The phylogenetic position of Apterosperma (Theaceae) based on morphological and karyotype characters

Classifications of Theaceae have usually placed the endangered monotypic genus Apterosperma in tribe Schimeae (x=18), whereas recent molecular phylogenetic evidence supports its transfer to tribe Theeae (x=15). Molecular data have not resolved the phylogenetic position of Apterosperma within Theeae. We investigated the chromosome number and karyotype of Apterosperma in the context of molecular and morphological phylogenetic evidence to provide further insight into the placement of Apterosperma within Theaceae. The chromosome number and karyotype was found to be 2n = 30 = 26m + 4sm, consistent with the transfer of Apterosperma to tribe Theeae. When the chromosome data were incorporated into a data set of 46 other nonmolecular characters, Apterosperma was placed as the first-diverging lineage within the clade comprising tribe Theeae. This supports its placement based on molecular data. The low intrachromosomal asymmetry (type 1A) of Apterosperma, presumably ancestral for the family, is also consistent with this placement. Character optimization strongly supports a base chromosome number of x=15 for tribe Theeae. Because of variable and sometimes conflicting chromosome count reports of species in tribes Schimeae and Stewartieae, the base chromosome number of Theaceae could be either x=15 or 17.     

Diversification and the adaptive radiation of the vangas of Madagascar

The vangas of Madagascar exhibit extreme diversity in morphology and ecology. Recent studies have shown that several other Malagasy species also are part of this endemic radiation, even as the monophyly of the clade remains in question. Using DNA sequences from 13 genes and representatives of all 15 vanga genera, we find strong support for the monophyly of the Malagasy vangids and their inclusion in a family along with six aberrant genera of shrike-like corvoids distributed in Asia and Africa. Biogeographic reconstructions of these lineages include both Asia and Africa as possible dispersal routes to Madagascar. To study patterns of speciation through time, we introduce a method that can accommodate phylogenetically non-random patterns of incomplete taxon sampling in diversification studies. We demonstrate that speciation rates in vangas decreased dramatically through time following the colonization of Madagascar. Foraging strategies of these birds show remarkable congruence with phylogenetic relationships, indicating that adaptations to feeding specializations played a role in the diversification of these birds. Vangas fit the model of an 'adaptive radiation' in that they show an explosive burst of speciation soon after colonization, increased diversification into novel niches and extraordinary ecomorphological diversity.     

A phylogenetic analysis of the palm subtribe Oncospermatinae (Arecaceae) based on morphological characters

Subtribe Oncospermatinae (Arecaceae: Arecoideae: Areceae) is a diverse group of spiny Old World palms. The subtribe includesOncosperma, a widespread Asian genus of five species, along with seven monotypic genera, all endemic to the Seychelles and Mascarene Islands of the western Indian Ocean. A phylogenetic analysis was conducted in order to test the monophyly of subtribe Oncospermatinae with respect to other Old World genera of tribe Areceae. A matrix of 38 morphological characters was scored for 29 taxa, including 11 species of the Oncospermatinae. A single most parsimonious tree was found, resolving the subtribe as a polyphyletic group of two distinct clades. One clade containingAcanthophoenix, Deckenia, Oncosperma, andTectiphiala was placed as sister to a large group that includes members of subtribes Archontophoenicinae, Arecinae, Iguanurinae, and Ptychospermatinae. The other clade of Oncospermatinae, including the Seychelles endemic generaNephrosperma, Phoenicophorium, Roscheria, andVerschaffeltia, was resolved as sister to the Madagascar endemic subtribe Masoalinae, and may have arisen in the western Indian Ocean region.     

Perenniporia cinereofusca sp. nov. (Polyporales,Basidiomycota) evidenced by morphological characters and phylogenetic analysis

A new Perenniporia species, P. cinereofusca, is described based on morphological and molecular characters. It is characterized by an annual growth habit, resupinate basidiocarps with gray to pale brown pore surface, tissues becoming black in 5% potassium hydroxide (KOH), a dimitic hyphal system with weakly dextrinoid skeletal hyphae and hyaline to pale yellowish, distinctly thick-walled and indextrinoid basidiospores (6.5–7.7 × 5.3–6.3 μm), and presence of dendrohyphidia and large rhomboid crystals. Both morphological and molecular evidence confirmed the placement of the new species in Perenniporia sensu stricto and showed its relationships with similar species in the genus.     

