Phylogeny of the Dictyoptera

Abstract. Relationships among six key dictyopteran taxa (Mantodea; Blattodea (excluding Cryptocercidae); Cryptocercidae; Mastotermes darwiniensis , Termopsidae and Kalotermitidae [Isoptera]) are analysed based on seventy morphological, developmental and behavioural characters. The fossil record and the 'living fossil' genera Cryptocercus, Mastotermes and Archotermopsis are discussed in detail. Exact analysis of the character state matrix by implicit enumeration (Hennig86) resulted in one cladogram, strongly supporting Blattodea + Cryptocercidae as a sister group to Mantodea, with the Isoptera as a sister group to that complex. Arrangements within the termites are equivocal, with Termopsidae and Mastotermes darwiniensis possible as the relatively most primitive element of Isoptera.     

Phylogeny of cockroaches (Insecta,Dictyoptera, Blattodea), with placement of aberrant taxa and exploration of out‐group sampling

We addressed the phylogeny of cockroaches using DNA sequence data from a broad taxon sample of Dictyoptera and other non‐endopterygotan insect orders. We paid special attention to several taxa in which relationships are controversial, or where no molecular evidence has been used previously: Nocticolidae, a family of small, often cave‐dwelling cockroaches, has been suggested to be the sister group of the predaceous Mantodea or of the cockroach family Polyphagidae; Lamproblatta, traditionally placed in Blattidae, has recently been given family status and placed as sister to Polyphagidae; and Saltoblattella montistabularis Bohn, Picker, Klass & Colville, a jumping cockroach, which has not yet been included in any phylogenetic studies. We used mitochondrial (COI + COII and 16S) and nuclear (18S and 28S) genes, and analysed the data using Bayesian inference (BI) and maximum likelihood (ML). Nocticolidae was recovered as sister to Polyphagidae. Lamproblatta was recovered as sister to Blattidae, consistent with the traditional placement (not based on phylogenetic analysis). However, because of the limited support for this relationship and conflict with earlier morphology‐based phylogenetic hypotheses, we retain Lamproblattidae. S. montistabularis was consistently placed as sister to Ectobius sylvestris Poda (Blaberoidea: Ectobinae), indicating that the saltatorial hindlegs of this genus are a relatively recent adaptation. Isoptera was placed within Blattodea as sister to Cryptocercidae. Nocticolidae + Polyphagidae was sister to Isoptera + Cryptocercidae, and Blaberoidea was sister to the remaining Blattodea.     

The phylogeny and classification of Embioptera (Insecta)

A phylogenetic analysis of the order Embioptera is presented with a revised classification based on results of the analysis. Eighty‐two species of Embioptera are included from all families except Paedembiidae Ross and Embonychidae Navás. Monophyly of each of the eight remaining currently recognized families is tested except Andesembiidae Ross, for which only a single species was included. Nine outgroup taxa are included from Blattaria, Grylloblattaria, Mantodea, Mantophasmatodea, Orthoptera, Phasmida and Plecoptera. Ninety‐six morphological characters were analysed along with DNA sequence data from the five genes 16S rRNA, 18S rRNA, 28S rRNA, cytochrome c oxidase I and histone III. Data were analysed in combined analyses of all data using parsimony and Bayesian optimality criteria, and combined molecular data were analysed using maximum likelihood. Several major conclusions about Embioptera relationships and classification are based on interpretation of these analyses. Of eight families for which monophyly was tested, four were found to be monophyletic under each optimality criterion: Clothodidae Davis, Anisembiidae Davis, Oligotomidae Enderlein and Teratembiidae Krauss. Australembiidae Ross was not recovered as monophyletic in the likelihood analysis in which one Australembia Ross species was recovered in a position distant from other australembiids. This analysis included only molecular data and the topology was not strongly supported. Given this, and because parsimony and the Bayesian analyses recovered a strongly supported clade including all Australembiidae, we regard this family also as monophyletic. Three other families – Notoligotomidae Davis, Archembiidae Ross and Embiidae Burmeister, as historically delimited – were not found to be monophyletic under any optimality criterion. Notoligotomidae is restricted here to include only the genus Notoligotoma Davis with a new family, Ptilocerembiidae Miller and Edgerly, new family, erected to include the genus Ptilocerembia Friederichs. Archembiidae is restricted here to include only the genera Archembia Ross and Calamoclostes Enderlein. The family group name Scelembiidae Ross is resurrected from synonymy with Archembiidae (new status) to include all other genera recently placed in Archembiidae. Embiidae is not demonstrably monophyletic with species currently placed in the family resolved in three separate clades under each optimality criterion. Because taxon sampling is not extensive within this family in this analysis, no changes are made to Embiidae classification. Relationships between families delimited herein are not strongly supported under any optimality criterion with a few exceptions. Either Clothodidae Davis (parsimony) or Australembiidae Ross (Bayesian) is the sister to the remaining Embioptera taxa. The Bayesian analysis includes Australembiidae as the sister to all other Embioptera except Clothididae, suggesting that each of these taxa is a relatively plesiomorphic representatative of the order. Oligotomidae and Teratembiidae are sister groups, and Archembiidae (sensu novum), Ptilocerembiidae, Andesembiidae and Anisembiidae form a monophyletic group under each optimality criterion. Each family is discussed in reference to this analysis, diagnostic combinations and taxon compositions are provided, and a key to families of Embioptera is included.     

