Morphological Characters Are Compatible with Mitogenomic Data in Resolving the Phylogeny of Nymphalid Butterflies (Lepidoptera: Papilionoidea: Nymphalidae)

Nymphalidae is the largest family of butterflies with their phylogenetic relationships not adequately approached to date. The mitochondrial genomes (mitogenomes) of 11 new nymphalid species were reported and a comparative mitogenomic analysis was conducted together with other 22 available nymphalid mitogenomes. A phylogenetic analysis of the 33 species from all 13 currently recognized nymphalid subfamilies was done based on the mitogenomic data set with three Lycaenidae species as the outgroups. The mitogenome comparison showed that the eleven new mitogenomes were similar with those of other butterflies in gene content and order. The reconstructed phylogenetic trees reveal that the nymphalids are made up of five major clades (the nymphaline, heliconiine, satyrine, danaine and libytheine clades), with sister relationship between subfamilies Cyrestinae and Biblidinae, and most likely between subfamilies Morphinae and Satyrinae. This whole mitogenome-based phylogeny is generally congruent with those of former studies based on nuclear-gene and mitogenomic analyses, but differs considerably from the result of morphological cladistic analysis, such as the basal position of Libytheinae in morpho-phylogeny is not confirmed in molecular studies. However, we found that the mitogenomic phylogeny established herein is compatible with selected morphological characters (including developmental and adult morpho-characters).     

Phylogeny of the Pyretophorus Series of Anopheles subgenus Cellia (Diptera: Culicidae)

A cladistic analysis based on morphological characters is presented for the Pyretophorus Series of subgenus Cellia of Anopheles. The cibarial armature was investigated for the first time since the pioneering work of Christophers (1933) , which is still used to define taxonomic series within Cellia, and for the first time ever using scanning electron microscopy. Christophers’ observations of characters of the cibarial armature were corroborated and several novel characters discovered. Despite difficulties in delimiting and encoding morphological characters at the species level, the analysis provided insights into relationships within the Pyretophorus Series. The data support the monophyly of the Series and establish the Oriental species as a monophyletic group. The Afrotropical species are basal and paraphyletic relative to the Oriental species, with Anopheles christyi placed in the most basal position. The pattern of relationships suggests that the capacity to transmit malaria is an ancestral condition subsequently lost independently in several lineages.     

Phylogenetic relationships within the Eurasian cattish family Siluridae (Pisces: Siluriformes), with comments on generic validities and biogeography

A numerical cladistic analysis, based on 23 terminal groups and 63 morphological characters, was done to infer phylogenetic relationships within the Eurasian catfish family Siluridae. Nine hundred and forty-five equally most parsimonious trees (134 steps, consistency index 0.634) were found that differ in their resolutions of four polychotomies. Strict consensus of these trees includes ten internal nodes, does not support monophyly of Silurus, Ompok and Kryptopterus , as usually defined, and offers ambiguous support for monophyly of Wallago. Silurus and Kryptopterus are each composed of two non-sister group clades, and Ompok is composed of at least two such clades. Heuristic searches constrained by monophyly of Silurus, Ompok or Kryptopterus yielded trees five or six steps longer than the shortest trees free of constraints. The strict consensus also infers a basal dichotomy that separates the Siluridae into a temperate Eurasian clade with about 20 nominal species and a subtropical/tropical south and southeast Asian clade with about 75 nominal species. The distributions of these clades overlap in a relatively narrow region of east Asia. A heuristic search for trees 1 step longer than the shortest trees yielded 253890 trees. A strict consensus of these trees also infers a basal dichotomy between the above-mentioned clades. This analysis revealed four additional putative synapomorphies of the Siluridae, pending further resolution of the family's outgroup relationships.     