Phylogenetic relationships within Maldanidae (Capitellida,Annelida), based on morphological characters

The Maldanidae annelid worms are reviewed and the phylogenetic relationships of their subgroups provided, based on Hennigian principles and maximum parsimony. Characters were coded as binary and multistate (transformation series). We used 33 terminal taxa (species), and 50 characters. Characters were treated as unordered and of equal weight, and analysis was run in TNT. Three equally most-parsimonious trees were obtained with heuristic searches, with lengths of 64 steps; CI = 0.95, and RI = 0.98. The monophyly of Maldanidae was supported with 100% of bootstrap and jackknife values. As a result of our analysis, Arenicolidae remains the sister-group of Maldanidae, and both should be referred to Maldanomorpha. Maldanidae was supported by the following synapomorphies: dorsal prostomium; prostomium keel-shaped and fused to peristomium; torus globose behind median chaetigers; median chaetigers greatly elongated; number of pre-anal segments reduced. The subfamily Bogueinae was not monophyletic; Boguea and Boguella were included within Rhodininae. Clymenura, previously included in the Clymenurinae, was included within Euclymeninae. The taxa Notoproctinae, Maldaninae, Nicomachinae and Euclymeninae were grouped in the Maldanoplaca, a new taxon. Eight further new clades have been found, but were not named.     

Avian higher-level phylogeny: well-supported clades and what we can learn from a phylogenetic analysis of 2954 morphological characters

It has been shown that increased character sampling betters the accuracy of phylogenetic reconstructions in the case of molecular data. A recently published analysis of avian higher-level phylogenetics based on 2954 morphological characters now provides an empirical example to test whether this is also true in the case of morphological characters. Several clades are discussed which are supported by multiple analyses of mutually independent molecular data (sequences of nuclear genes on different chromosomes and mitochondrial genes) as well as morphological apomorphies, but did not result from parsimony analysis of the large morphological data set. Incorrect character scorings in that analysis notwithstanding, it is concluded that in the case of morphological data, increased character sampling does not necessarily better the accuracy of a phylogenetic reconstruction. Because morphological characters usually have a strongly varying complexity, many simple and homoplastic characters may overrule fewer ones of greater phylogenetic significance in large data sets, thus producing a low ratio of phylogenetic signal to 'noise' in the data.     

Head capsule characters in the Hymenoptera and their phylogenetic implications

The head capsule of a taxon sample of three outgroup and 86 ingroup taxa is examined for characters of possible phylogenetic significance within Hymenoptera. 21 morphological characters are illustrated and scored, and their character evolution explored by mapping them onto a phylogeny recently produced from a large morphological data set. Many of the characters are informative and display unambiguous changes. Most of the character support demonstrated is supportive at the superfamily or family level. In contrast, only few characters corroborate deeper nodes in the phylogeny of Hymenoptera.     

The phylogenetic relationships of the physaloid genera (Solanaceae) based on morphological data

A group of genera, e.g., Chamaesaracha, Leucophysalis, Physaliastrum, Margaranthus, and Withania, in the subfamily Solanoideae (Solanaceae) is centered around the genus Physalis and has been named the physaloid group. It comprises a number of small and often poorly known genera, sometimes seen as united with Physalis and/or each other. A hypothesis of the phylogenetic relationships within this group, based on parsimony analyses of morphological data, is here presented for the first time. The result is discussed in relation to prevailing generic circumsciptions and taxonomic consequences. It is also compared with hypotheses of relationships based on cpDNA data.     

Towards a phylogenetic classification of dendrocoelid freshwater planarians (Platyhelminthes): a morphological and eclectic approach

