Phylogeny and the modalities of acoustic diversification in extant Eneopterinae (Insecta, Orthoptera, Grylloidea, Eneopteridae)

Calling with a tegminal stridulatory apparatus is widespread in crickets. However, the evolution of cricket stridulums has been poorly studied and then only on the basis of prephylogenetic models, which are unable to account for the huge diversity recently documented for acoustic features in crickets. The present paper focuses on the evolution of acoustic devices in the subfamily Eneopterinae. This is the first attempt to reconstruct the phylogeny of a large and diverse cricket clade in order to analyze the evolution of emitting structures using precise homology statements. In the first step, we reconstruct the phylogeny of this clade using a morphological data set of 193 characters and 45 taxa. The resultant phylogeny supports the monophyly of the subfamily and that of the 13 genera represented by at least two species in our taxonomic sample. Phylogenetic relationships within the subfamily also support the definition of five tribes: Eurepini, Eneopterini, Nisitrini, Xenogryllini and Lebinthini. In the second step, the evolution of acoustic devices is studied by optimization of venation characters defined on precise homology statements. As hypothesized by previous authors, losses of acoustic communication occur independently in the course of eneopterine evolution; however, they happen abruptly with no intermediate state. Our results also document for the first time the modalities of forewing evolution: the diversification of male forewing venation originates from two processes, a continuous and regular modification process, responsible for slight venation change; and an irregular, more intense punctuated process, allowing the emergence of different venations. This diversification process with sudden changes could be related to the occurrence of acoustic novelties in advertisement calls.     

The complex stridulatory behavior of the cricket Eneoptera guyanensis Chopard (Orthoptera: Grylloidea: Eneopterinae)

Crickets produce stridulated sounds by rubbing their forewings together. The calling song of the cricket species Eneoptera guyanensis Chopard, 1931 alternates two song sections, at low and high dominant frequencies, corresponding to two distinct sections of the stridulatory file. In the present study we address the complex acoustic behavior of E. guyanensis by integrating information on the peculiar morphology of the stridulatory file, the acoustic analysis of its calling song and the forewing movements during sound production. The results show that even if E. guyanensis matches the normal cricket functioning for syllable production, the stridulation involves two different closing movements, corresponding to two types of syllables, allowing the plectrum to hit alternately each differentiated section of the file. Transition syllables combine high and low frequencies and are emitted by a complete forewing closure over the whole file. The double-teeth section of the stridulatory file may be used as a multiplier for the song frequency because of the morphological multiplication due to the double teeth, but also because of an increase of wing velocity when this file section is used. According to available phylogenetic and acoustic data, this complex stridulation may have evolved in a two-step process.     

Functional forewing morphology and stridulation in crickets (Orthoptera, Grylloidea)

Forewing morphology and infrastructure are analysed in three cricket species ( Gryllus campestris, Oecanthus pellucens and Lerneca fuscipennis ) and checked in 20 other genera. A strong dorsoventral asymmetry is demonstrated for the first time. The upper side of the tegmina is covered with hexagonal, more or less high crests forming reliefs. On the lower side, annulate, flat and thick veins exist. The distribution of the hexagons and that of the different vein types is uneven on the forewing surfaces. Their significance in forewing functional morphology is discussed in relation to stridulation and to current knowledge on cricket sound production mechanisms. Their extent and distribution, together with the number and strength of the veins and the surface of wing cells, could account for the vibrational characteristics of the cricket forewing and explain the diverse results obtained by several authors on Gryllus and Oecanthus .     

Chromosomal polymorphism and speciation in the genusEunemobius (Orthoptera: Grylloidea)

Karyotypes are described for the three species of ground crickets belonging to the North American genusEunemobius. All species possess three pairs of metacentric autosomes and a submetacentric X-chromosome. The sex-determination mechanism is XX-XO with 2n=7 in males and 2n=8 in females. There is no evidence that chromosomal rearrangements have been involved in speciation in the group.     

