Higher-order phylogeny of modern birds (Theropoda, Aves: Neornithes) based on comparative anatomy. II. Analysis and discussion

In recent years, avian systematics has been characterized by a diminished reliance on morphological cladistics of modern taxa, intensive palaeornithogical research stimulated by new discoveries and an inundation by analyses based on DNA sequences. Unfortunately, in contrast to significant insights into basal origins, the broad picture of neornithine phylogeny remains largely unresolved. Morphological studies have emphasized characters of use in palaeontological contexts. Molecular studies, following disillusionment with the pioneering, but non-cladistic, work of Sibley and Ahlquist, have differed markedly from each other and from morphological works in both methods and findings. Consequently, at the turn of the millennium, points of robust agreement among schools concerning higher-order neornithine phylogeny have been limited to the two basalmost and several mid-level, primary groups. This paper describes a phylogenetic (cladistic) analysis of 150 taxa of Neornithes, including exemplars from all non-passeriform families, and subordinal representatives of Passeriformes. Thirty-five outgroup taxa encompassing Crocodylia, predominately theropod Dinosauria, and selected Mesozoic birds were used to root the trees. Based on study of specimens and the literature, 2954 morphological characters were defined; these characters have been described in a companion work, approximately one-third of which were multistate (i.e. comprised at least three states), and states within more than one-half of these multistate characters were ordered for analysis. Complete heuristic searches using 10 000 random-addition replicates recovered a total solution set of 97 well-resolved, most-parsimonious trees (MPTs). The set of MPTs was confirmed by an expanded heuristic search based on 10 000 random-addition replicates and a full ratchet-augmented exploration to ascertain global optima. A strict consensus tree of MPTs included only six trichotomies, i.e. nodes differing topologically among MPTs. Bootstrapping (based on 10 000 replicates) percentages and ratchet-minimized support (Bremer) indices indicated most nodes to be robust. Several fossil Neornithes (e.g. Dinornithiformes, Aepyornithiformes) were placed within the ingroup a posteriori either through unconstrained, heursitic searches based on the complete matrix augmented by these taxa separately or using backbone-constraints. Analysis confirmed the topology among outgroup Theropoda and achieved robust resolution at virtually all levels of the Neornithes. Findings included monophyly of the palaeognathous birds, comprising the sister taxa Tinamiformes and ratites, respectively, and the Anseriformes and Galliformes as monophyletic sister-groups, together forming the sister-group to other Neornithes exclusive of the Palaeognathae (Neoaves). Noteworthy inferences include: (i) the sister-group to remaining Neoaves comprises a diversity of marine and wading birds; (ii) Podicipedidae are the sister-group of Gaviidae, and not closely related to the Phoenicopteridae, as recently suggested; (iii) the traditional Pelecaniformes, including the shoebill (Balaeniceps rex) as sister-taxon to other members, are monophyletic; (iv) traditional Ciconiiformes are monophyletic; (v) Strigiformes and Falconiformes are sister-groups; (vi) Cathartidae is the sister-group of the remaining Falconiformes; (vii) Ralliformes (Rallidae and Heliornithidae) are the sister-group to the monophyletic Charadriiformes, with the traditionally composed Gruiformes and Turniciformes (Turnicidae and Mesitornithidae) sequentially paraphyletic to the entire foregoing clade; (viii) Opisthocomus hoazin is the sister-taxon to the Cuculiformes (including the Musophagidae); (ix) traditional Caprimulgiformes are monophyletic and the sister-group of the Apodiformes; (x) Trogoniformes are the sister-group of Coliiformes; (xi) Coraciiformes, Piciformes and Passeriformes are mutually monophyletic and closely related; and (xii) the Galbulae are retained within the Piciformes. Unresolved portions of the Neornithes (nodes having more than one most-parsimonious solution) comprised three parts of the tree: (a) several interfamilial nodes within the Charadriiformes; (b) a trichotomy comprising the (i) Psittaciformes, (ii) Columbiformes and (iii) Trogonomorphae (Trogoniformes, Coliiformes) + Passerimorphae (Coraciiformes, Piciformes, Passeriformes); and (c) a trichotomy comprising the Coraciiformes, Piciformes and Passeriformes. The remaining polytomies were among outgroups, although several of the highest-order nodes were only marginally supported; however, the majority of nodes were resolved and met or surpassed conventional standards of support. Quantitative comparisons with alternative hypotheses, examination of highly supportive and diagnostic characters for higher taxa, correspondences with prior studies, complementarity and philosophical differences with palaeontological phylogenetics, promises and challenges of palaeogeography and calibration of evolutionary rates of birds, and classes of promising evidence and future directions of study are reviewed. Homology, as applied to avian examples of apparent homologues, is considered in terms of recent theory, and a revised annotated classification of higher-order taxa of Neornithes and other closely related Theropoda is proposed. (c) 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 149, 1-95.     