We explore and review the taxonomic distribution of morphological features that may be used as supporting apomorphies for the monophyletic status of various taxa in future, more comprehensive phylogenetic analyses of the dendrocoelid freshwater planarians and their close relatives. Characters examined are: (i) musculature of the pharynx; (ii) anatomy of the eye; (iii) distribution of testis follicles along the longitudinal axis of the body; (iv) epidermis of the body wall; (v) musculature of the ventral body wall; (vi) presence or absence of a penis papilla; (vii) position of the mouth opening; and (viii) anterior adhesive organ. A phylogenetic hypothesis, in the form of a character state tree, is developed. This hypothesis suggests that the Kenkiidae should be resurrected as a separate family, sharing a sistergroup relationship with the Dendrocoelidae. This sistergroup relationship is supported by the shared presence of an apomorphic anterior adhesive organ. Although the shape of the anterior adhesive organ, both within and between kenkiids and dendrocoelids, may differ considerably between species, they are all constructed according to the same principles. The monophyly of the Kenkiidae is supported by the presence of an apomorphic thickening of the marginal zone of the epidermis and presumably also by the presence of prepharyngeal testes. Within the Kenkiidae, the genus Macrocotyla should be considered as a junior synonym of Kenkia, which is characterized by the presence of an apomorphic ‘planariid pharynx‐type 2’. The monophyly of the Dendrocoelidae is suggested by the presence of a dendrocoelid type of pharynx. It is shown that multicellular eye cups with numerous photoreceptor cells are not restricted to the Dugesiidae and the Terricola but are also characteristic for the Dendrocoelidae; the phylogenetic implications of this character state distribution are discussed. It is pointed out that in planarians there may be a relationship between: (i) body size, the size of an eye cup and the number of its constituent pigment cells; and (ii) the size of an eye cup and the number of photoreceptor cells that it contains. Some tentative phylogenetic groups are suggested on the basis of the following presumed apomorphic characters: presence of transverse fibres in the ventral subepidermal musculature, presence of an extra longitudinal muscle layer in the outer pharynx musculature, a mouth opening that has shifted anteriad. It is suggested that the presumed monophyly of the genus Bdellocephala is supported by the apomorphic absence of a penis papilla, combined with a penial lumen provided with many plicae.     

A phylogenetic study of the Anthozoa (phylum Cnidaria) based on morphological and molecular characters

The phylogenetic relationships within the Anthozoa were re-evaluated based on 41 morphological characters and nuclear sequences of 18S ribosomal DNA (29 anthozoans as ingroups and 3 hydrozoans as outgroups). The parsimony trees derived from the morphological data did not coincide closely with the molecular data, and the presence of several polytomies at some nodes of the trees resulted in ambiguities among the systematic relationships. On the other hand, the combined analysis using total evidence presents a more resolved and highly supported topology, as is indicated by higher bootstrap values and decay indices than either analysis alone. However, strict and semi-strict consensus trees derived from taxonomic congruence show a poorer resolution for the phylogeny of Anthozoa. The trees constructed from the molecular data, using neighbor-joining and maximum-likelihood methods, are nearly congruent with the result from the total evidence. Based on these results, Anthozoa is divided into three subclasses: Alcyonaria, Zoantharia, and Ceriantipatharia. The Ceriantipatharia now includes only one order, Ceriantharia, since the order Antipatharia is more closely related to orders within the Zoantharia. The Alcyonaria is a monophyletic group, in which the order Pennatulacea is basal, and orders Alcyonacea and Telestacea branch later. The order Gorgonacea is divided into two suborders, Holaxonia and Scleraxonia. Bellonela is more related to order Stolonifera, forming a monophyletic group. In Zoantharia, the order Zoanthinaria is basal, and the remaining taxa are divided into two clades: one includes the order Actiniaria and the other includes orders Antipatharia, Corallimorpharia, and Scleractinia. The latter two orders form a monophyletic group. This study presents a different phylogeny of actiniarians from the earlier hypothesis of scleractinian ancestry.     

The phylogenetic affinities of Crossley's babbler (Mystacornis crossleyi): adding a new niche to the vanga radiation of Madagascar

Crossley's babbler (Mystacornis crossleyi) is a passerine endemic to Madagascar. Traditionally, it has been classified as a babbler (Timaliidae), although affinities with warblers and vangas have been suggested. We investigated the phylogenetic affinities of Crossley's babbler using sequence data from two nuclear introns (myoglobin intron 2 and beta-fibrinogen intron 5) and one mitochondrial gene (ND2). We present for the first time (to our knowledge) a molecular phylogeny that confidently places this enigmatic species within the vangas (Vangidae). The inclusion of Crossley's babbler within the vangas adds another foraging niche--gleaning small invertebrates from the ground-to this already large adaptive radiation of songbirds.     