Phylogeny and evolution of Baptistonia (Orchidaceae, Oncidiinae) based on molecular analyses, morphology and floral oil evidences

Baptistonia is endemic to the Brazilian Atlantic Forest and is made up of 23 species, the phylogenetic relationships of which are analysed here. Three plastid regions (psbA-trnH, rpoB-trnC and trnS-G) were sequenced. Using seven inter-simple sequence repeat (ISSR) markers, we scored 183 loci of ISSR dominant markers. Trees were obtained by successively using (a) maximum parsimony and Bayesian analysis of individual and combined sequence data sets, (b) neighbour joining (NJ) and Dollo parsimony of the ISSR data and (c) NJ analysis of the floral oil composition. These approaches, as well as the results of a previous morphological analysis, were combined using a supertree method, yielding clearer relationships within the genus. Various results of our study indicate that the Baptistonia species are recently radiated. B. colorata and B. venusta (previously known as Oncidium trulliferum) are confirmed as being sister to the remaining species of the genus. Four taxonomic groups are clearly inferred. The evolution of some morphological traits and floral oil composition proved to be strongly linked to the phylogeny.     

Phylogeny of Henicopidae (Chilopoda: Lithobiomorpha): a combined analysis of morphology and five molecular loci

Abstract. Relationships in Henicopidae, the dominant southern temperate clade of Lithobiomorpha, are appraised based on parsimony analysis of forty-nine morphological characters and sequence data from five loci (nuclear ribosomal RNAs 18S and 28S, mitochondrial ribosomal RNAs 12S and 16S, protein-coding mitochondrial cytochrome c oxidase I). A combined analysis of these data used direct character optimization, and tested stability of hypotheses through parameter-sensitivity analysis. The morphology dataset highlighted the mandibles as a source of new characters. Morphology, as well as the most congruent parameters for the sequence data and combined analysis, resolved Zygethobiini within Henicopini. Groups retrieved by combined analysis of the sequences and combination with morphology for all parameters include Anopsobiinae/Henicopinae, Lamyctes + Lamyctinus + Henicops , Paralamyctes ( Paralamyctes ), and a clade that groups the southeastern Australian/New Zealand Paralamyctes ( Haasiella ) and P . ( Thingathinga ). Paralamyctes (including Haasiella ) is a Gondwanan clade in the most congruent cladograms based on all molecular data and combination with morphological data. Biogeographic analysis of subtrees for Paralamyctes resolved the interrelationships of Gondwana as (Patagonia ((New South Wales + southeastern Queensland) ((Tasmania) (southern Africa + India) (New Zealand + north Queensland)))).     

Phylogeny of Branchiopoda (Crustacea) based on a combined analysis of morphological data and six molecular loci

