Phylogenetic relationships of the Atoposauridae (archosauria,crocodylomorpha)

A systematic revision of the family Atoposauridae is presented, interpreting and reviewing their relationships. The seventy‐one characters have been divided into cranial, postcranial and metric which have been used in three separate cladistic analyses in order to discuss their concordance. Polarities, in each case, have been deduced from distinct outgroups. We have used a PHYLIP program (version 2.9). There are five valid taxa of Atoposauridae: Alligatorellus beaumonti (rejecting the two subspecific taxa) Alligatorium meyeri, Theriosuchus pusillus. Alligatorium depereti (which is transferred to a new genus Montsecosuchus) and Aloposaurus (discussed as a nomen dubium). The three cladograms are not fully concordant. The relative level of derivation in postcranial and cranial characters suggests congruence. The metric traits seem to retain, in general, a primitive condition. The final proposed cladogram clusters Alligatorium + Alligatorellus as the sister group of Montsecosuchus + Theriosuchus.     

Characteristics of sperm ultrastructure in the gall midges Porricondylinae (Insecta,Diptera, Cecidomyiidae), with phylogenetic considerations on the subfamily

 Sperm ultrastructure was studied in ten genera of the Porricondylinae (Cecidomyiidae). Sperm structure is remarkably simplified by the absence of the acrosome and the accessory tubules, as happens in all the cecidomyiid flies. All genera of the Porricondylinae show a peculiar 9+3 axonemal model except Diallactes, which retains the plesiomorphic condition of a 9+2 axoneme, and Winnertzia, which appears to have secondarily acquired a 9+0 model. A cladistic analysis of relevant sperm characters (based on the axonemal model, the number of mitochondrial derivatives and the size and structure of the centriolar adjunct) was performed to infer phylogenetic relationships among six tribes of the Porricondylinae. In this cladogram, the Porricondylini are the sister group to the Asynaptini, Heteropezini and Winnertzini and these four taxa form the sister group to the Dicerurini. The tribe Diallactini are regarded as the group with the most plesiomorphic characters within the family. Accepted: 15 March 1996     

Cladistic analysis and biogeography of the cicadas of the Indo-Pacific subtribe Cosmopsaltriina (Hemiptera: Cicadoidea: Cicadidae)

Abstract A consensus cladogram is presented for ninety-seven of the ninety-nine (sub)species of the cicada subtribe Cosmopsaltriina on the basis of a cladistic analysis of forty-nine characters. The consensus cladogram confirms the monophyly of the eight genera constituting the subtribe, and the monophyly of the subtribe as a whole. Cosmopsaltriina occur in Sulawesi, the Moluccas, New Guinea, Cape York, the Solomon Islands, Vanuatu, Fiji, Tonga and Samoa. The subtribe demonstrates a high rate of endemism on genus, species group and species level. The taxon-area and consensus-area cladograms resulting from Brooks parsimony analysis are discussed in relation to the palaeogeography of the area.     

Phylogeny and evolutionary zoogeography of the Hawaiian Pipunculidae (Diptera)

Within the scope of a world revision of the genus group Cephalops Fallén, the pipunculid fauna of the Hawaiian islands was revised. All 36 species proved to belong to a distinct monophyletic species group within the genus Cephalops, called the hawaiiensis subgroup. This subgroup seems to be endemic to the islands and probably originated from one founder ancestor. A cladistic analysis, based on characters of the male terminalia, resulted in a largely resolved cladogram where a number of distinct transformation series could be detected. A zoogeographical analysis, based on assumption 0 (Brooks parsimony analysis), 1 and 2 (Component analysis) agrees with the geological information, and the ancestry and colonization of the group throughout the history is discussed.     