Cladistic analysis of the alderflies of America north of Mexico (Megaloptera: Sialidae)

Abstract. A cladistic analysis of thirty-five morphological characters is used to produce a cladogram for the twenty-four American species of Sialis found north of Mexico, two European species, and three species of Protosialis . Three species groups ( americana, califomica and infumata ) proposed by Ross 1937 are shown to be monophyletic. Sialis, Protosialis , and the aequalis species group as currently constituted, are paraphyletic; S.nevadensis is nested within Protosialis and Protosialis is nested within Sialis . Character evolution is evaluated in the light of a phylogenetic hypothesis, and structural modifications are found to correlate with presumed functional constraints.     

Phylogeny and classification of the Culicidae (Diptera)

The generic relationships and higher classification of the family Culicidae are examined on the basis of a phylogenetic analysis. New and traditional morphological characters studied and compared throughout the Culicidae resulted in the acquisition of character data relative to the majority of species within each genus. Polymorphisms and morphological observations are discussed and additional information and illustrations are provided for the majority of characters and their character states. The analysis of seventy-three adult, pupal and fourth-instar larval characters coded for the thirty-eight currently recognized genera of mosquitoes resulted in relationships and groupings which differ significantly from traditional hypotheses. The analysis supports the monophyly of the subfamily Anophelinae and the tribes Culicini and Sabethini. The Anophelinae form the most basal clade of the family. The results indicate that Aedini is a paraphyletic assemblage with respect to the Mansoniini, each of which is monophyletic in itself. The Aedini + Mansoniini form a sister group to the Culicini + Sabethini, with the Aedini and the Culicini placed in ancestral relationships to the Mansoniini and the Sabethini, respectively. Based on the topography of generic relationships among more 'generalized' mosquitoes, the boundaries and relationships of the tribes Aedeomyiini, Uranotaeniini, Ficalbiini, Hodgesiini, Orthopodomyiini and Culisetini appear to be problematic. Relationships between genera of the tribe Aedini are generally poorly resolved due to a significant amount of polymorphism, especially within the genus Aedes as currently defined. There is no support for separate subfamily recognition for the genus Toxorhynchites , which is downgraded to tribal status as a result of the analysis. The results are discussed in relation to previous hypotheses based on subjective inference and cladistic analyses.     

Cladistic analysis of the Sepsidae (Cyclorrhapha: Diptera) based on a comparative scanning electron microscopic study of larvae

The result of a phylogenetic analysis of the Sepsidae based on larval characters is presented. It is shown that cyclorrhaphan larvae can be as rich a source of characters as Nematocera immatures when investigated using an SEM. The cladistic analysis comprised fifty-two species in sixteen genera of the Sepsidae and five outgroup species and used fifty-seven morphological characters. It found seven parsimonious trees which only differed with respect to the arrangement of some species within the genus Themira. The basal dichotomies of the phylogenetic trees are particularly well supported, indicating the conservative nature of larval characters. Orygma is confirmed as the sister group of all the remaining sepsids, the Sepsinae. There is good larval evidence that Ortalischema is the sister group of all remaining Sepsinae and that the Toxopodinae constitute an early radiation within the Sepsidae. According to larval data, some genera are paraphyletic ( Themira, Palaeosepsis ), but adult characters appear to contradict these findings. Among the traditionally recognized higher taxa within the Sepsidae, Hennig's Themira species-group and Steysbal's Sepsini have to be rejected as polyphyletic. However, Hennig's Sepsis species-group is confirmed as monophyletic and will probably constitute one major element of a future phylogenetic system of the Sepsidae. States of the strongly modified fore-legs of some adult sepsid males are mapped onto the phylogenetic tree, largely confirming Šulc's ideas about the evolution of these features. The origin and evolution of male sternites with brushes and a gland on the tibiae of the males ('osmeterium') are discussed. Whereas adult characters point to a sister-group relationship between the Sepsidae and the Ropalomeridae, larval characters appear to indicate a sister-group relationship between the Coelopidae and the Sepsidae. The evidence for both hypotheses is critically evaluated.     