The phylogenetic relationships of branchiopod crustaceans have been in the focus of a number of recent morphological and molecular systematic studies. Although agreeing in some respects, major differences remain. We analyzed molecular sequences and morphological characters for 43 branchiopods and two outgroups. The branchiopod terminals comprise all eight “orders”. The molecular data include six loci: two nuclear ribosomal genes (18S rRNA, 28S rRNA), two mitochondrial ribosomal genes (12S rRNA, 16S rRNA), one nuclear protein coding gene (elongation factor 1α), and one mitochondrial protein coding gene (cytochrome c oxidase subunit I). A total of 65 morphological characters were analyzed dealing with different aspects of branchiopod morphology, including internal anatomy and larval characters. The morphological analysis resulted in a monophyletic Phyllopoda, with Notostraca as the sister group to the remaining taxa supporting the Diplostraca concept (“Conchostraca” + Cladocera). “Conchostraca” is not supported but Cyclestheria hislopi is the sister group to Cladocera (constituting together Cladoceromorpha) and Spinicaudata is closer to Cladoceromorpha than to Laevicaudata. Cladocera is supported as monophyletic. The combined analysis under equal weighting gave results in some respects similar to the morphological analysis. Within Phyllopoda, Cladocera, Cladoceromorpha and Spinicaudata + Cladoceromorpha are monophyletic. The combined analysis is different from the morphological analysis with respect to the position of Notostraca and Laevicaudata. Here, Laevicaudata is the sister group to the remaining Phyllopoda and Notostraca is sister group to Spinicaudata and Cladoceromorpha. A sensitivity analysis using 20 different parameter sets (different insertion–deletion [indel]/substitution and transversion/transition ratios) show the monophyly of Anostraca, Notostraca, Laevicaudata, Spinicaudata, Cladoceromorpha, Cladocera, and within Cladocera, of Onychopoda and Gymnomera under all or almost all (i.e., 19 of 20) parameter sets. Analyses with an indel‐to‐transversion ratio up to 2 result in monophyletic Phyllopoda, with Laevicaudata as sister group to the remaining Phyllopoda and with Spinicaudata and Cladoceromorpha as sister groups. Almost all analyses (including those with higher indel weights) result in the same topology when only ingroup taxa are considered. © The Willi Hennig Society 2007.     

Detailed developmental morphology of the spermatophore of the Mediterranean field cricket, Gryllus bimaculatus (De Geer) (Orthoptera: Gryllidae)

The detailed developmental morphology of the spermatophore of a cricket (Gryllus bimaculatus, De Geer) is described for the first time. A specific behavioural reference point is identified from which time developmental processes proceed in a more or less robotic manner. The structures previously identified as forming part of the fully formed spermatophore, that is, the ampulla, the sperm tube and the attachment plate are all identified in detail during their development. Interestingly the attachment plate material is extruded as an amorphous mass from the ejaculatory duct prior to its insertion into the dorsal pouch, the sperm do not enter the sperm sac until the end of spermatophore formation, and the sperm tube itself is formed from a flattened layer of material which rolls up into a tube and involves the use of a die mechanism.     

Ontogeny of the cricket Phaeophilacris bredoides Kaltenbach (Orthoptera, Gryllidae)

Development of Phaeophilacris bredoides Kalt. was studied under stable laboratory conditions: temperature of 26°C, air humidity 60%, and 12L: 12D photoperiod (Knyazev, 1985). The life cycle of Ph. bredoides includes four stages: egg, pronymph, nymph, and adult. The duration of embryonic development is 28 days. The nymphal ontogeny lasts 230 days and consists of 25 instars. The duration of nymphal instars (days) is: 1st—16, 2nd—6, 3rd—8, 4th—10, 5th—10, 6th—15, 7th—10, 8th—8, 9th—9, 10th—11, 11th—11, 12th—8, 13th—9, 14th—10, 15th—7, 16th—9, 17th—7, 18th—7, 19th—7, 20th—10, 21st—11, 22nd—7, 23rd—9, 24th—6, and 25th—9. The duration of adult life is 126 days in males and 125 days in females. Three periods were distinguished in the imaginal ontogeny of males and females: pre-reproductive, reproductive, and post-reproductive. The pre-reproductive period begins with the molt to the adult and ends with the onset of reproductive behavior in males and with the first copulation in females. Its duration is 4 (3–6) days in males and 5 (2–7) days in females. The reproductive period in males starts with the onset of reproductive behavior on the 4th (3rd–6th) day and lasts 119 (98–135) days. In females it begins when they start responding to males’ courtship behavior and lasts 116 (97–133) days. The female reproductive period includes two alternating phases: copulation and egg-laying. The egg-laying phase is initiated by successful copulation. The post-reproductive period in females starts when oviposition ceases and in males, when their reproductive behavior disappears. This period lasts about 3 (2–3) days in males and 4 (2–7) days in females, until the insect dies.     

