A cladistic analysis of the nomadine bees (Hymenoptera: Apoidea)

Abstract. This study compares the results of Rozen's cladistic analysis of the larvae of fifteen genera of cleptoparasitic bees in the subfamily Nomadinae with an independent data set of adult characters for the same genera. Adult characters exhibited considerably higher levels of homoplasy and poorer resolution of cladistic relationships, with multiple equally parsimonious cladograms. However, comparison of a Nelson consensus tree based on adult characters with the cladogram based on larval characters reveals three components consistently supported in both analyses (the tribes Epeolini and Ammobatini, and Neopasites + Neolarra) , one component supported only by adult characters (Isepeolus + Protepeolus) , and one terminal component supported only by larval characters (Nomada + Ammobatini), as well as several more inclusive groupings based on larval characters that are difficult to compare with the adult consensus tree because it shows so much less resolution. When adult and larval characters are combined in a single data matrix, the resulting cladogram closely resembles the cladogram based on larval characters alone, although levels of homoplasy are considerably higher than in the larval analysis.
A preliminary analysis of adult characters for thirty-four genera in the Nomadinae also exhibited high levels of homoplasy and very large numbers of equally parsimonious cladograms. Nevertheless, certain consistent monophyletic groupings, most notably the Epeolini and Ammobatini, were also supported in this analysis. The one currently recognized tribe whose monophyly has received no support from any analysis is the Nomadini.
The relevance of these phylogenetic hypotheses to our understanding of host associations and variable features of egg morphology and oviposition behaviour in nomadine bees is briefly discussed.     

Phylogeny of Drosophilinae (Diptera: Drosophilidae), with comments on combined analysis and character support

Drosophilidae (Diptera) is a diverse, cosmopolitan family of flies. Here, we present a combined analysis phylogeny of Drosophilinae, one of the two subfamilies of Drosophilidae, based on data from six different data partitions, including both molecular and morphological characters. Although our data show support for the monophyly of the Hawaiian Drosophilidae, and the subgenus Sophophora, neither the genus Drosophila nor the subgenus Drosophila is monophyletic. Partitioned Bremer support (PBS) indicates that morphological data taken from Grimaldi's monograph (Grimaldi, 1990a), as well as sequences from the mitochondrial (mt) 16S rDNA and the nuclear Adh gene, lend much support to our tree's topology. This is particularly interesting in the case of Grimaldi's data, since his published hypothesis conflicts with ours in significant ways. Our combined analysis cladogram phylogeny reflects the catch-all designation that the name Drosophila has become, in that the cladogram does not support the monophyly of either the genus or subgenus Drosophila.     

Rooting with Multiple Outgroups: Consensus Versus Parsimony

Using outgroup(s) is the most frequent method to root trees. Rooting through unconstrained simultaneous analysis of several outgroups is a favoured option because it serves as a test of the supposed monophyly of the ingroup. When contradiction occurs among the characters of the outgroups, the branching pattern of basal nodes of the rooted tree is dependent on the order of the outgroups listed in the data matrix, that is, on the prime outgroup (even in the case of exhaustive search). Different equally parsimonious rooted trees (=cladograms) can be obtained by permutation of prime outgroups. An alternative to a common implicit practice (select one outgroup to orientate the tree) is that the accepted cladogram is the strict consensus of the different equally parsimonious rooted trees. The consensus tree is less parsimonious but is not hampered with extra assumption such as the choice of one outgroup (or more) among the initial number of outgroup terminals. It also does not show sister-group relations that are ambiguously resolved or not resolved at all.     

A CLADISTIC ANALYSIS OF PARKIA (LEGUMINOSAE: MIMOSOIDEAE)