Phylogeny of scolopendromorph centipedes (Chilopoda): morphological analysis featuring characters from the peristomatic area

Alternative schemes for the higher‐level systematics of the centipede order Scolopendromorpha have been established from characters of trunk segmentation, including the segmental position of spiracles, and the presence or absence of eyes. A comparative survey of the preoral chamber by light and scanning electron microscopy contributes 16 new characters of the epipharynx and hypopharynx, sampled from 26 species that represent 20 genera. These new data together with 49 additional morphological characters permit cladistic analysis of the major scolopendromorph groups. The shortest cladograms resolve blind Scolopendromorpha as a basal grade within which a clade now classified as Plutoniuminae + Scolopocryptopidae (supported by unreversed characters from the preoral chamber) is sister to the remaining scolopendromorphs. A unique row of bullet‐shaped sensilla between the labral and clypeal parts of the epipharynx provides a new autapomorphy of the Scolopendromorpha. Either 21 or 23 trunk segments optimize at the base of the Scolopendromorpha but in either case homoplasy is forced on the cladogram. New characters from the epipharynx give additional support for the monophyly of several traditional groupings, including Cryptopinae, Scolopendridae, Otostigmini, and Scolopendrini, and a basal resolution of Asanadini within Scolopendridae. Of the two competing hypotheses for the position of the enigmatic Mediterranean Plutonium zwierleini—being either sister to the cryptopid Theatops or sister to all other Scolopendromorpha—the former hypothesis is strongly supported; spiracles on all trunk segments in Plutonium are homoplastic with the state in Geophilomorpha. Observations on feeding behaviour are needed to illuminate convergence in characters of the epipharynx and mandible in Edentistoma (Otostigminae) and Campylostigmus (Scolopendrini). © The Willi Hennig Society 2008.     

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.     

Cladistics and the comparative morphology of linyphiid spiders and their relatives (Araneae, Araneoidea, Linyphiidae)

This paper provides the first quantitative cladistic analysis of linyphiid morphology. Classical and novel homology hypotheses for a variety of character systems (male and female genitalia, somatic morphology, spinneret silk spigot morphology, etc.) are critically examined and studied within a phylogenetic context. Critical characters have been illustrated. A sample of linyphiid taxa (nine genera in four subfamilies), five species of Pimoa (Pimoidae), and two other araneoid families (Tetragnathidae and Araneidae, represented by Tetragnatha and Zygiella , respectively) were used to study the implications of the phylogeny of Pimoidae for the systematics of linyphiids. The phylogenetic relationships of these 16 exemplar taxa, as coded for the 47 characters studied, were analysed using numerical cladistic methods. In the preferred cladogram Pimoidae and Linyphiidae are sister groups, Stemonyphantinae are sister group to the remaining linyphiids, and Mynogleninae are sister group to the clade composed of Erigoninae plus Linyphiinae. These results agree with the relationships recently proposed by Wunderlich, except by finding erigonines as the sister group to linyphiines rather than to mynoglenines.     

Information obtained in cladistic analysis.

The application of elementary equations from information theory to the elements involved in cladistic analysis is formalized mathematically. An equation is derived that quantifies the amount of information obtained by constructing a cladogram from a cladistic data matrix. Given particular conditions, the amount of information obtained increases monotonically with increases of the number of taxa involved and, so, may be used directly as a comparative measure of species richness for sister groups; in general, however, the amount of information obtained is related to the distribution of character states on the cladogram(s) deduced. An example is presented in which clades representing 11 phyla in the animal kingdom are compared in terms of information yielded. The amount of information obtained is consistent for different numbers of taxa and characters used in classifications. Speculative evolutionary explanations are presented for differences of information yielded among the phyla analyzed.     

A method for estimating the relative importance of characters in cladistic analyses

The method of character importance ranking (CIR) is proposed here as a means for estimating the relative "importance" of characters in cladistic analyses, especially those based on morphological features. CIR uses the weighting variable to incrementally remove one character at a time from the analysis, and then evaluates the impact of the removal on the shape of the cladogram. The greater the impact, the more important the character. The CIR method for determining which characters drive the shape of a particular cladogram has several applications. It identifies the characters with the strongest (though not necessarily most accurate) signal in a cladistic analysis; it permits the informed prioritization of characters for further investigation via genetic, developmental, and functional approaches; and it highlights characters whose definition, scoring, independence, and variation should be reviewed with particular care. The application of CIR reveals that at least some cladograms depend entirely on a single character.     