基于28S rDNA D2基因片段与形态特征的优茧蜂亚科系统发育研究

本研究选取优茧蜂亚科Euphorinae(膜翅目Hymenoptera:茧蜂科Braconidae)的8族19属23种作为内群,茧蜂其它6个亚科的8属8种作外群,首次结合同源核糖体28S rDNA D2基因序列片段和41个形态学特征对该亚科进行了系统发育学研究。利用"圆口类"的内茧蜂亚科Rogadinae、茧蜂亚科Braconinae、矛茧蜂亚科Doryctinae的3个亚科为根,以PAUP*4.0和MrBayes3.0B4软件分别应用最大简约法(MP)和贝叶斯法对优茧蜂亚科的分子数据和分子数据与非分子数据的结合体进行了分析;并以PAUP*4.0对优茧蜂亚科的28S rDNA D2基因序列的片段的碱基组成与碱基替代情况进行了分析。结果表明:优茧蜂亚科的28S rDNA D2基因序列片段的GC%含量在40.00%~49.25%之间变动,而对于碱基替代情况来讲,优茧蜂亚科各个成员间序列变异位点上颠换(transversion)大于转换(transition);不同的分析和算法所产生的系统发育树都表明目前根据形态定义出的优茧蜂亚科Euphorinae不是一个单系群,而是一个与蚁茧蜂亚科Neoneurinae和高腹茧蜂亚科Cenocoelinae混杂在一起的并系群;在优茧蜂亚科内部,悬茧蜂族Meterorini和食甲茧蜂族Microctonini(排除猎户茧蜂属Orionis)为单系群,而宽鞘茧蜂族Centistini、大颚茧蜂族Cosmophorini、优茧蜂族Euphorini、瓢虫茧蜂族Dinocampini为并系群;悬茧蜂族Meterorini在优茧蜂亚科Euphorinae内位于基部位置的观点得到部分的支持,同时食甲茧蜂族Microctonini被判定为相对进化的类群。此外对于优茧蜂亚科内各属之间的相互亲缘关系,不同算法所得到的系统发育属的结果不完全一致,这表明优茧蜂亚科内(属及族)的系统发育关系还有待于进一步研究。     

Generic-level relationships within the ant subfamily Dolichoderinae (Hymenoptera: Formicidae)

A cladistic analysis was undertaken to determine relationships among extant genera of the ant subfamily Dolichoderinae. Twenty-one of the twenty-two currently recognized genera within the subfamily were examined using 104 morphological characters. A single fully resolved, most-parsimonious tree was found when a combination of ordered and unordered characters was used. When all characters were coded as unordered, seventy most-parsimonious trees were found. The following results were found with both character coding methods. Leptomyrmex was placed basal to the remainder of the subfamily and the monophyletic sets Dolichoderus + Liometopum + Axinidris + Tapinoma + Technomyrmex, Froggattella + Iridomyrmex + Ochetellus + Papyrius + Philidris + Turneria , and Bothriomyrmex + Dorymyrmex + Forelius + Loweriella were suggested. The genera Linepithema and Doleromyrma showed a small amount of instability in moving between neighbouring sister groups when the character coding method changed. The genera Anillidris and Anonychomyrma were difficult to place as they showed major differences in their positions between the two character coding methods.     

槭树科植物广义形态学性状分支分析

通过45个广义的形态学性状对槭树科(Aceraceae)尤其是槭属(Acer L.)的主要类群做了分支分析,结果显示：1)槭属内由于各类群分布着较多的同塑性状状态,而难以为属下组间关系的解决提供更多有价值的信息;通过对具体的性状状态分布分析显示,对于象槭属这样在形态上分化较大的类群,由于多数分类性状在不同类群间经历了平行和逆转演化,因而在较低分类阶元水平很难选择合适的性状来通过分支分析构建其系统发育;2)鸡爪槭组(section Palmata)作为整个槭属的基部类群,虽然支持率较低,但与其它类群相比在槭属内维持了较多的原始性状;3)金钱槭属(Dipteronia Oliv.)的两个种作为单系得到了100％的靴带支持,且和槭属作为姐妹群也得到了较好的支持。     