A global phylogeny of apple snails: Gondwanan origin, generic relationships, and the influence of outgroup choice (Caenogastropoda: Ampullariidae)

Apple snails (Ampullariidae) are a diverse family of pantropical freshwater snails and an important evolutionary link to the common ancestor of the largest group of living gastropods, the Caenogastropoda. A clear understanding of relationships within the Ampullariidae, and identification of their sister taxon, is therefore important for interpreting gastropod evolution in general. Unfortunately, the overall pattern has been clouded by confused systematics within the family and equivocal results regarding the family's sister group relationships. To clarify the relationships among ampullariid genera and to evaluate the influence of including or excluding possible sister taxa, we used data from five genes, three nuclear and two mitochondrial, from representatives of all nine extant ampullariid genera, and species of Viviparidae, Cyclophoridae, and Campanilidae, to reconstruct the phylogeny of apple snails, and determine their affinities to these possible sister groups. The results obtained indicate that the Old and New World ampullariids are reciprocally monophyletic with probable Gondwanan origins. All four Old World genera, Afropomus, Saulea, Pila, and Lanistes, were recovered as monophyletic, but only Asolene, Felipponea, and Pomella were monophyletic among the five New World genera, with Marisa paraphyletic and Pomacea polyphyletic. Estimates of divergence times among New World taxa suggest that diversification began shortly after the separation of Africa and South America and has probably been influenced by hydrogeological events over the last 90 Myr. The sister group of the Ampullariidae remains unresolved, but analyses omitting certain outgroup taxa suggest the need for dense taxonomic sampling to increase phylogenetic accuracy within the ingroup. The results obtained also indicate that defining the sister group of the Ampullariidae and clarifying relationships among basal caenogastropods will require increased taxon sampling within these four families, and synthesis of both morphological and molecular data. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 98 , 61–76.     

1 Taxon sampling and inferences about diatom phylogeny

Proper taxon sampling is one of the greatest challenges to understanding phylogenetic relationships, perhaps as important as choice of optimality criterion or data type. This has been demonstrated in diatoms where centric diatoms may either be strongly supported as monophyletic or paraphyletic when analyzing SSU rDNA sequences using the same optimality criterion. The effect of ingroup and outgroup taxon sampling on relationships of diatoms is explored for diatoms as a whole and for the order Thalassiosirales. In the latter case, SSU rDNA and rbcL sequence data result in phylogenetic relationships that appear to be strongly incongruent with morphology and broadly incongruent with the fossil record. For example, Cyclotella stelligera Cleve & Grunow behaves like a rogue taxon, jumping from place to place throughout the tree. Morphological data place C. stelligera near the base of the freshwater group as sister to the extinct genus Mesodictyon Theriot and Bradbury, suggesting that it is an old, long branch that might be expected to “misbehave” in poorly sampled trees. Cyclotella stelligera and C. bodanica Grunow delimit the diameter of morphological diversity in Cyclotella, so increased sampling of intermediate taxa will be critical to resolving this part of the tree. Morphology is sampled for a much greater number of taxa and many transitional states of putative synapomorphies seem to suggest a robust morphological hypothesis. The Thalassiosirales are unstable with regards to taxon sampling in the genetic data, suggesting that perhaps the morphological hypothesis is (for now) preferable.     