Phylogeny of Androcymbium (Colchicaceae) based on morphology and DNA sequences

The inclusion of species of Colchicum within Androcymbium in a previous cpDNA phylogeny of the Colchicaceae, questioned the monophyly of Androcymbium, and it was proposed to unite the two genera in Colchicum. Here we expand the previous phylogenetic analyses of Androcymbium by increasing the taxon sampling and adding more data. The analysis include 29 of the 57 species of Androcymbium, more cpDNA (trnL intron, trnL-trnF IGS, trnY-trnD IGS, and trnH-psbA IGS), and nDNA (RNApol2 intron 23) regions, and morphological and life-history traits data. Both parsimony and Bayesian inference were used. According to our data there is no reason to expand Colchicum to include Androcymbium, but we support the inclusion of Bulbocodium and Merendera within Colchicum. Morphology and life history traits are the main arguments in favor of recognizing Androcymbium as a well-circumscribed genus. In the phylogeny two Androcymbium groups are clearly differentiated: (1) one including species from Western and Eastern South Africa, Namibia, and North Africa, and (2) one including species from the north west of South Africa and south of Namibia.     

Phylogeny of Ephemeroptera (mayflies) based on molecular evidence

This study represents the first molecular phylogeny for the Order Ephemeroptera. The analyses included 31 of the 37 families, representing approximately 24% of the genera. Fifteen families were supported as being monophyletic, five families were supported as nonmonophyletic, and 11 families were only represented by one species, and monophyly was not testable. The suborders Furcatergalia and Carapacea were supported as monophyletic while Setisura and Pisciforma were not supported as monophyletic. The superfamilies Ephemerelloidea and Caenoidea were supported as monophyletic while Baetoidea, Siphlonuroidea, Ephemeroidea, and Heptagenioidea were not. Baetidae was recovered as sister to the remaining clades. The mayfly gill to wing origin hypothesis was not supported nor refuted by these data. Mandibular tusks were supported as having at least one loss in Behningiidae and, together with the burrowing lifestyle, possibly two origins. The fishlike body form was supported as plesiomorphic for mayflies with multiple secondary losses. Topological sensitivity analysis was used as a tool to examine patterns concerning the stability of relationships across a parameter landscape, providing additional information that may not have been acquired otherwise.     

Phylogeny of the Brachytheciaceae (Bryophyta) based on morphology and sequence level data

Brachytheciaceae is often considered a taxonomically difficult group of mosses. For example, morphological variation has led to difficulty in generic delimitation. We used DNA sequence data (chloroplast psbT‐H and trnL‐F and nuclear ITS2) together with morphology (63 characters) to examine the relationships within this family. The combined unaligned length of the DNA sequences used in the phylogenetic analyses varied between 1277 and 1343 bp. For phylogeny reconstruction we performed direct optimization, as implemented in POY. Analyses were performed with three different gap costs and the morphological data partition was weighted both: (1) equal to gap cost, and (2) with a weight of one. The utility of sensitivity analysis has recently been cast into doubt; hence in this study it was performed only to explore the effects of weighting on homology statements and topologies and to enable more detailed comparisons between earlier studies utilizing the direct optimization method. The wide sequence length variation of non‐coding ITS2 sequences resulted in character optimizations (i.e., “alignments”) of very different lengths when various gap costs were applied. Despite this variation, the topologies of equally parsimonious trees remained fairly stable. The inclusion of several outgroups, instead of only one, was observed to increase the congruence between data sets and to slightly increase the resolution. An inversion event in the 9 bp loop region in the chloroplast psbT‐N spacer in mosses has been postulated to include only uninformative variation, thus possibly negatively impacting the phylogeny reconstruction. Despite this inversion, its variation within Brachytheciaceae was clearly congruent with information from other sources, but inclusion of these 9 bp in the analysis had only a minor effect on the phylogenetic results. In the most parsimonious topology, which was obtained with equal weighting of all data, Meteoriaceae and Brachytheciaceae were resolved as monophyletic sister groups, which had recently been suggested based on a few shared morphological characters. Our study revealed some new generic relationships within the Brachytheciaceae, which are discussed in light of the morphological characters traditionally used for generic delimitation.     