A phylogenetic analysis of basal Anseriformes, the fossil Presbyornis, and the interordinal relationships of waterfowl

A phylogenetic analysis of 123 morphological characters of basal waterfowl (Aves: Anseriformes) and other selected avian orders confirmed that the screamers (Anhimae: Anhitn-idae) are the sister-group of other waterfowl (Anseres), and that the magpie goose (Anseranatidae: Anseranas semipalmata) is the sister group of other modern waterfowl exclusive of screamers (Anatidae sensu stricto). The analysis also supports the traditional hypothesis of the gallinaceous birds (Galliformes) as the sister group of the Anseriformes. Presbyornis, a fossil from the early Eocene of Wyoming and averred by Olson & Feduccia as showing that the Anseriformes were derived from shorebirds (Charadriiformes), was found to represent the sister group of the Anatidae. Associated hypotheses by Olson & Feduccia concerning the implications of Presbyornis for the phylogenetic relationships of flamingos (Phoenicopteriformes), the position of the Anhimidae within the waterfowl, relationships among modern Anatidae, and a plausible evolutionary scenario for waterfowl also are rejected. Analyses revealed that cranial characters were critical to the establishment of the Galliformes as the sister group of the Anseriformes; exclusion of the Anhimidae, especially in combination with Anseranas, also undermined the support for this inference. Placement of Presbyornis as the sister group of the Anatidae casts doubt on the role suggested by Feduccia of ‘transitional shorebirds' in the origin of modern avian orders, and calls into question the concept of ‘fossil mosaics’. The phylogenetic hypothesis is used to reconstruct an evolutionary scenario for selected ecomorphological characters in the galliform-anseriform transition, to predict the most parsimonious states of these characters for Presbyornis, and to propose a phylogenetic classification of the higher-order taxa of waterfowl. This re-examination of Presbyornis also is used to exemplify the fundamental methodological shortcomings of the intuitive approach to the reconstruction of phylogenetic relationships.     

Comparative embryonic development of the skeleton of the domestic turkey (Meleagris gallopavo) and other galliform birds

Ossification sequences are poorly known for birds in general, even for common domestic and experimental species. Such sequences constitute a rich source of data on character evolution, and may even provide phylogenetic information. It is not clear, however, what factors influence ossification sequences and what the relative importance of phylogeny is to the sequences. Galliformes constitute a good group to examine these variables. These birds are osteologically conservative, have precocial young, but have a broad spectrum of body sizes and incubation periods. Here, I describe the embryonic ossification of the skeleton in the domestic turkey (Meleagris gallopavo), and compare it to the domestic chicken (Gallus gallus) and the Japanese quail (Coturnix coturnix). Ossification sequences in this group are not affected by egg size or incubation period. They also appear to be independent of both the spatial location and the embryonic tissue from which the osteogenic cells originated. Accumulation of a wider sample of ossification sequences from more morphologically variable avian taxa will be necessary in order to test functional and phylogenetic effects.     