PHYLOGENY OF THE NEMERTEAN SUBCLASS PALAEONEMERTEA (ANOPLA, NEMERTEA)

Abstract— A cladistic analysis based on 50 morphological characters was performed for 49 of the 98 species currently assigned to the subclass Palaeonemertea (phylum Nemertea), and six additional undescribed species. Thirty-five species were excluded from the parsimony analysis because of the high number of unknowns in the character matrix, and one species since it was considered a nomen nudum . An initial analysis suggested that the subclass Hoplonemertea is the sistergroup to the clade Palaeo- and Heteronemertea and the ingroup cladograms are rooted using a paraphyletic outgroup based on this information. Seventy-two equally most parsimonious cladograms were found; the consistency index was low but tree-length distribution for the character set is skewed to the left, and the cladograms are invariably shorter than trees based on random data. These cladograms suggested a character transformation series for the cerebral organ where this complex character reappeared several times after being absent. We considered this biologically implausible and the final discussion is based on three cladograms, one step longer than the most parsimonious, where the evolution of this character appears to be more realistic. The cladistic analysis indicates that many previously recognized genera (e.g. Cephalothrix, Procephalothrix and Cephalotrichella ), and higher taxa, are paraphyletic. It furthermore indicates that the previously suggested hypothesis of the Archinemertea as a monophyletic sistertaxon to Palaeonemertea is unsupported.     

Phylogeny of the tribe Colletieae, Rhamnaceae

A cladistic analysis based on 63 morphological characters was carried out on the tribe Colletieae including two presumed closely related genera, Geanothus and Noltea as outgroupS. In addition to a parsimony analysis of the equally weighted characters, analyses investigating the effects of character weighting, removal of a presumed hybrid species as well as the impact of uncertainly scored characters were undertaken. In all analyses Noltea was placed as sister group to a well supported monophyletic Colletieae. Nineteen different in group topologies were found, with the additional analyses mainly supporting two of them. Within the Colletieae a basal dichotomy divides Trevoa and Retanilla from the remainder of the tribe. While the Trevoa-Retanilla clade is fully resolved, the second lacks detailed resolution. Within this clade the Colletia species from a well supported monophyletic group, while monophyly of the disjunct genus Discaria could not be confirmed.     

A phylogenetic analysis of the nursery-web spider family Pisauridae, with emphasis on the genera Architis and Staberius (Araneae: Lycosoidea)

This study presents a general cladistic analysis of pisaurid genera, with emphasis on the Neotropical genera Architis Simon and Staberius Simon. The analysis was based on a matrix with 21 terminal taxa: eight species of Architis , Staberius spinipes (Taczanowski), seven exemplars of other pisaurid genera and five outgroup taxa. These terminals were scored for 59 morphological and three behavioural characters. An analysis with all characters equally weighted resulted in two most parsimonious trees, differing only in the position of Architis colombo Santos. In both trees Pisauridae arouse as a monophyletic group, and the exemplars of Architis composed a clade with S. spinipes . Contrary to hypotheses from the literature, Thalassius Simon emerged as sister-group of all remaining pisaurids, not as a close relative of Dolomedes Latreille. The genera Tinus F.O.P. -Cambridge and Thaumasia Perty appeared in the trees as the closest relatives of Architis and Staberius Simon. The analysis strongly supported S. spinipes as the sister-group of A. helveola (Simon), indicating that Architis as currently delimited is paraphyletic. Based on these results, Staberius is considered a subjective junior synonym of Architis . Additionally, the genus Mimicosa Petrunkevitch, originally described in Tetragnathidae, is transferred to Pisauridae and considered a junior synonym of Architis . Two species are proposed as junior synonyms of A. spinipes comb.n. Mimicosa spinosa Petrunkevitch and Staberius lemoulti Caporiacco.     

