Simultaneous analysis of basal Hymenoptera (Insecta): introducing robust-choice sensitivity analysis

Molecular characters are analysed on their own and in combination with morphological data to examine the phylogenetic relationships of the basal lineages of Hymenoptera ('Symphyta'). This study covers 47 sawfly genera and nine apocritan families and includes molecular sequences from five genes − 12S, 16S, 18S and 28S ribosomal genes and cytochrome oxidase 1 − as well as 343 morphological characters. A robust-choice sensitivity analysis is performed with the data. First, the simultaneous analysis is repeated three times, each time employing a different step matrix for weighting the transformations of the molecular characters. Then, the results of all three simultaneous analyses are summarized in a strict consensus in order to avoid basing the conclusions on a narrow set of assumptions. This methodology is discussed in the paper. The relationships among superfamilies largely confirm previous hypotheses, being (Xyeloidea (Tenthredinoidea s.l. (Pamphilioidea (Cephoidea (Siricoidea (Xiphydrioidea (Orussoidea Apocrita))))))), where Siricoidea is understood as Siricidae+Anaxyelidae. However, the relationships within Tenthredinoidea s.s. proposed here are novel: ({Argidae Pergidae}[ Athalia {(Diprionidae Cimbicidae) Tenthredinidae minus Athalia }]).  © 2003 The Linnean Society of London . Biological Journal of the Linnean Society , 2003, 79, 245–275.     

Phylogeny and classification of the Scopulini moths (Lepidoptera: Geometridae, Sterrhinae)

The phylogenetic relationships of the genera in the geometrid tribe Scopulini (Lepidoptera: Sterrhinae) were examined using 141 characters of adult morphology and ecology. The study material included 92 species, representing all previously recognized genera and covering the morphological variation and full geographical range of the tribe. The cladistic analysis resulted in 20 equally parsimonious trees and a strict consensus cladogram based on these was well resolved. A majority of the recovered synapomorphic characters have been used previously in the taxonomy of the tribe. However, many novel characters were found in the sclerotized structures of the thorax. Many previously recognized genera were found to be nonmonophyletic and based on the present revised, synapomorphy-based classification, the number of recovered genera is reduced considerably. Twenty new generic synonyms and 90 new or revived species combinations are proposed. Seven genera are considered valid, with the large genus Scopula Schrank including over 85% of all species in the tribe. The taxonomic history of the tribe is reviewed and the problems of earlier classifications are discussed. A key to the genera is presented, although an informal diagnosis is preferred. All recognized genera are illustrated and a revised world checklist of the Scopulini is presented.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 143 , 473−530.     

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.     

On the systematic status of the desert plated lizard (Angolosaurus skoogi): phylogenetic inference from DNA sequence analysis of the African Gerrhosauridae

The desert plated lizard ( Angolosaurus skoogi ), a 'sand sea' endemic of the northern Namib Desert, exhibits remarkable morphological convergence with other dune-dwelling lizards worldwide. This distinct ecomorphic condition sets Angolosaurus apart from the remaining genera in the family Gerrhosauridae. Indeed, a morphological phylogeny addressing generic relationships within the Cordyliformes (Cordylidae + Gerrhosauridae) identified Angolosaurus as the earliest diverging taxon among African gerrhosaurids. We re-evaluated the basal status of Angolosaurus , conducting a molecular phylogenetic analysis of the African and Madagascan gerrhosaurid genera. Our survey involved a comprehensive species-level comparison among the four nominal genera of mainland Africa ( Angolosaurus , Cordylosaurus , Tetradactylus and Gerrhosaurus ). Mitochondrial DNA sequence data from the cytochrome b , ND2, 12S and 16S rRNA genes were combined for analysis using both parsimony and maximum likelihood procedures. In contrast to the morphological hypothesis, our results do not depict Angolosaurus as the sister taxon to other African gerrhosaurids. Rather, the molecular analyses consistently place Angolosaurus within Gerrhosaurus , rendering the latter genus paraphyletic.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 78 , 253–261.     