Phylogenetic relationships among the phyllotini (rodentia: Sigmodontinae) using morphological characters

Thirty-three species representing all 14 genera of the South American rodent tribe Phyllotini and 5 problematic genera are surveyed for 96 multistate and binary dental, cranial, skeletal, external, and male reproductive tract characters. Wagner parsimony analysis confirmsCalomys as the most basal phyllotine genus, and as currently constituted it is likely paraphyletic. The results are consistent with the exclusion ofPseudoryzomys from the phyllotines and the separation ofReithrodon andNeotomys fromHolochilus at the tribal level. Several highly differentiated generic groups that include a radiation of altiplano endemics centered onAuliscomys and the largely southern Andean/PatagonianReithrodon group appear to form a clade. AGraomys generic group that includesAndalgalomys andEligmodontia is also apparent, but its relationships to other phyllotines are obscured by poorly resolved internal nodes in the more species-rich and probably paraphyletic genusPhyllotis. The significance and consequences of more intensive taxonomic sampling are discussed. The taxonomic consequences of the phylogeny are presented.     

Circumscription of Malvaceae (Malvales) as determined by a preliminary cladistic analysis of morphological, anatomical, palynological, and chemical characters

We report a phylogenetic analysis of “core” Malvales (Tiliaceae, Sterculiaceae, Bombacaceae, and Malvaceae) based on morphological, anatomical, palynological, and chemical features. The results of the analyses lead to the conclusion that Tiliaceae, Sterculiaceae, and Bombacaceae, as variously delimited, are paraphyletic; only the Malvaceae are likely monophyletic. The genera of “core” Malvales form a well-defined clade. Genera of “Tiliaceae” constitute the basal complex within “core” Malvales. The “Sterculiaceae” (most genera)+ “Bombacaceae” + Malvaceae form a clade on the basis of a monadelphous androecium; “Bombacaceae”+ Malvaceae also form a clade, which is diagnosable on the basis of monoloculate anthers. It is clear that the traditional classification, with its arbitrarily delimited evolutionary grades, is unsatisfactory, especially if one seeks to reflect phylogeny accurately. Thus, Malvaceae is redefined to refer to the most recent common ancestor of plants previously considered to be “Tiliaceae,” “Sterculiaceae,” “Bombacaceae,” and Malvaceae, and all of the descendants of that ancestor. This broadly circumscribed Malvaceae can be diagnosed by several presumed synapomorphies, but we draw special attention to the unusual floral nectaries that are composed of densely packed, multicellular, glandular hairs on the sepals (or less commonly on the petals or androgynophore).     

Palaeomorphology: fossils and the inference of cladistic relationships

Edgecombe, G.D. 2010. Palaeomorphology: fossils and the inference of cladistic relationships. —Acta Zoologica (Stockholm) 91 : 72–80 Twenty years have passed since it was empirically demonstrated that inclusion of extinct taxa could overturn a phylogenetic hypothesis formulated upon extant taxa alone, challenging Colin Patterson’s bold conjecture that this phenomenon ‘may be non‐existent’. Suppositions and misconceptions about missing data, often couched in terms of ‘wildcard taxa’ and ‘the missing data problem’, continue to cloud the literature on the topic of fossils and phylogenetics. Comparisons of real data sets show that no a priori (or indeed a posteriori) decisions can be made about amounts of missing data and most properties of cladograms, and both simulated and real data sets demonstrate that even highly incomplete taxa can impact on relationships. The exclusion of fossils from phylogenetic analyses is neither theoretically nor empirically defensible.     

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.     

Phylogenetic relationships of Lecythidaceae: a cladistic analysis using rbcL sequence and morphological data

This study examined in detail the rbcL sequence and morphological support for subfamilial relationships and monophyly of Lecythidaceae. Initially we needed to establish relationships of Lecythidaceae among other dicot families. To complete this we examined 47 rbcL sequences of 25 families along with molecular observations from several large analyses of rbcL data. All analyses strongly support the monophyly of the asterid III grouping. This analysis revealed Lecythidaceae to be paraphyletic and indicated potential outgroup relationships with Sapotaceae. Once relationships had been evaluated using molecular data we then concentrated on analyzing separate and combined morphological and molecular databases. The topology of the morphological data set was similar to the rbcL sequence and combined data sets except for the positioning of Napoleonaeoideae, Grias, Gustavia, and Oubanguia. According to the combined results, Planchonioideae, Lecythidoideae. and Foetidioideae are monophyletic, whereas the subfamily Napoleonaeoideae are paraphyletic. Nested within Napolconaeoideae, we found Asteronthos forms a strongly supported clade with Oubanguia (Scytopetalaceae). Foetidia, the only genus of Foetidioideae, is sister to Planchonioideae, and this clade is sister to Lecythidoideae. The [(Planchonioideae, Foetidioideae) Lecythidoideae are sister to Asteranthos/Oubanguia. Napoleonaeoideae are sister to the rest of Lecythidaceae.     