Termites and trees: a review of recent advances in termite phylogenetics

Summary: Modern termite phylogenetics is critically reviewed, with an emphasis on tree topologies as phylogenetic hypotheses. Studies have especially concentrated on (1) the position of Isoptera among the Dictyoptera and (2) the family group relationships within the Isoptera. The first of these problems is still controversial; although the weight of evidence now suggests that termites are nested within the cockroaches, thus making "Blattaria" as presently constituted paraphyletic. The exact position of termites within the cockroaches is uncertain, although Cryptocercus is the most plausible sister group.¶Family groups relationships are rather better resolved. Mastotermitidae is now generally accepted to be the most basal termite group. Termopsidae, Hodotermitidae and Kalotermitidae are all basal to (Termitidae + Serritermitidae + Rhinotermitidae), although their relative positions within that part of the tree are disputed. Most recent studies support a sister group relationship for Serritermitidae and (Termitidae + Rhinotermitidae). However, no study has yet unambiguously found the Rhinotermitidae monophyletic. The Termitidae are well established as monophyletic and as the most apical termite family. However, within the Termitidae the monophyly of none of the subfamilies is well established, making subfamily level analyses unreliable.¶A number of problem areas are identified: (1) poor taxon sampling is a universal problem, (2) higher taxonomic groupings are often assumed to be monophyletic a priori without adequate support, (3) datasets are collected from different taxa and character systems without consideration of the overall international effort.     

Phylogenetic relationships of Microdontinae (Diptera: Syrphidae) based on molecular and morphological characters

The intrasubfamilial classification of Microdontinae Rondani (Diptera: Syrphidae) has been a challenge: until recently more than 300 out of more than 400 valid species names were classified in Microdon Meigen. We present phylogenetic analyses of molecular and morphological characters (both separate and combined) of Microdontinae. The morphological dataset contains 174 characters, scored for 189 taxa (9 outgroup), representing all 43 presently recognized genera and several subgenera and species groups. The molecular dataset, representing 90 ingroup species of 28 genera, comprises sequences of five partitions in total from the mitochondrial gene COI and the nuclear ribosomal genes 18S and 28S. We test the sister‐group relationship of Spheginobaccha with the other Microdontinae, attempt to elucidate phylogenetic relationships within the Microdontinae and discuss uncertainties in the classification of Microdontinae. Trees based on molecular characters alone are poorly resolved, but combined data are better resolved. Support for many deeper nodes is low, and placement of such nodes differs between parsimony and Bayesian analyses. However, Spheginobaccha is recovered as highly supported sister group in both. Both analyses agree on the early branching of Mixogaster, Schizoceratomyia, Afromicrodon and Paramicrodon. The taxonomical rank in relation to the other Syrphidae is discussed briefly. An additional analysis based on morphological characters only, including all 189 taxa, used implied weighting. A range of weighting strengths (k‐values) is applied, chosen such that values of character fit of the resulting trees are divided into regular intervals. Results of this analysis are used for discussing the phylogenetic relationships of genera unrepresented in the molecular dataset.     

A phylogenetic hypothesis for Asilidae based on a total evidence analysis of morphological and DNA sequence data (Insecta: Diptera: Brachycera: Asiloidea)