Comparative vegetative anatomy and systematics of the Oncidiinae (Maxillarieae, Orchidaceae)

Subtribe Oncidiinae comprises a vegetatively heterogeneous assemblage of species that has persistently been incapable of organization. Anatomy was considered to be a possible means to resolve the perplexity of relationships amongst the constituent taxa. The consistent occurrence of a foliar hypodermis, homogeneous mesophyll, conical silica bodies in stegmata, and ubiquitous fibre bundles in leaves provides a matrix for linking the taxa, as do the parenchymatous pith and O-thickened endodermal cell walls in roots. However, the strict consensus of the 40 genera studied was completely unresolved, suggesting that vegetative characters alone are insufficient to assess the relationships amongst these taxa, a conclusion also reached for the remainder of Maxillarieae.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 152 , 91–107.     

Phylogenetic relationships of thorny catfishes (Siluriformes: Doradidae) inferred from molecular and morphological data

Doradidae is a putatively monophyletic group of South American freshwater catfishes containing 30 extant genera and 72 valid species. Only one study to date has attempted to estimate phylogenetic relationships among doradids. This morphological analysis partitioned species into two basal genera ( Wertheimeria and Francisodoras ) and a crown group of three subfamilies (Platydoradinae, Astrodoradinae and Doradinae) whose relationships were unresolved. No subsequent work has been done to resolve the subfamilial trichotomy or to assess whether postulated intergeneric relationships are accurate. We address this problem with complete sequences (2.5 kilobases, kb) of mitochondrial 12S and 16S rRNA genes and partial (1.3 kb) sequences of the nuclear elongation factor-1 alpha (EF1α) gene from representatives of 23 doradid genera (43 species) and 13 outgroups from additional siluriform families. Phylogenetic analysis of these data yields strong support for the monophyly of Doradidae and Astrodoradinae (as well as other relationships), but otherwise shows significant conflict with morphological results. A partial re-examination of published morphological data indicates that many characters may have been incorrectly polarized and many taxa have incorrect state assignments. Our results provide a framework for ongoing efforts to describe the species-level diversity of this poorly understood neotropical family.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 140 , 551–575.     

Phylogenetic analysis of Magnoliales and Myristicaceae based on multiple data sets: implications for character evolution

Magnoliales, consisting of six families of tropical to warm-temperate woody angiosperms, were long considered the most archaic order of flowering plants, but molecular analyses nest them among other eumagnoliids. Based on separate and combined analyses of a morphological matrix (115 characters) and multiple molecular data sets (seven variable chloroplast loci and five more conserved genes; 14 536 aligned nucleotides), phylogenetic relationships were investigated simultaneously within Magnoliales and Myristicaceae, using Laurales, Winterales, and Piperales as outgroups. Despite apparent conflicts among data sets, parsimony and maximum likelihood analyses of combined data converged towards a fully resolved and well-supported topology, consistent with higher-level molecular analyses except for the position of Magnoliaceae: Myristicaceae + (Magnoliaceae + (( Degeneria + Galbulimima ) + ( Eupomatia + Annonaceae))). Based on these results, we discuss morphological evolution in Magnoliales and show that several supposedly plesiomorphic traits are synapomorphies of Magnoliineae, the sister group of Myristicaceae (e.g. laminar stamens). Relationships within Annonaceae are also resolved with strong support ( Anaxagorea basal, then ambavioids). In contrast, resolution of relationships within Myristicaceae is difficult and still incomplete, due to a very low level of molecular divergence within the family and a long stem lineage. However, our data provide good evidence that Mauloutchia is nested among other Afro-Malagasy genera, contradicting the view that its androecium and pollen are plesiomorphic  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 125–186.     