Ontogenesis of the cricket Gryllus argentinus Sauss. (Orthoptera, Gryllidae)

The development of Gryllus argentinus Sauss. was studied under stable laboratory conditions: the temperature of 26°C, the air humidity of 60%, and the photoperiod of 12h light: 12 h dark. The life cycle of Gryllus argentinus includes four stages: egg, pronymph, nymph, and adult. The duration of embryonic development is 18 days. The depth of egg bedding in the peat is 9.63 ± 0.12 mm (n =145), the clutch containing 2–4 eggs. A female can lay over 1100 viable eggs during the entire oviposition period. Nymphal development includes 9 instars and lasts 97 days. The duration of nymphal instars (days) is: I—5; II—6; III—6; IV—6; V—8; VI—10; VII—13; VIII—14; IX—29. The duration of the adult life is 51 days in males and 69 days, in females. In the imaginal ontogenesis of males and females, three periods can be distinguished: pre-reproductive, reproductive, and postreproductive. Males start to emit the aggressive signal on the 6th (5–8th) day (the pre-reproductive period). They enter the reproductive period (start to emit the calling song) on the 9th (8–13th) day. Females enter the reproductive period (become capable of responding to the calling song and of copulation) on the 9th (8–10th) day. Oviposition starts on the day after the first copulation. The reproductive period lasts about 40 (15–59) days in males and 58 (21–70) days in females. The post-reproductive period starts in females at the moment of finishing the egg laying period and in males, with disappearance of reproductive behavior. The period ends in the animal’s death.     

Phylogeny of the Calliergonaceae (Bryophyta) based on molecular and morphological data

Phylogeny of the Calliergonaceae is reconstructed based on nuclear and chloroplast sequence data, and morphology. Depending on treatment of insertions and deletions, the total number of informative characters was 194 or 163. The study yielded good support for two major clades, one with Hamatocaulis, Scorpidium, and possibly Hygrohypnum ochraceum (Wils.) Loeske, and one with Calliergon, Loeskypnum, Straminergon, and Warnstorfia. Inclusion of H. ochraceum in the ingroup increased tree length, markedly decreased consistency and retention indices, and decreased support for the structure of the Hamatocaulis-Scorpidium clade, and its position within the family is challenged. When H. ochraceum was excluded, good support is found for the genera Hamatocaulis and Scorpidium. Within the other major subclade the monophyly of a clade with Loeskypnum, Straminergon, and Warnstorfia, of a portion of Calliergon, and of Warnstorfia fluitans plus W. pseudostraminea are well supported.     

Phylogeny of ants (Formicidae) based on morphology and DNA sequence data

In order to reconstruct ants' phylogeny, we analysed DNA sequences for two nuclear genes, abdominal-A and Ultrabithorax, from 49 species of ants and two outgroups. As these genes control the development of the first segments of the abdomen in insects, which are very variable in ants (petiole, postpetiole, and gaster constriction), we hypothesized that the morphological variations between the subfamilies may be correlated with mutations of some abd-A or Ubx regions. Contrarily to our hypothesis, these sequences are highly conserved. The differences observed concern mainly third codon positions and present some saturation. Phylogenetic reconstructions were carried out using the genetic raw sequence data and by combining them with a set of morphological data (Total Evidence). Relations among subfamilies of ants remains poorly resolved with molecular data only, but adding these data to morphological characters confirms and reinforce the topology of : a Poneroid complex [Ponerinae, Cerapachyinae, Leptanillinae and army ants], a Formicoid complex [Dolichoderinae, Formicinae] and a Myrmecoid complex [Myrmicinae, Myrmeciinae, Pseudomyrmecinae, Nothomyrmeciinae]. Our molecular results allow resolution near the branch tips and three subfamilies (Dolichoderinae, Formicinae and Pseudomyrmecinae) always appear as monophyletic. The Formicinae and the Dolichoderinae have close relationships. The Camponotini appear as a strong clade inside the Formicinae. The Ponerinae are separated in two parts: the Ectatommini and all other tribes. The Cerapachyinae, Dorylinae, and Ecitoninae belong to the same clade, the Cerapachyinae being confirmed in their subfamily status. The Myrmicinae appears to be very heterogeneous, with the Attini forming a very stable and well-separated group.     