Head structures of Karoophasma sp. (Hexapoda, Mantophasmatodea) with phylogenetic implications

External and internal head structures of adults of Karoophasma sp. were examined and described. The results are compared with conditions found in other representatives of Mantophasmatodea and members of other lower neopteran groups. The X-shaped apodeme of the frons, the unpigmented oval area enclosed by apical branches of the anterior tentorial arms, the oval sclerotisation at the base of the labrum, the sclerotized rounded apical part of the galea, and the loss of M. labroepipharyngalis are probably autapomorphic for Mantophasmatodea. Plesiomorphic features (groundplan of Neoptera) are the orthognathous condition, the absence of parietal ridges, the absence of a gula, the absence of a 'perforation of the corpotentorium', the multisegmented antennae inserted between the compound eyes, the general arrangement of the mouthparts, the shape and composition of the maxillae and labium, and the nearly complete set of muscles. The presence of a transverse muscle connecting the antennal ampullae is a potential synapomorphy of Orthoptera, Phasmatodea and Dictyoptera. Character states suggesting affinities with Grylloblattodea are the absence of ocelli, the elongation of the corpotentorium, and the very similar mandibles with widely separated bases and completely reduced molae. Whether predacious habits are a synapomorphic feature of Mantophasmatodea and Grylloblattodea is uncertain. The retained orthognathous condition in Mantophasmatodea and Mantodea is likely related with different specialized preying techniques in both groups, i.e. rapid forward pushes of the head–prothorax complex, and the use of raptorial legs, respectively.     

Pattern of Pads and Folds in the Foot of European Oscines (Aves, Passeriformes)

A comparative study of pads and folds in the foot is made on 56 passerine species, classified in 18 families or subfamilies of the suborder Oscines. A basic pattern of thirteen pads is recognized: twelve in the digits, each related to one phalanx; the thirteenth in the central part of the foot, ventral to the trochleas of tarsometatarsus. Certain pads in the anterior digits are divided in species with long and narrow pads. The number of folds varies, even in the same species, but a basic pattern is recognized. There is one fold at each joint. Except for the fold at the distal joint in the four digits, these folds are comparable with pads in certain non-Passerifor-mes. The folds at the proximal phalanx of the third and the fourth digit are interpreted as reduced pads. Some folds are separated parts of phalanx pads. – Most passerine birds have this basic pattern of pads and folds, clearly adapted to a life on branches or on twigs. Certain species, exemplified by Alauda arvensis, Plectrophenax nicalis, Certhia familiaris , and Regulus regulus , have different morphology of pads and folds, depending on a reduced number of papillae, and being adaptations to life on the ground, on tree trunks, or on very thin twigs. Relatively few papillae occur in certain species that winter in temperate or cold climates. This means a reduced area of contact with the substrate and is probably important in heat regulation by reducing the conductive heat flow through the papillae to the substrate.–The pads and folds have a restricted value for the classification on family or subfamily level.     

Deep phylogenetic divergence between Scolytoplatypus and Remansus,a new genus of Scolytoplatypodini from Madagascar (Coleoptera,Curculionidae, Scolytinae)

Scolytoplatypodini Blandford is a monotypic tribe of ambrosia beetles found in Asia, Madagascar and Africa. Only three species are currently known from Madagascar and four additional species are here described as new to science. Phylogenetic analyses of morphological and molecular data revealed that four of the seven endemic species are deeply separated from all other species by genetic and distinct morphological characters and therefore placed in a new genus Remansus Jordal. The split between this ancient lineage and Scolytoplatypus Schaufuss was estimated to approximate Palaeocene age (63 Ma), extending the minimum age of ambrosia feeding for this tribe to the beginning of the Palaeocene‒Eocene thermal maximum (PETM). In addition to the ancient origin of Remansus in Madagascar during the Palaeocene, a second origin occurred in Scolytoplatypus no more than 13 Ma. A geographical origin of the latter in South-Eastern Africa was unequivocally inferred from the phylogenies.     

Molecular Systematics of the Malagasy Babblers (Passeriformes: Timaliidae) and Warblers (Passeriformes: Sylviidae), Based on Cytochrome b and 16S rRNA Sequences

The phylogenetic relationships of the Timaliidae (babblers) and Sylviidae (warblers) have long challenged ornithologists. We focus here on three Malagasy genera currently assigned to the Timaliidae, Mystacornis, Oxylabes, and Neomixis, and on their relationships with other babblers and warblers using the sequences of two mitochondrial genes (cytochrome b and 16S rRNA). Maximum parsimony analyses show that the Malagasy “babblers” are not related to any of the other African and Asian babblers. The genus Mystacornis is neither a babbler nor a warbler. The other Malagasy “babblers” are members of warbler groups (the monophyly of the Sylviidae is not demonstrated). Oxylabes madagascariensis and Hartertula flavoviridis (we recognize Hartertula as a genus for the species flavoviridis, previously Neomixis flavoviridis) constitute, with two presumed sylviine taxa, Thamnornis chloropetoides and Cryptosylvicola randrianasoloi, a warbler radiation endemic to the island of Madagascar. The other Neomixis species (tenella, striatigula, and viridis) belong to another warbler group comprising cisticoline taxa. These results show that the Timaliidae did not disperse to Madagascar. Rather, the island has been colonized, independently, by at least two clades of warblers, probably originating from Africa, where the Sylviidae radiation has been the most extensive.     