A cladistic analysis of the tropical mimosoid genus Parkia was undertaken to examine relationships among the 31 species and to test the monophyly of the three recognized sections. The implications of the cladogram to the evolution of bat-pollination and biogeography were also explored. The analysis, based on 52 morphological characters, resulted in 408 most parsimonious trees. A consensus tree supports the monophyly of sections Parkia and Platyparkia, but section Sphaeroparkia is paraphyletic. The latter section is distinguished by a capitulum of all fertile flowers, a plesiomorphic attribute in this analysis. Characters supporting the monophyly of section Platyparkia include an inflorescence with distal nectar flowers having exserted styles, and fruits with seeds in two rows. Section Parkia is characterized by having sterile basal flowers with nectar flowers just above them, and calyx lobes included in bud. Bat-pollination was mapped onto one of the most parsimonious cladograms to examine the evolution of pollination syndromes within the genus. Our phylogeny is consistent with a single origin of bat-pollination and indicates that entomophilous species of Parkia are basal rather than secondarily derived. Sections Platyparkia and Parkia are separate lineages within the bat-pollinated clade and have independently developed capitulum types adapted to chiropterophily. Characters that are associated with chiropterophily include specialized nectar-producing flowers and basifixed anthers. Several characters, including presence of a staminodial fringe, a nectar ring, and pollen with verrucate sculpturing on the exine, probably represent increasing specialization for bat-pollination. The cladogram supports a South American origin for Parkia but is not consistent with a Gondwanan vicariance event as is usually hypothesized to explain its amphi-Atlantic distribution.     

Systematics of the genus Cidaria Treitschke (Lepidoptera, Geometridae, Larentiinae)

The genus Cidaria Treitschke is revised. Eight species of the genus which occur widely in the Palaearctic and northern India, are recognized, of which one, Cidaria luteata sp. nov. , is described as new while two subspecies of C. fulvata (Forster) are elevated to species, C. nugata Felder stat. rev., and C. distinctata Staudinger stat. nov. C. ochripennis Prout is proposed as a junior synonym of C. ochreata Staudinger. C. deletaria Hampson is excluded from the genus. All species of Cidaria and their genitalia are described and illustrated. The cladistic analysis of these eight species of Cidaria is carried out using two data matrices, the first comprising morphological characters alone and the second morphological and distributional characters. The most parsimonious cladogram of Cidaria is selected, and its monophyly defined. The character analysis shows that some distribution characters contribute to resolving the in group node. The choice of multiple trees is discussed.     

A COMPUTER ASSISTED ANALYSIS OF THE RELATIONSHIPS OF THE HIGHER CATEGORIES OF TURTLES

Abstract— A character matrix of 39 characters for 14 supregeneric categories of living and extinct turtles was examined using PAUP 2.41 and 3.0L. The Branch and Bound algorithm found a single most parsimonious cladogram of 55 steps, consistency index of 0.709, retention index of 0.848 and a rescaled consistency index of 0.601. The cladogram is identical to that proposed by Gaffney and Meylan (1988). The Pleurodira and Cryptodira are each shown to be monophyletic and are supported by synapomorphies involving complex structures of the basicranium and adductor musculature. These synapomorphies are judged to be relatively well-tested homologies. A paraphyletic Cryptodira occurs in 18% of 38 equally parsimonious trees 57 steps in length, but these trees are based on characters, such as absence of pterygoid teeth, that are susceptible to homoplasy in amniotes. We re-iterate the notion that it is better to choose fewer, well-analysed characters than large numbers of poorly analysed characters.     

Phylogenetic interrelationships of the barbets (Aves: Capitonidae) and toucans (Aves: Ramphastidae) based on morphology with comparisons to DNA-DNA hybridization