An hypothesis of phylogenetic relationships of Asilidae and its constituent taxa is presented, combining morphological and DNA sequence data in a total evidence framework. It is based on 77 robber fly species, 11 Asiloidea outgroup species, 211 morphological characters of the adult fly, and approximately 7300 bp of nuclear DNA from five genes (18S and 28S rDNA, AATS, CAD, and EF-1α protein-encoding DNA). The equally weighted, simultaneous parsimony analysis under dynamic homology in POY resulted in a single most parsimonious cladogram with a cost of 27,582 (iterative pass optimization; 27,703 under regular direct optimization). Six of the 12 included subfamily taxa are recovered as monophyletic. Trigonomiminae, previously always considered as monophyletic based on morphology, is shown to be non-monophyletic. Two of the three Trigonomiminae genera, Holcocephala Jaennicke, 1867 and Rhipidocephala Hermann, 1926, group unexpectedly as the sister taxon to all other Asilidae. Laphriinae, previously seen in the latter position, is the sister group of the remaining Asilidae. Five other subfamily taxa, i.e. Brachyrhopalinae, Dasypogoninae, Stenopogoninae, Tillobromatinae, and Willistonininae, are also shown to be non-monophyletic. The phylogenetic relationships among the higher-level taxa are partly at odds with findings of a recently published morphological study based on more extensive taxon sampling. The total evidence hypothesis is considered as the most informative one, but the respective topologies from the total-evidence, morphology-only, and molecular-only analyses are compared and contrasted in order to discuss the signals from morphological versus molecular data, and to analyze whether the molecular data outcompete the fewer morphological characters. A clade Apioceridae+Mydidae is corroborated as the sister taxon to Asilidae.     

Total‐evidence phylogenetic analysis and reclassification of the Phylinae (Insecta: Heteroptera: Miridae), with the recognition of new tribes and subtribes and a redefinition of Phylini

The subfamily Phylinae (Heteroptera: Miridae) is one of the largest subfamilies of plant bugs and in the most recent classification comprised six tribes: Pilophorini, Hallodapini, Auricillocorini, Phylini, Pronotocrepini, and Leucophoropterini. Phylogenetic analyses of the subfamily using dynamic homology (POY), parsimony (TNT), and model‐based (RAxML) methods are presented. A dataset comprising both morphological and molecular characters (16S, 18S, 28S, and COI–COII) was assembled for taxon samples of 164 ingroup and nine outgroup taxa. A reclassification of the subfamily based on the POY analysis is presented, recognizing nine tribes and nine subtribes. The Auricillocorini is synonymized with the Hallodapini and the Pronotocrepini with the Cremnorrhini; the Phylini was found to be polyphyletic and is redefined; the Semiini and Nasocorini are resurrected and redefined; and the Decomiini and Coatonocapsiniare presented as new tribes. The Hallodapini, rather than the Pilophorini, was found to be the sister‐group to the remaining Phylinae.     

外群选择对隧蜂科（膜翅目：蜜蜂总科）系统重建的影响

外群用于给树附根和推断祖先性状状态。通常,来自内群的姐妹群中的多个分类单元被共同选择作为外群。为了在经验上验证这一方法, 我们采用了3种外群选择策略： 姐妹群中的单一分类单元, 姐妹群中的多个分类单元和连续姐妹群中的多个分类单元。以隧蜂科（膜翅目： 蜜蜂总科）的系统发育重建为例, 我们评估了这3种策略对树拓扑结构的影响, 包括最大似然树、 最大简约树和贝叶斯树。初步结果表明： 相比其他两种策略, 采用姐妹群中的多个分类单元作为外群更有利于系统发育重建得到现已被广泛认可的隧蜂科系统发育关系; 相比最大似然法和贝叶斯法, 虽然隧蜂科系统发育关系没有被很好地解决, 但最大简约法在不同外群选择策略下得到了较为一致的拓扑结构     

Monophyletic Polyneoptera recovered by wing base structure

Phylogenetic relationships among the winged orders of Polyneoptera [Blattodea, Dermaptera, Embiodea (=Embioptera), Isoptera, Mantodea, Orthoptera, Phasmatodea, Plecoptera and Zoraptera] were estimated based on morphological data selected from the hindwing base structure. Cladistic analyses were carried out using hindwing base data alone and in combination with other, more general, morphological data. Both datasets resulted in similar trees and recovered the monophyly of Polyneoptera. Deepest phylogenetic relationships among the polyneopteran orders were not confidently estimated, but the monophyly of Mystroptera (= Embiodea + Zoraptera), Orthopterida (= Orthoptera + Phasmatodea) and Dictyoptera (= Blattodea + Mantodea + Isoptera) was supported consistently. In contrast, placements of Plecoptera and Dermaptera were unstable, although independent analysis of the wing base data supported their sister‐group relationship with two nonhomoplasious synapomorphies (unique conditions in the ventral basisubcostale, and in the articulation between the antemedian notal wing process and first axillary sclerite). Results from the combined wing base plus general morphology data were consistent, even if the wingless orders Grylloblattodea and Mantophasmatodea were included in the analysis. Generally, trees obtained from the present analyses were concordant with the results from other morphological and molecular analyses, but Isoptera were placed inappropriately to be the sister of Blattodea + Mantodea by the inclusion of the wing base data, probably as a result of morphological regressions of the order.     

