennig Society 2008.     

Calibration of avian molecular clocks

Molecular clocks can be calibrated using fossils within the group under study (internal calibration) or outside of the group (external calibration). Both types of calibration have their advantages and disadvantages. An internal calibration may reduce extrapolation error but may not be from the best fossil record, raising the issue of nonindependence. An external calibration may be more independent but also may have a greater extrapolation error. Here, we used the advantages of both methods by applying a sequential calibration to avian molecular clocks. We estimated a basal divergence within birds, the split between fowl (Galliformes) and ducks (Anseriformes), to be 89.8 +/- 6.97 MYA using an external calibration and 12 rate-constant nuclear genes. In turn, this time estimate was used as an internal calibration for three species-rich avian molecular data sets: mtDNA, DNA-DNA hybridization, and transferrin immunological distances. The resulting time estimates indicate that many major clades of modern birds had their origins within the Cretaceous. This supports earlier studies that identified large gaps in the avian fossil record and suggests that modern birds may have coexisted with other avian lineages for an extended period during the Cretaceous. The new time estimates are concordant with a continental breakup model for the origin of ratites.     

Phylogenetics, biogeography and classification of, and character evolution in, gamebirds (Aves: Galliformes): effects of character exclusion, data partitioning and missing data

The phylogenetic relationships, biogeography and classification of, and morpho‐behavioral (M/B) evolution in, gamebirds (Aves: Galliformes) are investigated. In‐group taxa (rooted on representatives of the Anseriformes) include 158 species representing all suprageneric galliform taxa and 65 genera. The characters include 102 M/B attributes and 4452 nucleic acid base pairs from mitochondrial cytochrome b (CYT B), NADH dehydrogenase subunit 2 (ND2), 12S ribosomal DNA (12S) and control region (CR), and nuclear ovomucoid intron G (OVO‐G). Analysis of the combined character data set yielded a single, completely resolved cladogram that had the highest levels of jackknife support, which suggests a need for a revised classification for the phasianine galliforms. Adding 102 M/B characters to the combined CYT B and ND2 partitions (2184 characters) decisively overturns the topology suggested by analysis of the two mtDNA partitions alone, refuting the view that M/B characters should be excluded from phylogenetic analyses because of their relatively small number and putative character state ambiguity. Exclusion of the OVO‐G partition (with > 70% missing data) from the combined data set had no effect on cladistic structure, but slightly lowered jackknife support at several nodes. Exclusion of third positions of codons in an analysis of a CYT B + ND2 partition resulted in a massive loss of resolution and support, and even failed to recover the monophyly of the Galliformes with jackknife support. A combined analysis of putatively less informative, “non‐coding” characters (CYT B/ND2 third position sites + CR +12S + OVO‐G sequences) yielded a highly resolved consensus cladogram congruent with the combined‐evidence cladogram. Traditionally recognized suprageneric galliform taxa emerging in the combined cladogram are: the families Megapodiidae (megapodes), Cracidae (cracids), Numididae (guineafowls), Odontophoridae (New World quails) and Phasianidae (pheasants, pavonines, partridges, quails, francolins, spurfowls and grouse) and the subfamilies Cracinae (curassows, chachalacas and the horned guan), Penelopinae (remaining guans), Pavoninae sensu lato (peafowls, peacock pheasants and argus pheasants), Tetraoninae (grouse) and Phasianinae (pheasants minus Gallus). The monophyly of some traditional groupings (e.g., the perdicinae: partridges/quails/francolins) is rejected decisively, contrasted by the emergence of other unexpected groupings. The most remarkable phylogenetic results are the placement of endemic African galliforms as sisters to geographically far‐distant taxa in Asia and the Americas. Biogeographically, the combined‐data cladogram supports the hypothesis that basal lineages of galliforms diverged prior to the Cretaceous/Tertiary (K‐T) Event and that the subsequent cladogenesis was influenced by the break‐up of Gondwana. The evolution of gamebirds in Africa, Asia and the Americas has a far more complicated historical biogeography than suggested to date. With regard to character evolution: spurs appear to have evolved at least twice within the Galliformes; a relatively large number of tail feathers (≥ 14) at least three times; polygyny at least twice; and sexual dimorphism many times. © The Willi Hennig Society 2006.     

The mitochondrial genome of Cygnus columbianus, the Whistling Swan.

The nucleotide sequence of the mitochondrial genome of the Whistling Swan, Cygnus columbianus, is reported. Many of the features common to avian mitochondrial genomes are present in C. columbianus and are described here. The gene order is the same as in Gallus gallus. The sequence of this mitochondrial genome allows relationships within the family Anatidae (swans, geese and ducks) to be reconsidered in the light of a large suite of mitochondrial characters. Protein-coding gene sequences of C. columbianus were concatenated to form a supergene, which was analyzed phylogenetically with similar constructs from previously published avian genomes. Relationships within Anatidae and between the Anatidae and the galliform birds were addressed. Three independent phylogenetic methods confirmed traditional classifications and the existence of the Galloanseres clade.     