Global diversity of freshwater birds (Aves)

Among the 10,000 birds species living on earth, 5% (e.g., 560) need imperatively freshwater habitat in order to satisfy at least one of their life history traits. About 11 completed families could even disappear if their wetland habitat left. About 10% (58) of these can be considered as endemic. Africa contains the biggest number of endemic (20) and more precisely Madagascar. Among freshwater species, ducks and geese have a major importance in human activities in northern hemisphere related to food resources (hunting) or birding. Guest editors: E.V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Evolutionary relationships of euthyneuran gastropods (Mollusca): a cladistic re-evaluation of morphological characters

Morphological characters of the Euthyneura available from the literature were re-evaluated in terms of terminology and primary homology. A total of 77 characters and 75 taxa were retained in a data matrix. Several assumptions on character weights and types were tested. In the cladistic analyses, it appeared that the data matrix was highly homoplastic, and only robust nodes (those which were little modified by variations in weight and coding of characters) were retained in a concensus tree. The evolutionary histories of all characters and monophylies of higher euthyneuran taxa were discussed. The following interrelationships of the taxa were obtained in a consensus tree: the clade Heterobranchia includes paraphyletic allogastropod taxa which emerge basally, and the clade Euthyneura. The latter includes the clade Pulmonata and at least 10 opisthobranch clades of unresolved relationship (Thecosomata, Gymnosomata, Acochlidioidea, Pyramidelloidea, Runcinoidea, Cephalaspidea, Sacoglossa, Umbraculoidea, Pleurobranchoidea, Nudibranchia). The Pulmonata include basommatophoran paraphyletic taxa and the clade Geophila (Onchidiidae, Soleolifera, Stylommatophora). The position of the Sacoglossa and the monophyly of the Notaspidea are also discussed.  © 2002 The Linnean Society of London, Zoological Journal of the Linnean Society , 2002, 135 , 403–470.     

Phylogeny and classification of Aedini (Diptera: Culicidae), based on morphological characters of all life stages