Beetle or roach: systematic position of the enigmatic Umenocoleidae based on new material from Zhonggou Formation in Jiuquan,Northwest China,and a morphocladistic analysis

The Umenocoleidae is among the most perplexing fossil insect taxa, widespread at lower and middle latitudes during the Early Cretaceous and probably becoming extinct after the mid-Cretaceous. This problematic taxon was first described from the Lower Cretaceous of China and has been variously assigned to stem-group Coleoptera, Protelytroptera, Blattaria, stem-group Dictyoptera, or Mantodea. The systematic position of the enigmatic family — especially its type species Umenocoleus sinuatus Chen et T’an, 1973 — has remained highly controversial owing to the lack of preserved phylogenetically informative or diagnostic characters. Here, we describe a new specimen of the type species from the type horizon of the type locality (Lower Cretaceous Zhonggou Formation at Jiuquan, Yumen City, China), and additionally two forewings of Ponopterix. We found that the Umenocoleidae is similar to stem-group Coleoptera in its forewing venation but differs in its multi-segmented antennae, orthognathous orientation of the head, subdivision of the pronotum by a transverse furrow, ultrastructure of the forewing punctures, and unfolded hindwings. Our phylogenetic analysis, using an extended matrix of 72 characters and 37 taxa, indicates that the Umenocoleidae is likely a specialized taxon of Dictyoptera, sister to Alienoptera. The beetle-like appearance is interpreted to be a result of convergent evolution.     

Phylogeny of the Anomala (Crustacea, Decapoda, Reptantia) based on the ossicles of the foregut

We present a cladistic analysis of the Anomala based on 66 ingroup species and 5 outgroup representatives. Based on a comparative analysis of the morphology of the foregut we scored 124 characters related to size, shape, and fusion of foregut ossicles and other foregut structures. Our parsimony analysis resulted in 30 equally parsimonious trees which differ mainly at the lower hierarchical level. Our study reveals two large clades within Anomala. One large clade consists of Galatheoidea and Chirostyloidea. The internal relationships show a monophyletic Porcellanidae nested within a group comprising paraphyletic Galatheidae, and Munididae as well as Munidopsidae. The other large clade contains Aegla as sister group to a monophyletic group consisting of the Hippoidea and a clade formed by Lomis and the Paguroidea. Coenobitidae are nested within paraphyletic Diogenidae and Lithodidae are nested within paraphyletic Paguridae. The results are discussed in the context of other morphological and molecular analyses. Furthermore, some aspects of carcinization are touched upon; in particular, an anomalan stem species with a, at least to some extent, ventrally folded pleon is suggested.     

An investigation into the cladistic relationships and monophyly of therocephalian therapsids (Amniota: Synapsida)

A comprehensive phylogenetic investigation was performed to elucidate the cladistic relationships and possible monophyly of therocephalian therapsids (Amniota: Synapsida). The phylogenetic positions of 30 therapsid taxa were examined under maximum parsimony, including 23 therocephalian genera. The analysis incorporated 110 cranial and postcranial characters in order to assess the interrelationships of basal therocephalians and eutherocephalians and their relationships to Cynodontia, representing the most complete review of therocephalian phylogeny to date. The analysis supports the hypothesis that Therocephalia represents the monophyletic sister taxon to Cynodontia, with as many as 15 morphological synapomorphies, in contrast with other recent analyses of lesser taxon sampling. The results also support the hypothesis that Scylacosauridae is more closely related to Eutherocephalia than to the basal therocephalian family Lycosuchidae, supporting a ‘Scylacosauria’ clade. The taxa suggested here to be neotenic forms (e.g. Ictidosuchoides and Ictidosuchops) are positioned near the base of a monophyletic Baurioidea. Neotenic development of the therocephalian feeding apparatus and evolutionary parallelism with cynodonts are suggested to have been important trends in the early evolution of baurioid therocephalians into the Late Permian and Early Triassic.     