A phylogenetic analysis of the interrelationships of the barbets (Capitonidae) and the toucans (Aves: Ramphastidae, Superfamily Ramphastoidea) is presented. Thirty-two morphological characters from the literature and independent osteological observations were analysed. Character polarity was determined by outgroup comparison to the Picidae, Indicatoridae, Galbulidae, Bucconidae and Coraciiformes. Four alternative phylogenetic hypotheses were compared: (1) the overall most parsimonious morphological phylogeny, (2) the most parsimonious morphological phylogeny in which the capitonids and ramphastids were hypothesized as monophyletic sister groups, and (3) and (4) the most parsimonious hypotheses for the evolution of the morphological characters within two proposed DNA-DNA hybridization phylogenies of the ramphastoids. The analysis focused on the higher level relationships of ramphastids and capitonids and interrelationships among capitonid genera. Two cladistic analyses were performed using 26 phylogenetically informative characters, and the PAUP and CONTREE computer alogorithms. The most parsimonious morphological phylogeny required fewer character changes and had a lower consistency index than any of the alternative hypotheses but congruence between the most parsimonious phylogeny and the second, revised DNA-DNA hybridization hypothesis was very high. Based on these results the monophyly of the Capitonidae is rejected. The ramphastids and the Neotropical capitonids form a well corroborated clade within the pantropical ramphastoid radiation. Neither the African, Asian nor New World capitonids is monophyletic. The genus Trachyphonus is the sister group to all other capitonids and ramphastids. The sister group to the ramphastids is the genus Semnornis. The interrelationships of the Old World capitonids excluding Trachyphonus are not completely resolved by these morphological data but one of the alternative phylogenetic resolutions is presented as a preliminary hypothesis. The clades in this resolved phylogeny are diagnosed and the palaeontology and biogeography of the ramphastoids arc-reviewed in light of this new evidence. A phylogenetic classification is proposed in which the Capitonidae is rejected and the capitonids and ramphastids are placed in seven subfamilies of the Ramphastidae.     

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。     

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.     

Testing the monophyly of the New Zealand and Australian endemic family Conoesucidae Ross based on combined molecular and morphological data (Insecta: Trichoptera: Sericostomatoidea)

Conoesucidae (Trichoptera, Insecta) are restricted to SE Australia, Tasmania and New Zealand. The family includes 42 described species in 12 genera, and each genus is endemic to either New Zealand or Australia. Although monophyly has been previously assumed, no morphological characters have been proposed to represent synapomorphies for the group. We collected molecular data from two mitochondrial genes (16S and cytochrome oxidase I), one nuclear gene (elongation factor 1-α) (2237–2277 bp in total), and 12 morphological characters to produce the first phylogeny of the family. We combined the molecular and morphological characters and performed both a maximum parsimony analysis and a Bayesian analysis to test the monophyly of the family, and to hypothesize the phylogeny among its genera. The parsimony analysis revealed a single most parsimonious tree with Conoesucidae being a monophyletic taxon and sistergroup to the Calocidae. The Bayesian inference produced a distribution of trees, the consensus of which is supported with posterior probabilities of 100% for 15 out of 22 possible ingroup clades including the most basal branch of the family, indicating strong support for a monophyletic Conoesucidae. The most parsimonious tree and the tree from the Bayesian analysis were identical except that the ingroup genus Pycnocentria changed position by jumping to a neighbouring clade. Based on the assumption that the ancestral conoesucid species was present on both New Zealand and Australia, a biogeographical analysis using the dispersal-vicariance criteria demonstrated that one or two (depending on which of the two phylogenetic reconstructions were applied) sympatric speciation events took place on New Zealand prior to a single, late dispersal from New Zealand to Australia.     

The phylogenetic interrelationships of the higher taxa of apterygote hexapods

The phylogeny of the basal hexapods, the so-called apterygote insects, was studied using parsimony analysis procedures. Most analyses took into account 47 characters mainly based on external morphology, and 19 taxa including 14 apterygote representatives, 3 pterygotes and also 2 distantly related myriapods were used as outgroups. The binary and multistate characters are discussed in detail and treated as unordered and equally weighted. Other analyses were performed using a second data set in which 28 characters, based on internal anatomy and already used in a previous work ( Bitsch & Bitsch 1998 ), were added to the first data set. This second matrix was restricted to 12 terminal taxa, the same as those of our previous work. The results of the different analyses are generally congruent. They strongly support the monophyly of several orders (Protura, Collembola, Archaeognatha) and of two groupings (Ectognatha, Dicondylia). Three other assemblages (Ellipura, Diplura, Entognatha) appear as parsimonious phylogenetic hypotheses, but they are never supported by the cladistical analyses and are based on a very small number of autapomorphies; so, the monophyly of each of them is not firmly established. Archaeognatha appears as the sister group of the Dicondylia. The three unresolved representatives of the Zygentoma are found as the sister group of the Pterygota. The results are discussed in the light of current concepts in hexapod phylogeny.     