Higher‐level relationships within Aedini, the largest tribe of Culicidae, are explored using morphological characters of eggs, fourth‐instar larvae, pupae, and adult females and males. In total, 172 characters were examined for 119 exemplar species representing the existing 12 genera and 56 subgenera recognized within the tribe. The data for immature and adult stages were analysed separately and in combination using equal (EW) and implied weighting (IW). Since the classification of Aedini is based mainly on adult morphology, we first tested whether adult data alone would support the existing classification. Overall, the results of these analyses did not reflect the generic classification of the tribe. The tribe as a whole was portrayed as a polyphyletic assemblage of Aedes and Ochlerotatus within which eight (EW) or seven (IW) other genera were embedded. Strict consensus trees (SCTs) derived from analyses of the immature stages data were almost completely unresolved. Combining the adult and immature stages data resulted in fewer most parsimonious cladograms (MPCs) and a more resolved SCT than was found when either of the two data subsets was analysed separately. However, the recovered relationships were still unsatisfactory. Except for the additional recovery of Armigeres as a monophyletic genus, the groups recovered in the EW analysis of the combined data were those found in the EW analysis of adult data. The IW analysis of the total data yielded eight MPCs consisting of three sets of two mutually exclusive topologies that occurred in all possible combinations. We carefully studied the different hypotheses of character transformation responsible for each of the alternative patterns of relationship but were unable to select one of the eight MPCs as a preferred cladogram. Overall, the relationships within the SCT of the eight MPCs were a significant improvement over those found by equal weighting. Aedini and all existing genera except Ochlerotatus and Aedes were recovered as monophyletic. Ochlerotatus formed a polyphyletic assemblage basal to Aedes. This group included Haemagogus and Psorophora, and also Opifex in a sister‐group relationship with Oc. (Not.) chathamicus. Aedes was polyphyletic relative to seven other genera, Armigeres, Ayurakitia, Eretmapodites, Heizmannia, Udaya, Verrallina and Zeugnomyia. With the exception of Ae. (Aedimorphus), Oc. (Finlaya), Oc. (Ochlerotatus) and Oc. (Protomacleaya), all subgenera with two or more species included in the analysis were recovered as monophyletic. Rather than leave the generic classification of Aedini in its current chaotic state, we decided a reasonable and conservative compromise classification would be to recognize as genera those groups that are ‘weighting independent’, i.e. those that are common to the results of both the EW and IW analyses of the total data. The SCT of these combined analyses resulted in a topology of 29 clades, each comprising between two and nine taxa, and 30 taxa (including Mansonia) in an unresolved basal polytomy. In addition to ten genera (Armigeres, Ayurakitia, Eretmapodites, Haemagogus, Heizmannia, Opifex, Psorophora, Udaya, Verrallina and Zeugnomyia), generic status is proposed for the following: (i) 32 existing subgenera of Aedes and Ochlerotatus, including nine monobasic subgenera within the basal polytomy, i.e. Ae. (Belkinius), Ae. (Fredwardsius), Ae. (Indusius), Ae. (Isoaedes), Ae. (Leptosomatomyia), Oc. (Abraedes), Oc. (Aztecaedes), Oc. (Gymnometopa) and Oc. (Kompia); (ii) three small subgenera within the basal polytomy that are undoubtedly monophyletic, i.e. Ae. (Huaedes), Ae. (Skusea) and Oc. (Levua), and (iii) another 20 subgenera that fall within the resolved part of the SCT, i.e. Ae. (Aedes), Ae. (Alanstonea), Ae. (Albuginosus), Ae. (Bothaella), Ae. (Christophersiomyia), Ae. (Diceromyia), Ae. (Edwardsaedes), Ae. (Lorrainea), Ae. (Neomelaniconion), Ae. (Paraedes), Ae. (Pseudarmigeres), Ae. (Scutomyia), Ae. (Stegomyia), Oc. (Geoskusea), Oc. (Halaedes), Oc. (Howardina), Oc. (Kenknightia), Oc. (Mucidus), Oc. (Rhinoskusea) and Oc. (Zavortinkius). A clade consisting of Oc. (Fin.) kochi, Oc. (Fin.) poicilius and relatives is raised to generic rank as Finlaya, and Downsiomyia Vargas is reinstated from synonymy with Finlaya as the generic name for the clade comprising Oc. (Fin.) leonis, Oc. (Fin.) niveus and their relatives. Three other species of Finlaya?Oc. (Fin.) chrysolineatus, Oc. (Fin.) geniculatus and Oc. (Fin.) macfarlanei? fall within the basal polytomy and are treated as Oc. (Finlaya) incertae sedis. Ochlerotatus (Ochlerotatus) is divided into three lineages, two of which, Oc. (Och.) atropalpus and Oc. (Och.) muelleri, are part of the basal polytomy. The remaining seven taxa of Oc. (Ochlerotatus) analysed, including the type species, form a reasonably well‐supported group that is regarded as Ochlerotatus s.s. Ochlerotatus (Rusticoidus) is retained as a subgenus within Ochlerotatus s.s. Ochlerotatus (Nothoskusea) is recognized as a subgenus of Opifex based on two unique features that support their sister‐group relationship. A new genus, Tanakaius gen. nov. , is proposed for Oc. (Fin.) togoi and the related species Oc. (Fin.) savoryi. The taxonomic status and generic placement of all currently valid species of Aedini are listed in an appendix. © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 142 , 289?368.     