裸藻类植物的分支系统学研究

本文选取了裸藻类的33个属级分类单位,以及它们的35个性状,利用分支系统学的原理和方法,对性状的演化极性进行了分析,同时对性状间的极性关系进行了和谐性分析,使性状间极性关系处在较为合理的状态,然后建立了分支分析的数据矩阵。应用徐克学建立的“演化极端结合法”进行微机运算．得简约系数远小于1(o．2159)的分支谱系图。根据分支诺系图对裸藻类的系统发育关系进行了探讨,井与已有的关于裸藻类分类系统和演化假设进行了比较。在此基础上按照裸藻类的亲缘关系及单系原则,对裸藻类的分类等级进行了划分,初步提出了建立1门1纲5目的分类系统。按照在分支谱系中的演化地位,认为裸藻属的5个亚属,明显地都应是独立的属。同时对裸藻类的共生起源与演化的关系也进行了讨论。     

Phylogenetic evidence for the herbaceous origin of angiosperms

The ancestral angiosperm is commonly interpreted as an arborescent to shrubby magnolialean with large, multiparted, complex flowers. We examined this hypothesis using a phylogenetic analysis of new and reevaluated characters polarizabled with outgroup comparison. Our cladistic analysis of basal angiosperms placed the nonmagnolialeanChloranthaceae andPiperaceae at the bottom of the tree. We further inferred the probable ancestral states of characters not polarizable with outgroup comparison by examining their distribution among taxa at the base of our cladogram. The sum of ancestral character states suggests that the protoangiosperm was a diminutive, rhizomatous to scrambling perennial herb, with small, simple flowers.     

Multivariate and cladistic analyses of the purple-flowered species ofErysimum (Cruciferae) from the Iberian Peninsula

A taxonomically difficult purple-flowered group within the genusErysimum, restricted to the Iberian Peninsula, is analyzed by multivariate and cladistic analyses. 51 specimens have been scored for 14 characters. Both principal components and discriminant analyses provide support to the recognition of the five species considered by the author, namelyE. linifolium, E. lagascae, E. baeticum, E. popovii, andE. cazorlense. Cladistic analysis, using 7 characters resulted in a single most parsimonious cladogram containing no homoplasies. The pattern of morphologic divergence follows a clear NW.-SE. trend, which is congruent with the topology of the cladogram. This trend significantly affects growth-form as well as fruit characters, both providing the main grounds for species recognition. The different behavior and significance of several characters in both kinds of analysis is discussed. The co-occurrence of morphologically similar individuals differing in the flower color is discussed, too. Possible explanations for this phenomenon involve hybridization in a wide sense or, alternatively, rejecting the assumption of monophyly for the group.     

PHYLOGENETIC RELATIONSHIPS OF THE CAMAENIDAE (PULMONATA: STYLOMMATOPHORA: HELICOIDEA)

The land snail family Camaenidae is defined by the lack of characters,such as dart sac and glands, which are present in related familiesAlthough the Camaenidae is regarded as monophyletic, no synapomorphiesunite the family It is possible that the Camaenidae is para-or polyphyletic This was investigated using a cladistic analysisof the super-family Helicoidea which was based on anatomicalcharacters from previous studies by Tilher (1989) Camaenidaewas split into two geographical groups (American and Australasian).Successive weighting generated a single most parsimonious cladogram(length 198, c.i. 0.61, r.i. 0.64), which suggested that theCamaenidae is a polyphyletic taxon. Evidence from this analysisdemonstrated that the American component of the Camaenidae isclosely related to the Hehcidae and Helminthoglyptidae, andthe Australasian component is sister group to the Bradybaenidae ^*Current address Department oJ Zoology. James Cook Universityof North Queensland. Townsville. Queensland 4811, Australia (Received 30 July 1994; accepted 20 June 1995)     