Phylogenetic analysis of pelecaniformes (aves) based on osteological data: implications for waterbird phylogeny and fossil calibration studies

Background

Debate regarding the monophyly and relationships of the avian order Pelecaniformes represents a classic example of discord between morphological and molecular estimates of phylogeny. This lack of consensus hampers interpretation of the group''s fossil record, which has major implications for understanding patterns of character evolution (e.g., the evolution of wing-propelled diving) and temporal diversification (e.g., the origins of modern families). Relationships of the Pelecaniformes were inferred through parsimony analyses of an osteological dataset encompassing 59 taxa and 464 characters. The relationships of the Plotopteridae, an extinct family of wing-propelled divers, and several other fossil pelecaniforms (Limnofregata, Prophaethon, Lithoptila, ?Borvocarbo stoeffelensis) were also assessed. The antiquity of these taxa and their purported status as stem members of extant families makes them valuable for studies of higher-level avian diversification.

Methodology/Principal Findings

Pelecaniform monophyly is not recovered, with Phaethontidae recovered as distantly related to all other pelecaniforms, which are supported as a monophyletic Steganopodes. Some anatomical partitions of the dataset possess different phylogenetic signals, and partitioned analyses reveal that these discrepancies are localized outside of Steganopodes, and primarily due to a few labile taxa. The Plotopteridae are recovered as the sister taxon to Phalacrocoracoidea, and the relationships of other fossil pelecaniforms representing key calibration points are well supported, including Limnofregata (sister taxon to Fregatidae), Prophaethon and Lithoptila (successive sister taxa to Phaethontidae), and ?Borvocarbo stoeffelensis (sister taxon to Phalacrocoracidae). These relationships are invariant when ‘backbone’ constraints based on recent avian phylogenies are imposed.

Conclusions/Significance

Relationships of extant pelecaniforms inferred from morphology are more congruent with molecular phylogenies than previously assumed, though notable conflicts remain. The phylogenetic position of the Plotopteridae implies that wing-propelled diving evolved independently in plotopterids and penguins, representing a remarkable case of convergent evolution. Despite robust support for the placement of fossil taxa representing key calibration points, the successive outgroup relationships of several “stem fossil + crown family” clades are variable and poorly supported across recent studies of avian phylogeny. Thus, the impact these fossils have on inferred patterns of temporal diversification depends heavily on the resolution of deep nodes in avian phylogeny.     

Proopiomelanocortin (POMC) and testing the phylogenetic position of turtles (Testudines)

Morphological and molecular studies have inferred multiple hypotheses for the phylogenetic relationships of Testudines. The hypothesis that Testudines are the only extant anapsid amniotes and the sister taxon of diapsid amniotes is corroborated by morphological studies, while the hypothesis that Testudines are diapsid amniotes is corroborated by more recent molecular and morphological studies. In this study, the placement of Testudines is tested using the full length cDNA sequence of the polypeptide hormone precursor proopiomelanocortin (POMC). Because only extant taxa have been used, the hypotheses being tested are limited to the following (1) Testudines as the sister taxon of Archosauria, (2) Testudines included in Archosauria and the sister taxon of Crocodilia, (3) Testudines as the sister taxon of Lepidosauria, (4) Testudines as the sister taxon of Sauria, and (5) Testudines as the sister taxon of a monophyletic Mammalia–Sauria clade. Neither Maximum likelihood, Bayesian, or maximum parsimony analyses are able to falsify the hypothesis of (Archosauria (Lepidosauria, Testudines)) and as such is the preferred inference from the POMC data.     

Phylogenetic relationships of family groups in Pentatomoidea based on morphology and DNA sequences (Insecta: Heteroptera)