A revision of the higher taxonomy of the Afrotropical freshwater crabs (Decapoda: Brachyura) with a discussion of their biogeography

The higher taxonomy of the 20 known genera of Afrotropical freshwater crabs is revised to reflect the evolutionary relationships revealed by the consensus of a series of recent morphological and molecular phylogenetic studies of the group. The Afrotropical freshwater crab genera fall into two monophyletic groups, one from Socotra with two genera (Potamidae) and another that includes the remaining 18 genera. The latter group, which includes the bulk of the region's freshwater crab fauna, forms a well-supported monophyletic clade. We recognize two monophyletic sister groups (subfamilies) within the Potamonautidae, one for seven genera from Africa (the Potamonautinae) and one for 11 genera from Africa, the Seychelles, and Madagascar (the Deckeniinae). The Deckeniinae includes two monophyletic groups (tribes), one with seven genera from Madagascar (the Hydrothelphusini), and one with four genera from Africa and the Seychelles (the Deckeniini). The Deckeniini is further divided here into two subtribes, the Deckeniina and the Globonautina. The Platythelphusidae is not recognized, and the Deckeniidae and Globonautinae are lowered in rank. There is no phylogenetic support for the continued inclusion of any genus from the Afrotropical region in the Gecarcinucidae which is treated here as an exclusively Oriental family. The Afrotropical freshwater crabs (excluding those from Socotra) form a monophyletic assemblage that has no representatives outside of the region. The wider biogeographical implications of the taxonomic revision are discussed.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 399–413.     

Phylogenetic analysis of the Camaenidae (Mollusca: Stylommatophora) with special emphasis on the American taxa

The monophyly of the land snail family Camaenidae has been in doubt due to a disjunct bihemispheric distributional pattern and to the lack of morphological synapomorphies. A cladistic analysis is presented using an ingroup composed of representatives of the three subfamilies distributed in Australia and 52 other species with American distribution. Bradybaenidae, Helicidae and Helminthoglyptidae were used as outgroups. Fifty morphological characters were treated as unordered and analysed using Pee-Wee ver. 2.9, a program for parsimony analysis using implied weights. The results of the analysis support Camaenidae as a monophyletic family (synapomorphies: oval genital orifice, absence of penial sheath). Two of the three Australasian subfamilies, Sinumeloninae and Camaeninae, are monophyletic in the strict consensus tree. The American taxa are classified in eight genera and arranged into two main clades. Caracolus is proposed as the sister group of the American Continental Camaenidae. The genus Solaropsis , previously excluded from this family by different authors, is reassigned to Camaenidae. Shell characters proved to be phylogenetically informative in defining Pleurodonte , Caracolus , Solaropsis , Isomeria and Labyrinthus . © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society, 2003, 138, 449–476.     

Python phylogenetics: inference from morphology and mitochondrial DNA

We used nucleotide sequences from four mitochondrial genes and structural features of the mitochondrial control region, combined with a revised, previously published, morphological data set to infer phylogenetic relationships among the pythons. We aimed to determine which of two competing hypotheses of relationships of the genera Aspidites and Python best explains the evolutionary and bioegeographical history of the family. All analyses of the combined data recover a set of relationships in which (1) the genus Python is paraphyletic with the two east Asian species, P. reticulatus and P. timoriensis , as the sister lineage to the seven Australo-Papuan python genera. We support recognition of a distinct genus for the P. reticulatus  +  P. timoriensis clade; (2) the remaining species of the genus Python form a clade which is the sister lineage to the remainder of the family; (3) the genus Aspidites is embedded among the Australo-Papuan genera. The seemingly primitive characteristics of Aspidites may be better interpreted as reversals or specializations that have accompanied a switch to burrowing in this genus. Resolution of the relationships among the Australo-Papuan lineages is weak, possibly because of rapid diversification early in the history of the radiation. We assessed the tempo of the Indo-Australian python radiation using a maximum likelihood framework based on the birth–death process. We find strong support for elevated speciation rates during the period when Australia collided with the proto-Indonesian archipelago. The data support an origin for pythons outside Australia, followed by a radiation into Australia during the mid-Tertiary.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 603–619.     