Phylogenetic analysis of the genus <Emphasis Type="Italic">Argia</Emphasis> Rambur, 1842 (Odonata: Coenagrionidae), based on morphological characters of larvae and mitochondrial DNA sequences

The study of the evolutionary interrelationships among the species encompassed in the Neotropical genus Argia (Zygoptera: Coenagrionidae) has been neglected. The goal of this study is to infer the phylogenetic relationships among 36 species of Argia Rambur, 1842, using complementary data sets (i.e., larval morphology and mitochondrial DNA). The morphological data set comprises 76% of the larvae currently described for this genus and includes 97 morphological characters. From those, 47 characters have not been previously used in taxonomic studies involving dragonflies’ larvae. This is the first cladistic study based on larvae morphology for species within the suborder Zygoptera. Data partitions were analyzed individually, as well as total evidence, using parsimony and Bayesian inference as criteria for optimal-tree selection. The results support the monophyly of the North American species of Argia. This genus can be identified by the combination of eight synapomorphies, four of which are exclusively found in Argia. According to the optimal trees, the individual data sets (i.e., morphology and DNA sequences) have a high level of homoplasy, resulting in soft polytomies and low support for several nodes. The specific relationships of the terminal units differ between the phylogenies; nonetheless, there is historical congruence among them. Within Argia, five clades were consistently recovered. Most of those clades have been identified, at least in part, in previous phylogenetic and taxonomic studies. Indubitably, the morphological characters from larvae have historical signal useful for cladistic and taxonomic inference. Therefore, it should be a priority to pay more attention to this source of characters.     

Phylogeny reconstruction in the neogene bryozoan metrarabdotos: A paleontologic evaluation of methodology

An adequate stratigraphic record can not only aid in both cladistic and stratophenetic reconstruction of phytogenies, but can also serve in estimating the temporal consistency of the resulting phylogenetic trees. For hypothetical data sets, cladistically constructed trees can be as consistent with the temporal distribution of sampled populations or species as those constructed stratophenetically. Empirical testing in taxonomic groups with sufficiently dense fossil records is needed to show whether, and under what conditions, this potential can be realized. A stratophenetic tree and cladistic trees based on several approaches to character weighting were constructed for Caribbean Neogene species of the bryozoan Metrarabdotos with multiple‐character data from closely spaced sequential populations. The modular morphology and highly punctuated evolutionary pattern of these species blur the distinction between continuous and discrete characters, so that all available characters are potentially of equal significance in establishing phytogenies, rather than just those with discrete states conventionally used in cladistic analysis. However, only the cladistic trees generated with all characters weighted to emphasize contribution to species discrimination have temporal consistencies that are clearly significant statistically and approach that of the stratophenetic tree in magnitude. These results provide a start toward establishing general guidelines for cladistic analysis of taxa with stratigraphie records too sparse for stratophenetic reconstruction.     

五味子科的系统发育:核糖体DNA ITS区序列证据

The Schisandraceae is one of the most important taxa for understanding the origin and evolution of primitive angiosperms due to its basal position in the recent cladograms of the angiosperm based both on several gene sequences and on morphological characters, but phylogenetic relationships within the family are still unresolved. The sequences of nrDNA ITS region of 15 species representing four sections of Schisandra Michx., two sections of Kadsura Kaempf. ex Juss. and one outgroup, Illicium fargesii Finet et Gagnep., were used to reconstruct the phylogeny of Schisandraceae. Fourteen most parsimonious trees (Length=259, CI=0.934 and RI=0.889) were obtained from the analysis with I．fargesii as the outgroup. In the consensus tree, the genus Schisandra was found to be divided into three clades. Sect. Pleiostema formed a clade together with sect. Maximowiczia, sect. Sphaerostema was weakly supported to be the sister group of a clade comprising S. bicolor var. tuberculata and Kadsura species. In particular, S. bicolor var. tuberculata, a species of sect. Schisandra, was nested within Kadsura. It seems from this result and the morphological characters that Schisandra might not be a monophyletic group. According to the present molecular phylogeny, both elongated hypocarpium and deciduous habit originated independently at least twice in the Schisandraceae, and therefore, the value of the present morphological characters used in the classification of the family Schisandraceae should be reevaluated.     