Phylogenetic relationships within the Pentatomoidea are investigated through the coding and analysis of character data derived from morphology and DNA sequences. In total, 135 terminal taxa were investigated, representing most of the major family groups; 84 ingroup taxa are coded for 57 characters in a morphological matrix. As many as 3500 bp of DNA data are adduced for each of 52 terminal taxa, including 44 ingroup taxa, comprising the 18S rRNA, 16S rRNA, 28S rRNA, and COI gene regions. Character data are analysed separately and in the form of a total evidence analysis. Major conclusions of the phylogenetic analysis include: the concept of Urostylididae is restricted to that of earlier authors; the Saileriolinae is raised to family rank and treated as the sister group of all Pentatomoidea exclusive of Urostylididae sensu stricto; a broadly conceived Cydnidae, as recognized by Dolling, 1981 , is not supported; the placement of Thaumastellidae within the Pentatomoidea is affirmed and the taxon is recognized at family rank rather than as a subfamily of Cydnidae, although its exact phylogenetic position within the Pentatomoidea remains equivocal; the Parastrachiinae is treated as also including Dismegistus Amyot & Serville and placed within a broadly conceived Corimelaenidae, the latter group being treated at family rank; the family‐group taxa Dinidoridae and Tessaratomidae probably represent a monophyletic group, but the recognition of monophyletic subgroups will benefit from additional representation in the sequence data set; and the Lestoniidae is treated as the sister group of the Acanthosomatidae. The Acanthosomatidae and Scutelleridae are consistently recovered as monophyletic. The monophyly of the Pentatomidae appears unequivocal, inclusive of the Aphylinae and Cyrtocorinae, on the basis of morphology, the latter two taxa not being represented in the molecular data set. © The Willi Hennig Society 2008.     

Molecular systematics of the Middle American genus Hypopachus (Anura: Microhylidae)

We present the first phylogenetic study on the widespread Middle American microhylid frog genus Hypopachus. Partial sequences of mitochondrial (12S and 16S ribosomal RNA) and nuclear (rhodopsin) genes (1275 bp total) were analyzed from 43 samples of Hypopachus, three currently recognized species of Gastrophryne, and seven arthroleptid, brevicipitid and microhylid outgroup taxa. Maximum parsimony (PAUP), maximum likelihood (RAxML) and Bayesian inference (MrBayes) optimality criteria were used for phylogenetic analyses, and BEAST was used to estimate divergence dates of major clades. Population-level analyses were conducted with the programs NETWORK and Arlequin. Results confirm the placement of Hypopachus and Gastrophryne as sister taxa, but the latter genus was strongly supported as paraphyletic. The African phrynomerine genus Phrynomantis was recovered as the sister taxon to a monophyletic Chiasmocleis, rendering our well-supported clade of gastrophrynines paraphyletic. Hypopachus barberi was supported as a disjunctly distributed highland species, and we recovered a basal split in lowland populations of Hypopachus variolosus from the Pacific versant of Mexico and elsewhere in the Mesoamerican lowlands. Dating analyses from BEAST estimate speciation within the genus Hypopachus occurred in the late Miocene/early Pliocene for most clades. Previous studies have not found bioacoustic or morphological differences among these lowland clades, and our molecular data support the continued recognition of two species in the genus Hypopachus.     

2 Influence of mixed host populations on success of the parasitic dinoflagellate Amoebophrya

Proper taxon sampling is one of the greatest challenges to understanding phylogenetic relationships, perhaps as important as choice of optimality criterion or data type. This has been demonstrated in diatoms where centric diatoms may either be strongly supported as monophyletic or paraphyletic when analyzing SSU rDNA sequences using the same optimality criterion. The effect of ingroup and outgroup taxon sampling on relationships of diatoms is explored for diatoms as a whole and for the order Thalassiosirales. In the latter case, SSU rDNA and rbcL sequence data result in phylogenetic relationships that appear to be strongly incongruent with morphology and broadly incongruent with the fossil record. For example, Cyclotella stelligera Cleve & Grunow behaves like a rogue taxon, jumping from place to place throughout the tree. Morphological data place C. stelligera near the base of the freshwater group as sister to the extinct genus Mesodictyon Theriot and Bradbury, suggesting that it is an old, long branch that might be expected to "misbehave" in poorly sampled trees. Cyclotella stelligera and C. bodanica Grunow delimit the diameter of morphological diversity in Cyclotella , so increased sampling of intermediate taxa will be critical to resolving this part of the tree. Morphology is sampled for a much greater number of taxa and many transitional states of putative synapomorphies seem to suggest a robust morphological hypothesis. The Thalassiosirales are unstable with regards to taxon sampling in the genetic data, suggesting that perhaps the morphological hypothesis is (for now) preferable.