A Phylogenetic Analysis of the Evolution of the Stridulatory Apparatus in True Crickets (Orthoptera, Grylloidea)

The stridulatory apparatus (or stridulum) is currently assumed ancestral in crickets. Models of its subsequent evolution consider only one modality of evolutionary change: the stridulum would have been progressively lost in multiple cricket lineages. A phylogenetic test of this hypothesis is presented here. The morpho-functional types of stridulum have been optimized on the cladistic phylogenies of two monophyletic cricket clades, and parsimonious evolutionary scenarios of the evolution of the stridulum in these clades have been derived. The phylogenetic patterns thus obtained support the hypothesis that the stridulum has been lost several times convergently in crickets. They indicate, however, that the loss of the stridulum could be reversible, and that several modalities of evolutionary change exist for the stridulum. Phylogenetic analysis thus reveals an unsuspected complexity in the evolution of acoustic communication in crickets.     

Phylogeny of the parasitic wasp subfamily Charipinae (Hymenoptera, Cynipoidea, Figitidae)

The Charipinae are a major group of hyperparasitoids of Hemiptera. Here, we present the first cladistic analysis of this subfamily's internal relationships, based on 96 morphological characters of adults. The data matrix was analysed using uniformly weighted parsimony. The effects of using alternative weighting schemes were explored by performing additional searches employing implied weights criteria. One of the caveats of implied weights analysis is that it lacks an objective criterion for selecting the value of the concavity function. In the present study, differential weighting was used to explore the sensitivity of our results to the alternative assumptions made in the analysis and to select one of the most parsimonious trees under equal weights, which we regard as being the hypothesis that minimizes the amount of ad hoc assumptions. The validity of the two existing tribes and the monophyly of all the genera of Charipinae were tested, in particular the cosmopolitan and highly species-rich Alloxysta and Phaenoglyphis , which appear repeatedly in ecological and biochemical studies of host–parasitoid associations. The evolution of several major characters and the relationships between genera are discussed. On the basis of the phylogenetic results, we discuss a number of taxonomic issues. A new classification of the subfamily is proposed in which no tribes are maintained, Carvercharips is synonymyzed with Alloxysta , and the creation of a new genus from Nepal is justified. Our analysis points to the need for a world revision of the basal genus Phaenoglyphis , which is shown as paraphyletic.     

Phylogeny of Aphroditiformia (Polychaeta) based on molecular and morphological data

The phylogeny of Aphroditiformia, benthic polychaetes carrying dorsal elytra, is assessed from nuclear 18S rDNA, mitochondrial cytochrome c oxidase subunit I (COI), and 31 morphological characters. Two non-elytrabearing taxa, Palmyra and Pisione, are included to assess their relationship to the elytrabearers. The data are analysed both separately and combined, with parsimony, maximum likelihood and Bayesian analyses. In total, 19 terminal taxa are examined, including 12 elytrabearing taxa from all scale-worm groups, Palmyra, Pisione, and five outgroup taxa. The results show that Palmyra and Pisione are nested within Aphroditiformia. Palmyra is sister to Aphrodita, and both Pisione and Pholoe are positioned within Sigalionidae, suggesting that both family names Pisionidae and Pholoidae should be treated as junior synonyms of Sigalionidae.