Evolutionary systematics of the Indian mouse Mus famulus Bonhote, 1898: molecular (DNA/DNA hybridization and 12S rRNA sequences) and morphological evidence

The genus Mus encompasses 38 species of mice divided into four subgenera: Mus , Pyromys , Nannomys and Coelomys . Each of these four taxa is characterized by discrete morphological as well as biochemical traits. We used two different molecular approaches to determine the relationships between these subgenera: DNA/DNA hybridization and 12S rRNA mitochondrial sequences. We compared the resulting phylogenies from each method and with phylogenies derived from morphological data. The degree of resolution of each molecular approach is discussed. The two molecular studies indicate that Mus , Pyromys , Nannomys and Coelomys are clearly distinct monophyletic groups, as previously indicated by morphological data and other biochemical and molecular approaches. There is one divergence between previous morphological and the molecular and morphological studies presented here: the position of the Indian species Mus famulus . This taxon, which was formerly included in the subgenus Coelomys , is demonstrated here to belong to the subgenus Mus. We also propose the following relationships within Mus sensu lato : Mus and Pyromys are the closest relatives, followed by Nannomys and Coelomys , whose relationships are still unclear. This arrangement is more robustly supported by DNA/DNA hybridization than by 12S rRNA data. A molecular time scale for the evolution within Mus sensu lato is proposed, using as a reference the Mus/Rattus divergence estimated by the fossil record at around 12 mya.  © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society , 2003, 137 , 385–401.     

A new phylogenetic classification for the gymnophthalmid genera Cercosaura , Pantodactylus and Prionodactylus (Reptilia: Squamata)

Because of the poor state of knowledge of many of the gymnophthalmid genera, systematic revision is necessary to render the classification consistent with evolutionary history. To that end, I conducted a review of the species of three genera of the Cercosaurinae which appear to form a monophyletic group: Cercosaura , Pantodactylus , and Prionodactylus . Phylogenetic analysis of 61 morphological characters was conducted after specimens of all species were examined to evaluate the composition of each taxon. The phylogenetic reconstruction suggested that the genus Prionodactylus was paraphyletic. A new phylogenetic classification is proposed that synonymizes Pantodactylus and Prionodactylus with Cercosaura. Cercosaura is redefined to include 11 species and seven subspecies. A key is provided to distinguish among species.  © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society, 2003, 137 , 101−115.     

The evolutionary history of the Chydoridae (Crustacea: Cladocera)

Although much is known about the evolutionary history of the pelagic 'cladocerans', there is little information on benthic families such as the Chydoridae. In this study, we examine the phylogenetic history of 37 chydorid species using sequence variation in two mitochondrial genes, COI and 16S rDNA, and one nuclear gene, 18S rDNA. The four recognized subfamilies of chydorids (Eurycercinae, Saycinae, Aloninae and Chydorinae) were well supported, being separated by large sequence divergences of 14.3–16.4%. By contrast, the existing taxonomic system appears to be less clear at a generic level, since many genera (e.g. Alona , Chydorus , Pleuroxus ) consist of an amalgam of distantly related species. However, among those genera which are monophyletic, levels of divergence are very high, suggesting that they originated somewhere in the mid-Palaeozoic. The factors involved in promoting diversification in this group are discussed.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79, 629–643.     

Morphological analysis, phylogenies and classification in Velloziaceae

A morphological (non-molecular) cladistic analysis of the Velloziaceae is presented. The terminal taxa are 47 species of Velloziaceae plus four taxa as outgroups: Acanthochlamys bracteata (Acanthochlamydaceae), Encholirium scrutor (Bromeliaceae), Thoracocarpus bissectus (Cyclanthaceae) and Pandanus racemosus (Pandanaceae). The species of Velloziaceae sampled represent a significant proportion of the morphological diversity of the family, including all recognized genera and sections. The analysis revealed two major groups within Velloziaceae, supported mainly by stomata, vascular bundles in the pedicel, aquiferous tissues, filaments, anthers, pollen, stigma, seeds, ploidy and chemical characters. Comparison between this and the previous phylogenetic hypotheses for Velloziaceae, together with the two conflicting current classifications, suggests that there is incompatibility concerning the support of current genera and sections. The only three groups supported in all phylogenies are Barbacenioideae sensu Menezes (= Barbacenia sensu Smith & Ayensu), Xerophyta sect. Barbacenioides and Xerophyta sect. Xerophyta sensu Smith & Ayensu.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 157–173.     