Cladistic analysis of the Lythraceae sensu lato based on morphological characters

GRAHAM, S. A., CRISCI, J. V. & HOCH, P. C, 1993. Cladistic analysis of the Lythraceae sensu lato based on morphological characters. The Lythraceae s. I. comprise 31 mostly highly distinctive genera distributed worldwide. Cladistic analyses of the family were undertaken using 26 characters from anatomy, floral morphology, pollen, and seed morphology. Of four outgroup hypotheses, the Onagraceae offered the best heuristic assessment, generating 3746 trees with a tree length of 83 and consistency index and (C.I.) of 0.41. A strict consensus tree from the 3746 trees produced a basal monophyletic group of five genera (Sonneratia, Duabanga, Punka, Lagerstroemia, Lawsonia) characterized by anthotelic (determinate) inflorescences and wet stigmas, and a second clade with blastotelic (indeterminate) inflorescences, dry stigmas, and reduced carpel number. Successive weighting with Onagraceae as outgroup generated five most parsimonious trees with a tree length of 164 and C.I. of 0.75. Internal branches are weakly supported by only seven non-homoplasious characters. Significant results of the analysis include: (1) recognition of the two major clades and a total of seven monophyletic groups within the family; (2) the paraphyly of subfamily Lythroideae (Lythraceae s. s .); and (3) indication that genera endemic to the New World have been derived from more than one ancient Old World evolutionary line. The current taxonomic classification of the family is not closely correlated with results of the cladistic analysis.     

A phylogenetic analysis of Pisum based on morphological characters, and allozyme and RAPD markers

Cladistic analyses of 17 wild and cultivated pea taxa were performed using morphological characters, and allozyme and RAPD (random amplified polymorphic DNA) markers. Both branch-and-bound and bootstrap searches produced cladograms that confirmed the close relationships among the wild species and cultivars of Pisum proposed by a variety of systematic studies. Intraspecific rankings were supported for northern P. humile, southern P. humile, P. elatius and P. sativum, which together comprise a single-species complex. P. fulvum, while clearly the most divergent of the pea taxa, could also be assigned to the same species complex without violating the hierarchial logic of the cladogram. Its inclusion or exclusion depends on whether the level of interfertility it displays with other pea taxa or its overall morphological and chromosomal distinction are emphasized. As suggested by previous studies, northern P. humile was the most likely sister taxon to cultivated P. sativum; although, rigorous phylogenetic evaluation revealed a close genealogical affinity among P. elatius, northern P. humile and P. sativum. Despite their limited number, the 16 morphological characters and allozyme markers used precisely organized the pea taxa into established taxonomic groupings, perhaps in part reflecting the role morphology has played historically in pea classification. The RAPD data also generally supported these same groupings and provided additional information regarding the relationships among the taxa. Given that RAPDs are relatively quick and easy to use, are refractory to many environmental influences, can be generated in large numbers, and can complement traditional characters that may be limited in availability, they provide a valuable new resource for phylogenetic studies.     

Determination of phylogenetic position of Pipizini (Diptera: Syrphidae): based on molecular biological and morphological data

Based on the sequence analysis of 5.8S subunit and internal transcribed spacers (ITS ) of ribosomal RNA gene (rDNA), the molecular phylogenetic tree of representative species of Pipizini and three groups of Syrphidae with different feeding habits (seven species belong to six genera) was constructed. Meanwhile, the phylogenetic tree of tribes (including Pipizini and other 17 tribes of Syrphidae) was constructed using morphological characteristics of adults and larvae and the number of chromosomes. Both the results show that the relationship between Pipizini and predatory groups is closer than that between Pipizini and saprophagous groups. So it is suggested that Pipizini be transferred from Milesiinae to Syrphinae.     