Review of morphological evidence on the phylogeny of basal Hymenoptera (Insecta), with a discussion of the ordering of characters

In a previous study of the phylogeny of basal Hymenoptera, Vilhelmsen (2001; Zool. J. Linn. Soc . 131 : 393–442) compiled an extensive morphological data matrix for a phylogenetic analysis of basal Hymenoptera, comprising 38 hymenopteran genera. In this study, his characters are revised. This results in a cladogram whose relationships largely agree with those proposed by Vilhelmsen, except that the relationships at the base of the Hymenoptera are unresolved. The revised data matrix is expanded by 17 sawfly and three apocritan taxa. Moreover, 112 new morphological characters from different parts of the larval and adult morphology are also added to the data matrix, including 82 from a recent study of the terminal abdominal segments of male Hymenoptera. The addition of the new characters leads to Xyelidae, again, being the sister-group of all other Hymenoptera. The relationships among the sawfly families as proposed by Vilhelmsen are confirmed, except that the relationships among Syntexis , Siricidae and Xiphydriidae + Vespina are unresolved and that the monophyly of Apocrita is not convincingly supported. A separate analysis is performed which includes all extant genera of Xyelidae. The internal phylogeny of Xyelidae is determined as (( Macroxyela Megaxyela ) Xyelecia ( Xyela Pleroneura )).  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79 , 209–243.     

The ecological tale of Gonyleptidae (Arachnida,Opiliones) evolution: phylogeny of a Neotropical lineage of armoured harvestmen using ecological,behavioural and chemical characters

The large Neotropical family Gonyleptidae comprises nearly 820 species divided into 16 subfamilies. The majority of publications on harvestman ecology, behaviour and scent gland secretion chemistry have focused on this family. We used the information available in the literature and combined it with an intensive search for ecological, behavioural and chemical data to infer the phylogeny of the Gonyleptidae. We included 28 species belonging to 14 of the 16 gonyleptid subfamilies in the ingroup and four species belonging to the families Cosmetidae, Stygnidae and Manaosbiidae in the outgroup. We performed the analyses using equally weighted characters and coded 63 characters comprising 153 states, which makes this the largest non‐morphological, non‐molecular phylogenetic data matrix published to date. We obtained five most parsimonious trees, and the strict consensus resulted in six collapsed nodes. The results show that the monophyly of Gonyleptidae is equivocal because Metasarcinae is placed at a basal polytomy with the outgroups Cosmetidae and Stygnidae. Gonyleptinae, Pachylinae and Progonyleptoidellinae are polyphyletic groups, but the remaining subfamilies are monophyletic and have several synapomorphies. Based on the resulting topology, we discuss the performance of ecological, behavioural and chemical characters, and map a selected set of characters to discuss their evolutionary patterns in the family.     

Phylogenetic analysis of the Diplomonadida (Wenyon, 1926) Brugerolle, 1975: evidence for heterochrony in protozoa and against Giardia lamblia as a "missing link".

A suite of 23 ultrastructural characters was used in a phylogenetic analysis of the protozoan order Diplomonadida. A single most parsimonious solution was found, with a length of 38 transformations and a consistency index of 0.84. The cladogram supports previous hypotheses of the relationships of the genera in the suborder Diplomonadina, as well as the inclusion of the genera Enteromonas and Trimitus in the order. Heterochrony is suggested in the change to binary axial symmetry, as hypermorphosis resulting from delayed cytokinesis in the ancestor. Hypotheses regarding a pivotal position for Giardia lamblia in the evolution of eukaryotes are inconsistent with the phylogeny proposed here.     

Phylogenetic Analysis of the Diplomonadida (Wenyon, 1926) Brugerolle, 1975: Evidence for Heterochrony in Protozoa and Against Giardia lamblia as a "Missing Link"

ABSTRACT. A suite of 23 ultrastructural characters was used in a phylogenetic analysis of the protozoan order Diplomonadida. A single most parsimonious solution was found, with a length of 38 transformations and a consistency index of 0.84. The cladogram supports previous hypotheses of the relationships of the genera in the suborder Diplomonadina, as well as the inclusion of the genera Enteromonas and Trimitus in the order. Heterochrony is suggested in the change to binary axial symmetry, as hypermorphosis resulting from delayed cytokinesis in the ancestor. Hypotheses regarding a pivotal position for Giardia lamblia in the evolution of eukaryotes are inconsistent with the phylogeny proposed here.     