Molecular phylogeny of Anomalodesmata (Mollusca: Bivalvia) inferred from 18S rRNA sequences

The origin of the anomalodesmatan bivalves and the relationships of the constituent families are far from being settled. Phylogenetic uncertainties result from the morphological heterogeneity of the Anomalodesmata and from parallel/convergent evolution of several character complexes due to similar life habits. Here, we assess these problems with 26 near-complete anomalodesmatan 18S rRNA sequences from 12 out of 15 families and a selection of heteroconch outgroup taxa. The robustly monophyletic Anomalodesmata share insertions in the V2 and V4 expansion regions. Both parsimony and maximum-likelihood analyses confirm their position among the basal heterodonts rooting between Carditidae and Lucinidae or, together with the latter, between Carditidae and the remaining Heterodonta. There is no support for monophyletic Myoida, nor for a close relationship of Anomalodesmata with any myoid taxon. At the base of the Anomalodesmata is an unstable cluster of long-branch species belonging to the Poromyidae, Verticordiidae, Lyonsiellidae and Thraciidae. The remaining Anomalodesmata split consistently but with varying branch support into three major clades: the Cuspidariidae excluding Myonera ; a 'thraciid' clade consisting of (Euciroidae, ( Myonera ( Thracia, Cleidothaerus , Myochamidae))); and a 'lyonsiid' clade with Laternulidae, Pandoridae, diphyletic Lyonsiidae due to a robust clade of Lyonsia norwegica and the clavagellid Brechites vaginiferus . Tests of various alternative topologies showed that all are significantly longer but optimal likelihood trees with monophyletic carnivorous taxa and/or Thraciidae are not significantly less likely. These results differ greatly from previous morphological studies. Palaeontological data and homology decisions for selected characters are evaluated in the light of the molecular trees.  © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society , 2003, 139 , 229–246.     

A phylogeny of the families of fossil and extant tetraodontiform fishes (Acanthomorpha, Tetraodontiformes), Upper Cretaceous to Recent

A data matrix is presented of 210 morphological characters (mostly osteological, some external) for 20 extant taxa of the ten Recent families of tetraodontiform fishes and 36 fossil tetraodontiforms. The oldest of these are from the Upper Cretaceous (95 Mya); most are from the Lower to Middle Eocene (50–58 Mya). There are two outgroup taxa (a zeiform and a caproid). A cladistic analysis of this matrix for only the extant taxa produced two equally parsimonious trees that call into question the monophyly of some of the previously accepted major higher-level tetraodontiform clades. Inclusion in the analysis of the large number of available fossil taxa helps to resolve relationships between family level clades. The new phylogenetic hypothesis, together with stratigraphic and biogeographical data, is used to discuss scenarios of the origin and evolution of the major clades of the order.  © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society , 2003, 139 , 565−617.     

Weintrauboa, a new genus of pimoid spiders from Japan and adjacent islands, with comments on the monophyly and diagnosis of the family Pimoidae and the genus Pimoa (Araneoidea, Araneae)

The spider genus Weintrauboa new genus (Araneae, Pimoidae) is described to place two species of pimoids from Japan and adjacent islands that were formerly classified in the linyphiid genus Labulla . Weintrauboa contortipes (Karsch) new comb., the type species, and W. chikunii (Oi) new comb. are redescribed. Parsimony analysis of morphological characters provides robust support for the monophyly of the genus Weintrauboa and corroborates the monophyly of Pimoa , Pimoidae, and the clade Linyphiidae plus Pimoidae. New diagnoses for Pimoa and Pimoidae are provided.  © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society , 2003, 139 , 261–281     

The utility of morphological characters in gastropod phylogenetics: an example from the Calyptraeidae