A phylogenetic analysis of the Gruiformes (Aves) based on morphological characters,with an emphasis on the rails (Rallidae)

The order Gruiformes, for which even familial composition remains controversial, is perhaps the least well understood avian order from a phylogenetic perspective. The history of the systematics of the order is presented, and the ecological and biogeographic characteristics of its members are summarized. Using cladistic techniques, phylogenetic relationships among fossil and modern genera of the Gruiformes were estimated based on 381 primarily osteological characters; relationships among modern species of Grues (Psophiidae, Aramidae, Gruidae, Heliornithidae and Rallidae) were assessed based on these characters augmented by 189 characters of the definitive integument. A strict consensus tree for 20,000 shortest trees compiled for the matrix of gruiform genera (length = 967, CI = 0.517) revealed a number of nodes common to the solution set, many of which were robust to bootstrapping and had substantial support (Bremer) indices. Robust nodes included those supporting: a sister relationship between the Pedionomidae and Turnicidae; monophyly of the Gruiformes exclusive of the Pedionomidae and Turnicidae; a sister relationship between the Cariamidae and Phorusrhacoidea; a sister relationship between a clade comprising Eurypyga and Messelornis and one comprising Rhynochetos and Aptornis; monophyly of the Grues (Psophiidae, Aramidae, Gruidae, Heliornithidae and Rallidae); monophyly of a clade (Gruoidea) comprising (in order of increasingly close relationship) Psophia, Aramus, Balearica and other Gruidae, with monophyly of each member in this series confirmed; a sister relationship between the Heliornithidae and Rallidae; and monophyly of the Rallidae exclusive of Himantornis. Autapomorphic divergence was comparatively high for Pedionomus, Eurypyga, Psophia, Himantornis and Fulica; extreme autapomorphy, much of which is unique for the order, characterized the extinct, flightless Aptornis. In the species-level analysis of modern Grues, special efforts were made to limit the analytical impacts of homoplasy related to flightlessness in a number of rallid lineages. A strict consensus tree of 20,000 shortest trees compiled (length = 1232, CI = 0.463) confirmed the interfamilial relationships resolved in the ordinal analysis and established a number of other, variably supported groups within the Rallidae. Groupings within the Rallidae included: monophyly of Rallidae exclusive of Himantornis and a clade comprising Porphyrio (including Notornis) and Porphyrula; a poorly resolved, basal group of genera including Gymnocrex, Habroptila, Eulabeornis, Aramides, Canirallus and Mentocrex; an intermediate grade comprising Anurolimnas, Amaurolimnas, and Rougetius; monophyly of two major subdivisions of remaining rallids, one comprising Rallina (paraphyletic), Rallicula, and Sarothrura, and the other comprising the apparently paraphyletic ''long-billed'' rails (e.g. Pardirallus, Cyanolimnas, Rallus, Gallirallus and Cabalus and a variably resolved clade comprising ''crakes'' (e.g. Atlantisia, Laterallus and Porzana, waterhens (Amaurornis), moorhens (Gallinula and allied genera) and coots (Fulica). Relationships among ''crakes'' remain poorly resolved; Laterallus may be paraphyletic, and Porzana is evidently polyphyletic and poses substantial challenges for reconciliation with current taxonomy. Relationships among the species of waterhens, moorhens and coots, however, were comparatively well resolved, and exhaustive, fine-scale analyses of several genera (Grus, Porphyrio, Aramides, Rallus, Laterallus and Fulica) and species complexes (Porphyrio porphyrio -group,Gallirallus philippensis -group and Fulica americana -group) revealed additional topological likelihoods. Many nodes shared by a majority of the shortest trees under equal weighting were common to all shortest trees found following one or two iterations of successive weighting of characters. Provisional placements of selected subfossil rallids (e.g. Diaphorapteryx, Aphanapteryx and Capellirallus ) were based on separate heuristic searches using the strict consensus tree for modern rallids as a backbone constraint. These analyses were considered with respect to assessments of robustness, homoplasy related to flightlessness, challenges and importance of fossils in cladistic analysis, previously published studies and biogeography, and an annotated phylogenetic classification of the Gruiformes is proposed.     

Comparative morphology of mandibular structures in lycid larvae and its phylogenetic implications (Polyphaga, Hexapoda)

The morphology of the larval mandibular structures of the family Lycidae (Polyphaga) is characterized by facultative dicondyly, posterior articulation with long mandibular rods, lateral location of the anterior condyle and its articulation with a paired non‐cranial structure. It is compared to that of Eucrustacea, Chilopoda, Entognatha, Microcoryphia and Zygentoma and found to be more reminiscent of the Entognatha. The phylogenetic implications of this conclusion are discussed, with the Pterygota and Dicondylia hypothesized to be non‐monophyletic.