Phylogenetic relationships in the tribe Oxyptilini (Lepidoptera,Pterophoridae, Pterophorinae) based on morphological data of adults

The monophyly of the tribe Oxyptilini and phylogenetic relationships of the genera embraced in this tribe were examined using 171 (75 binary and 96 multistate) characters of adult morphology. The study material included 98 species of 30 genera, representing all previously recognized genera of Oxyptilini, together with the genera Sphenarches, Antarches, Diacrotricha, and Cosmoclostis, four species of Oidaematophorini, three species of Platyptiliini, as well as three and two other species belonging to Pterophorini and Exelastini respectively. Two Agdistis species were used as outgroups. The cladistic analysis resulted in six equally parsimonious trees. A majority of the recovered synapomorphic characters have previously been used in the taxonomy of the subfamily. However, 25 novel characters were found. The monophyly of Oxyptilini was supported, although only with homoplastic characters and low amounts of tree confidence; the genera Capperia, Procapperia, Paracapperia, Oxyptilus, Megalorhipida, and Trichoptilus were found to be nonmonophyletic; Sphenarches and Antarches were recovered as members of Oxyptilini; the two genera Cosmoclostis and Diacrotricha were placed out of Oxyptilini, inside the tribe Pterophorini; and close affinity of the genus Dejongia to Stangeia, Stenodacma, Megalorhipida, Trichoptilus, and Buckleria species was revealed. Four new combinations, Cosmoclostis lanceata (Arenberger) comb. nov. , Nippoptilia regulus (Meyrick) comb. nov. , Capperia tadzhica (Zagulajev) comb. nov. , and Buckleria negotiosus (Meyrick) comb. nov. are proposed; Capperia insomnis Townsend was considered as a senior synonym of Procapperia hackeri Arenberger syn. nov. , Buckleria negotiosus (Meyrick) as a senior synonym of Buckleria vanderwolfi Gielis syn. nov. , and Oxyptilus variegatus Meyrick syn. nov. as a junior synonym of Oxyptilus secutor Meyrick. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 484–547.     

The evolutionary radiation of modern birds (Neornithes): reconciling molecules, morphology and the fossil record

The pattern, timing and extent of the evolutionary radiation of anatomically modern birds (Neornithes) remains contentious: dramatically different timescales for this major event in vertebrate evolution have been recovered by the 'clock-like' modelling of molecular sequence data and from evidence extracted from the known fossil record. Because current synthesis would lead us to believe that fossil and nonfossil evidence conflict with regard to the neornithine timescale, especially at its base, it is high time that available data are reconciled to determine more exactly the evolutionary radiation of modern birds. In this review we highlight current understanding of the early fossil history of Neornithes in conjunction with available phylogenetic resolution for the major extant clades, as well as recent advancements in genetic methods that have constrained time estimates for major evolutionary divergences. Although the use of molecular approaches for timing the radiation of Neornithes is emphasized, the tenet of this review remains the fossil record of the major neornithine subdivisions and better-preserved taxa. Fossils allowing clear phylogenetic constraint of taxa are central to future work in the production of accurate molecular calibrations of the neornithine evolutionary timescale.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 141 , 153–177.     

Sylviornis neocaledoniae (Aves,Galliformes, Megapodiidae), oiseau Géant éteint de l'ile des Pins (Nouvelle-Calédonie)

Sylviornis neocaledoniae is represented in the sites of Kanumera, Isle of Pines, by a few cranial elements (palatines, quadrates, quadratojugal) but mainly by post-cranial remains. The fact that palatines are fused in the sagittal plane proves the palate of Sylviornis to be neognathous. The flightless bird's post-cranial skeleton is highly modified, but some anatomical characters show it can be assigned to Galliformes, and, amongst them, is closer to recent or extinct Megapodes than to any other family, Sylviornis neocaledoniae was probably exterminated by man who preserved its mythological memory under the name of Du.     