The phylogeny and classification of Scaphopoda (Mollusca): an assessment of current resolution and cladistic reanalysis

The first cladistic analysis of phylogeny in the class Scaphopoda (Steiner 1992a,1996) examined relationships among family and selected sub-family taxa using morphological data. A preferred/ consensus tree of relationships illustrated monophyly of the orders Dentaliida and Gadilida, partial resolution among dentaliid families, and complete resolution among gadilid taxa. However, several alternative replications of the analysis, including use of a revised data matrix, did not produce the reported tree number or level of resolution; in all cases, monophyly of the Dentaliida was not supported by strict consensus of resultant parsimonious trees. Reanalysis, using unordered characters and outgroup rooting, only clearly resolves monophyly of the Gadilida and the sister relationship of the Entalinidae with the remaining gadilid families. These analyses emphasize the need for more comparative data and thorough parsimony analysis in scaphopod cladistic phylogenetics, as relationships in this class are still some way from resolution.     

A comprehensive phylogenetic analysis of termites (Isoptera) illuminates key aspects of their evolutionary biology

The first comprehensive combined molecular and morphological phylogenetic analysis of the major groups of termites is presented. This was based on the analysis of three genes (cytochrome oxidase II, 12S and 28S) and worker characters for approximately 250 species of termites. Parsimony analysis of the aligned dataset showed that the monophyly of Hodotermitidae, Kalotermitidae and Termitidae were well supported, while Termopsidae and Rhinotermitidae were both paraphyletic on the estimated cladogram. Within Termitidae, the most diverse and ecologically most important family, the monophyly of Macrotermitinae, Foraminitermitinae, Apicotermitinae, Syntermitinae and Nasutitermitinae were all broadly supported, but Termitinae was paraphyletic. The pantropical genera Termes, Amitermes and Nasutitermes were all paraphyletic on the estimated cladogram, with at least 17 genera nested within Nasutitermes, given the presently accepted generic limits. Key biological features were mapped onto the cladogram. It was not possible to reconstruct the evolution of true workers unambiguously, as it was as parsimonious to assume a basal evolution of true workers and subsequent evolution of pseudergates, as to assume a basal condition of pseudergates and subsequent evolution of true workers. However, true workers were only found in species with either separate- or intermediate-type nests, so that the mapping of nest habit and worker type onto the cladogram were perfectly correlated. Feeding group evolution, however, showed a much more complex pattern, particularly within the Termitidae, where it proved impossible to estimate unambiguously the ancestral state within the family (which is associated with the loss of worker gut flagellates). However, one biologically plausible optimization implies an initial evolution from wood-feeding to fungus-growing, proposed as the ancestral condition within the Termitidae, followed by the very early evolution of soil-feeding and subsequent re-evolution of wood-feeding in numerous lineages.     

PHYLOGENETIC RELATIONSHIPS OF SCOTUSSA AND LEIOTETTIX (ORTHOPTERA: ACRIDIDAE)

Abstract— A cladistic analysis of the South American grasshopper genera Scotussa and Leiotettix was performed in order to test the monophyly of these genera. Eurotettix, Chlorus and the Dichroplus bergi species group were included as terminal taxa. The genus Atrachelacris was used to root the tree. Twenty-nine characters from external morphology, male genitalia and female ovipositor were used in the analysis. In order to test for association between the structural change that occurred in the ovipositor valves of Scotussa and the functional change of the oviposition habits, the data matrix was partitioned and two analyses were performed. Characters from the female ovipositor were excluded from the data set used in the first analysis and another analysis was performed where all the characters were included in the analysis. Information on oviposition habits was then mapped on the cladogram, to determine the transformation for performance. Both analyses yielded only one most parsimonious tree and produced congruent results, confirming the monophyly of Leiotettix and Scotussa and corroborating their close relationship. Characters from the female ovipositor valves were informative not only at the species level but also at higher levels in the cladogram. The results also support the hypothesis of association between the structural change that occurred in the ovipositor valves of Scotussa with the functional change in the oviposition habits. However, this association did not seem to be correlated with the adaptive radiation in the genus.     

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.