Organismal taxonomy is often based on a single or a small number of morphological characters. When they are morphologically simple or known to be plastic, we may not have great confidence in the taxonomic conclusions of analyses based on these characters. For example, calyptraeid gastropod shells are well known for their simplicity and plasticity, and appear to be subject to frequent evolutionary convergences, but are nevertheless the basis for calyptraeid taxonomy. In a case like this, knowing how the pattern of relationships inferred from morphological features used in traditional taxonomy compares to the patterns of relationships inferred from other morphological characters or DNA sequence data would be useful. In this paper, I examine the relative utility of traditional taxonomic characters (shell characters), anatomical characters and molecular characters for reconstructing the phylogeny of calyptraeid gastropods. The results of an ILD test and comparisons of the recovered tree topologies suggest that there is conflict between the DNA sequence data and the morphological data. Very few of the nodes recovered by the morphological data were recovered by any other dataset. Despite this conflict, the inclusion of morphological data increased the resolution and support of nodes in the topology recovered from a combined dataset. The RIs and CIs of the morphological data on the best estimate topology were not any worse than these indices for the other datasets. This analysis demonstrates that although analyses can be misled by these convergences if morphological characters are used alone, these characters contribute significantly to the combined dataset.  © 2003 The Linnean Society of London . Biological Journal of the Linnean Society , 2003, 78 , 541–593.     

Two new species from Gabon show the need to reduce Commitheca to the synonymy of Pauridiantha (Rubiaceae, Pauridiantheae)

Two new Pauridiantha species (Pauridiantheae, Rubiaceae) from Gabon are described and illustrated. Pauridiantha pleiantha is characterized by a combination of entirely glabrous stems, stipules with a narrow base, and large inflorescences. Pauridiantha smetsiana is characterized by entirely glabrous stems, narrow, early deciduous stipules, leaves red-tinged when dry, and small inflorescences. The two new species, especially P. smetsiana , close the morphological gap between Commitheca and Pauridiantha in combining typical Commitheca characters, such as glabrous stems and a narrow stipular base, with Pauridiantha characters, such as eucamptodromous venation. A detailed morphological comparison (including ovary structure, seed morphology, exotesta anatomy, and pollen morphology) between the two genera is given; in conclusion, Commitheca is relegated to the synonymy of Pauridiantha . The necessary new specific combinations are provided.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 141 , 105–117.     

Phylogeny of the lice (Insecta, Phthiraptera) inferred from small subunit rRNA

There has been much argument about the phylogenetic relationships of the four suborders of lice (Insecta: Phthiraptera). Lyal's study of the morphology of lice indicated that chewing/biting lice (Mallophaga) are paraphyletic with respect to sucking lice (Anoplura). To test this hypothesis we inferred the phylogeny of 33 species of lice from small subunit (SSU) rRNA sequences (18S rRNA). Liposcelis sp. from the Liposcelididae (Psocoptera) was used for outgroup reference. Phylogenetic relationships among the four suborders of lice inferred from these sequences were the same as those inferred from morphology. The Amblycera is apparently the sister-group to all other lice whereas the Rhynchophthirina is apparently sister to the Anoplura; these two suborders are sister to the Ischnocera, i.e. (Amblycera (Ischnocera (Anoplura, Rhynchophthirina))). Thus, the Mallophaga (Amblycera, Ischnocera, Rhynchophthirina) is apparently paraphyletic with respect to the Anoplura. Our analyses also provide evidence that: (i) each of the three suborders of lice that are well represented in our study (the Amblycera, Ischnocera, and Anoplura) are monophyletic; (ii) the Boopiidae is monophyletic; (iii) the genera Heterodoxus and Latumcephalum (Boopiidae) are more closely related to one another than either is to the genus Boopia (also Boopiidae); (iv) the Ricinidae and Laemobothridae may be sister-taxa; (v) the Philopteridae may be paraphyletic with respect to the Trichodectidae; (vi) the genera Pediculus and Pthirus are more closely related to each other than either is to the genus Pedicinus ; and (vii) in contrast to published data for mitochondrial genes, the rates of nucleotide substitution in the SSU rRNA of lice are not higher than those of other insects, nor do substitution rates in the suborders differ substantially from one another.