Phylogenetic relationships in Magnoliaceae subfam. Magnolioideae: a morphological cladistic analysis

Relationships within Magnolioideae have been the subject of persistent debate; the main point at issue mostly being the disposition of tribes, genera and sections. A morphological cladistic analysis of the subfamily using Liriodendron as the out-group showed that Magnolioideae consisted of a large basal polytomy, but with five resolved and variously supported clades. Manglietia constituted a clade with sect. Rytidospermum of Magnolia subg. Magnolia. Kmeria and Woonyoungia formed a pair. Pachylarnax, Parakmeria and Manglietiastrum were grouped together, and sect. Splendentes and Dugandiodendron also formed a pair. The largest and best supported clade consisted of Magnolia subg. Magnolia sects. Oyama and Maingola, Magnolia subg. Yulania, Michelia, Aromadendron, Alcimandra, Elmerrillia, Paramichelia and Tsoongiodendron, with sect. Oyama of Magnolia subg. Magnolia is sister to the remainder. Although Magnolia sect. Maingola, Aromadendron, Alcimandra and Elmerrillia constituted a poorly resolved subclade, Aromadendron formed a monophyletic clade with Alcimandra. Within the Michelia/Magnolia subgen. Yulania subclade, Paramichelia was sister to Tsoongiodendron. These results are supported by similar placement of taxa within various molecular analyses of the family, but the low level of resolution indicates that more morphological data are needed to improve phylogenetic signal. Our results support the molecular analyses in suggesting that Magnolia is best considered to be a large and diverse genus, but that the relationships between the taxa within it require more detailed clarification, with more extensive sampling and a combined molecular and morphological approach being needed.     

Systematic relationships within the genus Trichomanes sensu lato (Hymenophyllaceae, Filicopsida): cladistic analysis based on anatomical and morphological data

Systematics within the genus Trichomanes sensu lato (Hymenophyllaceae, Filicopsida) have continued to be controversial because of the difficulty in identifying homologies and informative characters within high morphological diversity. Systematic relationships are investigated in this study by using a cladistic approach with 31 anatomical and morphological characters from 20 taxa which correspond to the sections defined by Morton. The results broadly confirm Morton's four subgenera: Trichomanes, Didymoglossum, Pachychaetum and Achomanes. The monophyly of Pachychaetum remains fragile and the section Lacostea appears to be excluded from its traditional subgenus (Achomanes). In addition, by producing two major 'ecological' clades (terrestrial and epiphytic), the analysis shows that several selected characters appear to be strongly linked with the ecology which may have a significant influence on the topology.     

Revision and cladistic analysis of Isoctenus and description of a new neotropical genus (Araneae,Ctenidae, Cteninae)

A cladistic analysis was applied to test the monophyly of the genus Isoctenus. The data matrix comprised 28 taxa scored for 53 morphological and two behavioural characters. The analysis resulted in two equally parsimonious trees of 89 steps. The strict consensus was used to discuss the relationships of Isoctenus and related Cteninae genera. Ctenopsis Schmidt is synonymized with Isoctenus. Isoctenus foliifer Bertkau, I. strandi Mello‐Leitão, I. eupalaestrus Mello‐Leitão, I. janeirus (Walckenaer), I. coxalis (Pickard‐Cambridge), I. corymbus Polotow, Brescovit & Pellegatti‐Franco and I. malabaris Polotow, Brescovit & Ott are maintained in Isoctenus. Four species currently included in Ctenus are transferred to Isoctenus: I. griseolus (Mello‐Leitão) comb. nov., I. taperae (Mello‐Leitão) comb. nov., I. herteli (Mello‐Leitão) comb. nov. and I. minusculus (Keyserling) comb. nov. The following specific names are synonymized: Ctenus sanguineus Walckenaer, C. semiornatus Mello‐Leitão and Ctenopsis stellata Schmidt with Isoctenus janeirus (Walckenaer), Ctenus mourei Mello‐Leitão with Isoctenus herteli (Mello‐Leitão) and Ctenus pauper Mello‐Leitão with Isoctenus strandi Mello‐Leitão. Isoctenus sigma Schenkel, described from French Guiana, is transferred to Ctenus. Four species are newly described: Isoctenus areia sp. nov. from Paraíba, Brazil, I. charada sp. nov. and I. segredo sp. nov. from Paraná, Brazil, and I. ordinario sp. nov. from south and south‐eastern Brazil and north‐eastern Argentina. Isoctenus latevittatus Caporiacco is considered species inquirenda. Parabatinga gen. nov. is proposed to include Ctenus brevipes Keyserling. The following synonymies are established: Ctenus taeniatus Keyserling, C. tatarandensis Tullgren, C. anisitsi Strand, C. atrivulvus Strand, C. mentor Strand, C. brevipes brevilabris Strand, Isoctenus masculus Mello‐Leitão and Ctenus birabeni Mello‐Leitão with Parabatinga brevipes (Keyserling) comb. nov. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 583–614.     