Evolutionary constraints in hind wing shape in Chinese dung beetles (Coleoptera: Scarabaeinae)

This study examines the evolution hindwing shape in Chinese dung beetle species using morphometric and phylogenetic analyses. Previous studies have analyzed the evolution of wing shape within a single or very few species, or by comparing only a few wing traits. No study has analyzed wing shape evolution of a large number of species, or quantitatively compared morphological variation of wings with proposed phylogenetic relationships. This study examines the morphological variation of hindwings based on 19 landmarks, 119 morphological characters, and 81 beetle species. Only one most parsimonious tree (MPT) was found based on 119 wing and body characters. To better understand the possible role of the hindwing in the evolution of Scarabaeinae, additional phylogenetic analyses were proposed based on the only body features (106 characters, wing characters excluded). Two MPT were found based on 106 body characters, and five nodes were collapsed in a strict consensus. There was a strong correlation between the morphometric tree and all phylogenetic trees (r>0.5). Reconstructions of the ancestral wing forms suggest that Scarabaeinae hindwing morphology has not changed substantially over time, but the morphological changes that do occur are focused at the base of the wing. These results suggest that flight has been important since the origin of Scarabaeinae, and that variation in hindwing morphology has been limited by functional constraints. Comparison of metric disparity values and relative evolutionary sequences among Scarabaeinae tribes suggest that the primitive dung beetles had relatively diverse hindwing morphologies, while advanced dung beetles have relatively similar wing morphologies. The strong correlation between the morphometric tree and phylogenetic trees suggest that hindwing features reflect the evolution of whole body morphology and that wing characters are suitable for the phylogenetic analyses. By integrating morphometric and cladistic approaches, this paper sheds new light on the evolution of dung beetle hind wings.     

A Phylogenetic Study of Tribe Euphorbieae (Euphorbiaceae)

A phylogenetic investigation of a monophyletic lineage of spurge plants, tribe Euphorbieae, was conducted to elucidate evolutionary relationships, to clarify biogeographic patterns, and to reexamine the previous classification of Euphorbieae. Cladistic analyses of the 52 morphological characters of 61 species resulted in 2922 equally most parsimonious trees of 193 steps with a consistency index of 0.34. The strict consensus tree indicates genus Anthostema of subtribe Anthosteminae as a likely sister group to all other members of tribe Euphorbieae. The morphological data support a monophyletic origin of subtribe Euphorbiinae, but the subtribes Anthosteminae and Neoquillauminiinae did not form monophyletic groups. Although the previous taxonomic treatments within tribe Euphorbieae have supported the generic status of Pedilanthus, Monadenium, Synadenium, Chamaesyce, and Elaeophorbia, the results of this analysis do not support generic placement of them based on cladistic principles. Recognition of these groups as genera results in Euphorbia becoming a paraphyletic group. One solution to this problem in Euphorbieae is to divide the largest genus Euphorbia into several monophyletic genera and to keep the generic ranks for previously recognized genera. The distribution of basal endemic genera in Euphorbieae showed African and east Gondwanan affinities and strongly indicated that the ancestor of Euphorbieae originated prior to the breakup of Gondwanaland from an old group in Euphorbiaceae. However, some recent African taxa of Euphorbia should be interpreted by transoceanic dispersal from the New World ancestors.     