Higher Level Phylogeny of Curculionidae (Coleoptera: Curculionoidea) based mainly on Larval Characters, with Special Reference to Broad-Nosed Weevils

A cladistic analysis of Curculionidae was performed using 49 characters (41 from larvae, three from pupae, and five from adults). Illustrations of characters of immatures are provided. The analysis involved 19 terminal units and a hypothetical ancestor determined by the outgroup comparison method used to root the tree. One most parsimonious cladogram was obtained based on the complete data set and the following phylogenetic hypothesis is proposed: Ithycerinae, Microcerinae, and Brachycrinae sensu stricto are broad-nosed weevils placed sequentially at the base of the cladogram. The remaining weevil subfamilies form two major natural groups: one constituted by the sister taxa Rhynchophorinae—Platypodinae; the other with Erirhininae at the base, as sister taxon of the "Curculionidae sensu stricto " which show an unresolved trichotomy involving Curculioninae, Cossoninae—Scolytinae, and the clade including the Entiminae and allied subfamilies. This latter clade of broad-nosed weevils has Thecesterninae at the base; the next branch is Amycterinae, the sister taxon of the clade comprising two groups: one constituted by Aterpinae, Rhytirrhininae, and Gonipterinae; the other is Entiminae whose units form two main clades: one constituted by the sister tribes Pachyrhynchini—Ectemnorhinini, and the other by Alophini, Sitonini, and Entimini. When the analysis was done using only immature characters, results congruent with those based on the complete data set were obtained, except for the placement of Erirhininae. According to the results the hypothesis of monophyly of broad-nosed weevils is not accepted; the Entiminae are justified as monophyletic and their natural classification into tribes is proposed and the phylogenetic position and relationships of higher taxa of Curculionidae are discussed. This paper shows the importance of immature characters in recognition of natural groups and relationships in Curculionidae.     

Cladistics of Sunaristes,a genus of harpacticoid copepods associated with hermit crabs

Four species of Sunaristes (Canuellidae) are known to live in association with hermit crabs and exhibit a degree of host specificity. Although hermit crabs are common in many parts of the world, Sunaristes is notably absent from waters of the New World. The phylogeny of Sunaristes is here examined through a cladistic analysis. The reconstructed phologeny indicates that S. inaequalis and S. japonicus form a sister group, and its closest relative is S. tranteri. The sister taxon of this 3-species clade is S. paquri. It is assumed that Sunaristes evolved in the Paleocene on the north shore of the Tethys seaway. The constructed cladogram forms the basis for the analysis of vicariant events that led to the speciation and development of the present pattern of distribution of Sunaristes.     

Bartolius pierrei n. g., n. sp. (Digenea: Gymnophallidae) from the Península Valdés, Argentina

Bartolius pierrei n. g., n. sp. (Digenea: Gymnophallidae) is described from metacercariae and naturally and cultivated obtained adults from southern Argentina. The second intermediate host is Darina solenoides (King) (Bivalvia: Mactridae) and the definitive host is Larus dominicanus Lichtenstein (Aves: Laridae). The diagnostic characters are as follows: Body small, oval. Oral sucker without lateral projections, twice size of ventral sucker (except in young metacercariae). Caeca short (in adults), without dorsal diverticula. Ventral sucker in posterior third of body. Ventral pit absent. Seminal vesicle bipartite. Ovary post-testicular. Vitelline glands paired, compact, close to ventral sucker. Uterus in fore- and hindbody. Genital atrium tubular. Genital pore inconspicuous, close to anterior margin of ventral sucker. Excretory vesicle Y-shaped with very short stem. Excretory formula: 2[(2+2)+(2+2)]=16. Bartolius is distinguished from other genera of the Gymnophallidae by the post-testicular position of the ovary.     