Relationships, classification and evolution of Scleranthus (Caryophyllaceae) as inferred from analysis of morphological characters

Study of floral anatomy, micromorphology, palynology and onotogeny has revealed new characters for phylogenetic analysis in the genus Scleranthus . Cladistic analysis of these characters, along with those previously available, suggests that the genus consists of Eurasian/Mediterranean and Australasian sister clades. Gynoecial morphology and development are closely similar in all species, suggesting the genus is monophyletic despite its disjunct northern and southern hemisphere distribution. Variation in pollen:ovule ratios and their implications for the evolution of Scleranthus species are also discussed and it is concluded that a range of breeding strategies intermediate between autogamy and xenogamy exists in the genus. Single-stamened species of Scleranthus are likely to be obligate autogams, despite their comparatively high pollen:ovule ratios in relation to autogamic species of other genera.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 140 , 15–29.     

Increased taxon and character sampling reveals novel intergeneric relationships in the Cracidae (Aves: Galliformes)

The Cracidae is one of the most endangered bird families in the World. Several studies have been published recently on the evolution and conservation of cracids. Phylogenetic analyses using a fragment of 661 bp of the mitochondrial cytochrome b gene for 39 different species of cracids corroborated most relationships found in previous studies. The present work attempts to refine the former phylogenetic hypothesis by increasing taxon sampling and combining molecular with osteological, integumentary and behavioural characters using Maximum Parsimony (MP) and Bayesian analyses. We present both separate and combined total evidence analyses with our molecular data, 152 osteological and 74 integumentary + behavioural characters. While supporting most aspects of the molecular-based hypotheses, the tree based on the combined matrix suggests several modifications of the generic composition for each of the two subfamilies: Penelopinae and Cracinae, and supports the merging of the genera Pipile with Aburria and Mitu with Pauxi . These results suggest that increased taxon and character sampling from a diversity of sources may be at least as important as increased sampling of only one type. Besides, of a total of 891 characters we had 437 parsimony-informative sites (almost half of the analyzable sites) proving the efficiency of a total-evidence approach.     

Phylogeny, classification, and biogeography of the cycloteline Therevinae (Insecta: Diptera: Thereavidae)

Twenty-one members of the Laurasian group of Therevinae (Diptera: Therevidae) are compared using 65 adult morphological characters. Cladistic analysis using parsimony on the 17 ingroup and 4 outgroup taxa provides a well-supported hypothesis of relationships among taxa within the Gyclotelini, tribe nov. The Cyclotelini is a monophyletic assemblage of mostly New World genera, including Anolinga , gen. nov. , Breviperna Irwin, Coleiana , gen. nov. , Crebraseta , gen. nov. , Cyclotelus Walker, Mesonana , gen.nov. , and Ozodiceromyia Bigot. In addition, three Old World genera, Ammothereva Lyneborg, Bugulaverpa , gen. nov. , and Procyclotelus Nagatomi & Lyneborg, are included in the tribe. These ten genera are divided into two monophyletic genus-groups, the Brevipema-group and the Cyclotelus-group. Keys are provided for the genera of Cyclotelini. The tribe, the two informal genus-groups, and all genera are diagnosed; five new genera and six new species are proposed. The biogeographical histories of the genera are discussed in terms of their cladistic relationships using methods of cladistic biogeography. Two major vicariant events account for the current distribution of the tribe. The first relates to the Beringian land bridge connecting western North America and eastern Asia. Second, New World cyclotelines were profoundly affected by the Early Eocene breakup of the archipelagic bridge between North and South America, and the distributions support the hypotheses favouring the continental origin of the Greater Antilles.