Preliminary morphological analysis of relationships between the spider wasp subfamilies (Hymenoptera: Pompilidae): revisiting an old problem

No qualitative cladistic analysis has been performed previously for the subfamily classification of Pompilidae (Hymenoptera). In 1994 Shimizu proposed six subfamilies, but their validity and relationships remain inconclusive. The objective of this study was to perform a quantitative analysis of phylogenetic relationships of the Pompilidae, with emphasis on testing the validity of proposed subfamilies. Two cladistic analyses were performed based on morphological evidence. First, a maximum-parsimony analysis of Shimizu's original morphological data matrix (72 taxa by 54 characters) was conducted, with the data subjected to a heuristic search for the first time with phylogenetic software. The resulting strict-consensus cladogram yielded a monophyletic Ceropalinae that was sister group to a large polytomy containing members of the remaining five subfamilies. In a second analysis, several of Shimizu's characters were re-examined, and new characters and more taxa were added to the data set. Terminal taxa were coded as species rather than as generic abstractions, and 20 additional morphological characters were introduced. The analysis was based on 77 morphological characters derived from the adults of 84 taxa. This second analysis suggested that Notocyphinae sensu Shimizu (1994) was nested within Pompilinae and that Epipompilinae sensu Shimizu (1994) was nested within Ctenocerinae; neither should retain their status as a separate subfamily. Lastly, Chirodamus s .s., which historically has been a member of the Pepsinae, is placed within the Pompilinae with reservations rather than erecting a new subfamily. After these allowances were made, a strict consensus tree gave the following relationships: (Ceropalinae + (Pepsinae + (Ctenocerinae + Pompilinae))).     

Phylogeny of Dendroctonus bark beetles (Coleoptera: Curculionidae: Scolytinae) inferred from morphological and molecular data

Bark beetles in the genus Dendroctonus may attack and kill several species of coniferous trees, some of them causing major economic losses in temperate forests throughout North and Central America. For this reason, they have been widely studied. However, various aspects of the taxonomy and evolutionary history of the group remain contentious. The genus has been subdivided in species groups according to morphological, biological, karyological or molecular attributes, but the evolutionary affinities among species and species groups within the genus remain uncertain. In this study, phylogenetic relationships among Dendroctonus species were reassessed through parsimony‐based cladistic analysis of morphological and DNA sequence data. Phylogenetic inference was based on 36 morphological characters and on mitochondrial DNA sequences of the cytochrome oxidase I (COI) gene. Analyses were carried out for each dataset, as well as for the combined data analysed simultaneously, under equal and implied weights. According to the combined analysis, the genus Dendroctonus is a monophyletic group defined by at least three synapomorphic characters and there are four main lineages of varied composition and diversity within the genus. Within these lineages, several monophyletic groups match, to some extent, species groups defined by previous authors, but certain groups proposed by those authors are polyphyletic or paraphyletic.     

Morphological phylogenetics of termites (Isoptera)

Isoptera (termites) are an ecologically important order, with both a high abundance and biomass in tropical ecosystems. However, there have been few phylogenetic hypotheses for termites, and we present here the first comprehensive cladistic analysis for the group. We analysed relationships between all seven termite families, including representatives of all known feeding group, plus a number of systematically critical taxa. Termite species richness is biased towards the higher termites (Termitidae), and our taxon sampling reflects this. Our analysis was based essentially on morphological characters (96 workers, 93 soldiers) plus seven biological characters. The cladistic analysis gave four equally parsimonious trees, representing two islands of topologies. The strict consensus tree is fully resolved for the higher termites, but less so for the lower termites. Overall there is low statistical support for the suggested topology, and this can be explained by the high incongruence between the data sets (worker, soldier and biological). This study highlights the particular problems of coding morphological characters in social insects with multiple castes. Without the input of additional data sets, e.g. alates, biological, behavioural and molecular, it will not be possible to obtain a well-supported termite phylogeny.