The Craniolingual Morphology of Waterfowl (Aves,Anseriformes) and Its Relationship with Feeding Mode Revealed Through Contrast-Enhanced X-Ray Computed Tomography and 2D Morphometrics

Within Anseriformes, waterfowl (ducks, geese, and swans) exhibit three specialized feeding modes that are distinctive among Aves: filter-feeding with fine and dense keratinous lamellae on a flat, mediolaterally expanded bill; cropping or grazing vegetation with large and robust lamellae with a dorsoventrally expanded bill; and sharp lamellae associated with a narrow bill used in acquiring mixed invertebrates and fish underwater mainly by grasping. Here we assess morphometric variation in cranial and hyolingual structures as well as hyolingual myology in a diverse sample of Anatidae to explore the relationship of tongue variation and feeding mode. Phylogenetically informed principal component analysis (phyl.PCA) of cranial-lingual measurements for 67 extant and two extinct anatids recovers grazers and filter-feeding taxa in largely distinct areas of morphospace, while underwater graspers and other mixed feeders show less distinct clustering. The relationship between morphometric differences in skeletal features and muscular variation was further explored through a reassessment of hyolingual musculature enabled by contrast-enhanced X-ray computed tomography (CT) imagery acquired from three exemplar species (Branta canadensis, Chen caerulescens, and Aythya americana) with distinctive ecologies and morphologies of the bony hyoid. Data for these duck and geese exemplars reveal further significant, and previously unstudied, morphological differences between filter-feeding and grazing species. Grazers have a larger hyolingual apparatus with highly-developed extrinsic hyoid muscles; while filter-feeding species are characterized by relatively more diminutive extrinsic hyoid muscles and larger intrinsic hyoid muscles. The feeding modes of two extinct taxa (i.e., Presbyornis and Thambetochen) were also estimated from morphometric data. The results indicate a derived terrestrial browsing or grazing ecology for Thambetochen but do not unequivocally support a specialized filter-feeding ecology for Presbyornis, which is recovered with mixed feeders including swans. The combination of detailed, CT-mediated acquisition of fine muscular anatomy with morphometric approaches shows promise for illuminating form–function relationships in extant taxa more generally.     

Ossification sequence of the avian order anseriformes,with comparison to other precocial birds

Ossification sequences are poorly known for most amniotes, and yet they represent an important source of morphogenetic, phylogenetic, and life history information. Here, the author describes the ossification sequences of three ducks, the Common Eider Somateria mollissima dresseri, the Pekin Duck Anas platyrhynchos, and the Muscovy Duck Cairina moschata. Sequence differences exist both within and among these species, but are generally minor. The Common Eider has the most ossified skeleton prior to hatching, contrary to what is expected in a subarctic migrant species. This may be attributed to a tradeoff between growth rate and locomotory performance. Growth rate is higher in hatchlings with more cartilaginous skeletons, but this may compromise locomotion. No major ossification sequence differences were observed in the craniofacial skeleton when compared with Galliformes, which suggests that the influence of adult morphology on ossification sequence might be relatively minor in many taxa. Galliformes and Anseriformes, while both highly ossified at hatching, differ in the location of their late‐stage ossification centers. In Anseriformes, these are most often located in the appendicular skeleton, whereas in Galliformes they are in the thoracic region and form the ventilatory apparatus. J. Morphol., 2008. © 2008 Wiley‐Liss, Inc.     

A new genus of mantidflies discovered in the Oriental region,with a higher‐level phylogeny of Mantispidae (Neuroptera) using DNA sequences and morphology

A remarkable new genus and two new species of Mantispidae (Neuroptera) are described from the Oriental region. Allomantispa Liu, Wu, Winterton & Ohl gen.n. , currently including A. tibetana Liu, Wu & Winterton sp.n. and A. mirimaculata Liu & Ohl sp.n. The new genus is placed in the subfamily Drepanicinae based on a series of morphological characteristics and on the results of total evidence phylogenetic analyses. Bayesian and Parsimony analyses were undertaken using three gene loci (CAD, 16S rDNA and COI) combined with 74 morphological characters from living and fossil exemplars of Mantispidae (17 genera), Rhachiberothidae (two genera) and Berothidae (five genera), with outgroup taxa from Dilaridae and Osmylidae. The resultant phylogeny presented here recovered a monophyletic Mantispidae with ?Mesomantispinae sister to the rest of the family. Relationships among Mantispidae, Rhachiberothidae and Berothidae support Rhachiberothidae as a separate family sister to Mantispidae. Within Mantispidae, Drepanicinae are a monophyletic clade sister to Calomantispinae and Mantispinae. In a combined analysis, Allomantispa gen.n. was recovered in a clade comprising Ditaxis McLachlan from Australia, and two fossil genera from the Palaearctic, ?Promantispa Panfilov (Kazakhstan; late Jurassic) and ?Liassochrysa Ansorge & Schlüter (Germany; Jurassic), suggesting a highly disjunct and relictual distribution for the family. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:464B06E8‐47E6‐482E‐8136‐83FE3B2E9D6B .     

A phylogenetic test of the calcichordate scenario

The calcichordate scenario of Jefferies and colleagues purports to explain the origin and early evolution of the phyla Echinodermata and Chordata. Calcichordate proponents have argued that echinoderms are the sister group of the chordates and urochordates are the sister group of the craniates. These phylogenetic hypotheses, which differ from the traditional groupings, are derived primarily from morphological interpretations of carpoids (solutes, cornutes, and mitrates), an enigmatic fossil group usually held to be primitive stem-group echinoderms. Although the scenario has received only limited support, it has yet to be falsified. The difficulty with falsifying the calcichordate scenario is proving that the morphological interpretations, for example, that carpoids possessed notochords, dorsal hollow nerve cords, and other typical chordate or craniate characters, are incorrect. Here, rather than argue over the interpretation of fossils, the phylogenetic hypotheses embedded within the scenario are tested. It is found that the calcichordate scenario fails such a test, even if both the Recent and fossils forms are coded according to the calcichordate scenario. It is argued that: (1) the erection of scenarios must follow the construction of a cladogram; and (2) fossils are unable to dictate the relationships among phyla. □ Calcichordate scenario, Carpoidea, Deuterostomia, Echinodermata, Chordata, phylogeny, cladistics.