CLADISTIC ANALYSIS OF PATTERNS OF ENDOTHECIAL THICKENINGS IN THE POALES/RESTIONALES

The three-dimensional structure of the endothecial thickenings in the anthers was investigated in 87 species from 70 genera, chosen to provide representative coverage of the families Cyperaceae, Restionaceae, Anarthriaceae, Ecdeiocoleaceae, Centrolepidaceae, Joinvilleaceae, Flagellariaceae, Poaceae, Xyridaceae, and Eriocaulaceae. There is complex variation in the patterns of endothecial thickening: the Eriocaulaceae, Flagellariaceae, and most Poaceae have thickenings with a complete baseplate; the Cyperaceae and most of the Restionaceae are characterized by helical thickenings; some genera in the Bambusoideae have annular thickenings; and U-shaped thickenings occur in the Xyridaceae and Eriocaulaceae and in some Poaceae and Restionaceae. Joinvillea and Ecdeiocolea have unique thickening types. Endothecial characters were subjected to cladistic analysis. Including endothecial characters in an existing cladogram of the group indicates that there is no single, well-corroborated cladogram available for the Poales/Restionales.     

A cladistic phylogeny of the genus Littorina (Gastropoda): implications for evolution of reproductive strategies and for classification

Gross anatomical characters of all 18 species of Littorina are used to construct a phylogenetic hypothesis for the genus, by the method of cladistic analysis. The resulting cladogram suggests that of the four subgenera of Littorina, one (Littorina) is paraphyletic. It is uncertain whether the genus Mainwaringia should be included in Littorina. It is shown that the non-planktotrophic Littorina species in the northern Atlantic comprise a monophyletic group, with the sister-species L. kurila and/or L. subrotundata in the northern Pacific. Invasion of the Atlantic by a minimum of two Pacific species, across the Arctic migration route established during the late Cenozoic, is sufficient to account for the modern distribution of the subgenera Littorina and Neritrema. The importance of the cladogram as a basis for hypotheses of adaptation is illustrated by a discussion of spawn and development in Littorina.     

Weighted ancestral area analysis and a solution of the redundant distribution problem

A new cladistic method for the estimation of ancestral areas is based on reversible parsimony in combination with a weighting scheme that weights steps in positionally plesiomorphic branches more highly than steps in positionally apomorphic branches. By applying this method to cladograms of human mitochondrial DNA, the method is superior to previously proposed algorithms. The method is also an appropriate tool for the solution of the redundant distribution problem in area cladograms. Under the assumption of allopatric speciation, redundant distributions, i.e., sympatry of sister groups, show that dispersal has occurred; thus, the ancestral area of at least one sister group was smaller than the combined distribution of its descendants. With the weighted ancestral area analysis, the ancestral areas can be confined and at least some dispersal events can be distinguished from possible vicariance events. As applied to a cladogram of the Polypteridae, weighted ancestral area analysis is superior to Brooks parsimony analysis (assumption 0) and component analysis under assumptions 1 and 2 (Nelson and Platnick, 1981, Systematics and biogeography: Cladistics and vicariance. Columbia Univ. Press, New York.) in resolving redundancies. The results of the weighted ancestral area analysis may differ from the results of dispersal-vicariance analysis, because the rules of dispersal-vicariance analysis indirectly favor the questionable assumption that the ancestral species occupied only one unit area.     

Sophisticated falsification and research cycles: Consequences for differential character weighting in phylogenetic systematics

Practicing phylogenetic systematics as a sophisticated falsification research program provides a basis for claiming increased knowledge of sister species relationships and synapomorphies as evidence for those cladistic propositions. Research in phylogenetic systematics is necessarily cyclic, and the place where the positive shift in understanding occurs is subsequent to discovering the most parsimonious cladogram(s). A priori differential character weighting is inconsistent with seeking the maximally corroborated cladogram (sensu Popper), because weighting adds to background knowledge, the evidence being then less improbable than it would be otherwise. Also, estimating weights from character state frequencies on a cladogram is inconsistent with the view that history is unique. Sophisticated falsification provides the place in the cycle of phylogenetic systematic research where weight of evidence can be evaluated and these inconsistencies do not apply. On balance, phylogenetic systematics appears to achieve greater coherence and generality as a result of focusing on the foundations for claiming increased knowledge and avoiding efforts to differentially weight characters.