The adult head structures of Tipulomorpha (Diptera,Insecta) and their phylogenetic implications

Schneeberg, K. and Beutel, R.G. 2011. The adult head structures of Tipulomorpha (Diptera, Insecta) and their phylogenetic implications. —Acta Zoologica (Stockholm) 92 : 316–343. Head structures of adults of Tipula paludosa, Limonia sp. and Trichocera saltator were examined and described. The results are compared with conditions found in other dipterans and other antliophoran groups, notably Nannochoristidae. Several potential synapomorphies of a dipteran–nannomecopteran–siphonapteran clade are present in Tipuloidea and Trichocera, the labro‐epipharyngeal food channel, the loss of the galea and the postpharyngeal pumping apparatus. The sensorial field of the maxillary palpomere 3, a potential dipteran–nannomecopteran synapomorphy, is also present but modified. The presence of M. clypeolabralis, labellae and mandibular stylets are groundplan apomorphies of Diptera, with secondary loss of the mandibles in Tipuloidea, Trichoceridae and many other groups. Tipuloidea is supported by the origin of M. tentorioscapalis anterior on the head capsule, the reduction of M. frontobuccalis anterior and the loss of the ocelli. The reduced tentorium, the origin of two further antennal muscles on the head capsule, the maxillary sensorial field with sensilla in individual pits, the lacking dorsal prelabial concavity and the unpaired salivary channel entering the head are apomorphies of Tipulidae. Closer affinities of Tipulidae and Cylindrotomidae are suggested by pseudotracheae of the advanced type, which have evolved independently in this lineage. The results do neither support a basal placement of Tipuloidea nor close affinities with Brachycera.     

Complete mitochondrial genomes of three Oxya grasshoppers (Orthoptera) and their implications for phylogenetic reconstruction

Oxya is a genus of grasshoppers (Orthoptera: Acridoidea) attacking rice and other gramineous plants in Africa and Asia. In the present study, we characterized complete mitochondrial genomes (mitogenomes) of three species, Oxya japonica japonica (15,427 bp), Oxya hainanensis (15,443 bp) and Oxya agavisa robusta (15,552 bp) collected from China. The three mitogenomes contained a typical gene set of metazoan mitogenomes and shared the same gene order with other Acridid grasshoppers, including the rearrangement of tRNA^Asp and tRNA^Lys. Analyses of pairwise genetic distances showed that ATP8 was the least conserved gene, while COI the most conserved. To determine the position of Oxya grasshoppers in the phylogeny of Acrididae, we reconstructed phylogenetic trees among 64 species from across 11 subfamilies using nucleotide sequences of mitogenomes. While the tree confirms traditional classifications of Acrididae at major higher-levels, it suggests a few modifications for classifications at lower-levels.     

Laying the foundations of evolutionary and systematic studies in crickets (Insecta,Orthoptera): a multilocus phylogenetic analysis

Orthoptera have been used for decades for numerous evolutionary questions but several of its constituent groups, notably crickets, still suffer from a lack of a robust phylogenetic hypothesis. We propose the first phylogenetic hypothesis for the evolution of crickets sensu lato, based on analysis of 205 species, representing 88% of the subfamilies and 71% tribes currently listed in the database Orthoptera Species File (OSF). We reconstructed parsimony, maximum likelihood and Bayesian phylogenies using fragments of 18S, 28SA, 28SD, H3, 12S, 16S, and cytb (~3600 bp). Our results support the monophyly of the cricket clade, and its subdivision into two clades: mole crickets and ant‐loving crickets on the one hand, and all the other crickets on the other (i.e. crickets sensu stricto). Crickets sensu stricto form seven monophyletic clades, which support part of the OSF families, “subfamily groups”, or subfamilies: the mole crickets (OSF Gryllotalpidae), the scaly crickets (OSF Mogoplistidae), and the true crickets (OSF Gryllidae) are recovered as monophyletic. Among the 22 sampled subfamilies, only six are monophyletic: Gryllotalpinae, Trigonidiinae, Pteroplistinae, Euscyrtinae, Oecanthinae, and Phaloriinae. Most of the 37 tribes sampled are para‐ or polyphyletic. We propose the best‐supported clades as backbones for future definitions of familial groups, validating some taxonomic hypotheses proposed in the past. These clades fit variously with the morphological characters used today to identify crickets. Our study emphasizes the utility of a classificatory system that accommodates diagnostic characters and monophyletic units of evolution. Moreover, the phylogenetic hypotheses proposed by the present study open new perspectives for further evolutionary research, especially on acoustic communication and biogeography.     

Genets and 'genet-like' taxa (Carnivora, Viverrinae): phylogenetic analysis, systematics and biogeographic implications

The subfamily Viverrinae is a taxon of uncertain systematic status. This study consists of cladistic analyses based on morphological characters of specimens belonging to the genera Genetta , Osbornictis , Poiana and Prionodon . Two levels of analysis are carried out, one concerning generic relationships (intergeneric analysis) and one dealing with the interrelationships of species within the genus Genetta (intrageneric analysis). In the first analysis, different outgroups were used in order to test the ingroup topology.
With regard to the intergeneric analysis, Osbornictis , Poiana and Prionodon , together with Genetta johnstoni , constitute a monophyletic group (including Nandinia ), which is the sister-group of a clade formed by the other species of genets. Thus, the genus Genetta is regarded as paraphyletic. Prionodon appears to be a derived taxon. The Poiana – Prionodon clade is well supported, especially by ultrastructural hair characters. The cladogram topology in the intrageneric analysis indicates an ecological transition from the rain forest genets to the savanna genets. This supports a rain forest origin of the genus Genetta , a conclusion which may be generalized to the entire study group. © 2002 The Linnean Society of London, Zoological Journal of the Linnean Society , 2002, 134 , 317–334.     

蝗虫分子系统学研究进展

本文从遗传多样性、近缘种鉴别以及分子进化和系统发育重建 3个方面综述了蝗虫分子系统学的研究进展。     

A molecular phylogenetic approach to the systematics of Cylindropuntieae (Opuntioideae,Cactaceae)

Previous taxonomic schemes for the Cylindropuntieae were re‐evaluated in the light of a molecular phylogeny derived from a Bayesian, maximum‐likelihood and parsimony reconstructions with three plastid regions (atpB‐rbcL, psbA‐trnH and trnK/matK data sets). The reconstruction revealed that Corynopuntia and Grusonia as currently defined were polyphyletic. Quiabentia, Micropuntia, Pereskiopsis and Cylindropuntia were the only genera recovered as monophyletic. Grusonia s.s. (only including G. bradtiana) is nested in a polytomy with the rest of the species of Corynopuntia. Grusonia s.l. (G. bradtiana plus Corynopuntia) and Corynopuntia as currently defined are polyphyletic because G. pulchella is sister to Pereskiopsis. Some previous taxonomic proposals for Cylindropuntia and Grusonia recognized polyphyletic series and subgenera that do not conform to the strongly supported monophyletic groups here recovered. This study proposes redefining the polyphyletic Grusonia excluding G. pulchella in order to recognize a strongly supported monophyletic genus and the acceptance of a monotypic Micropuntia (G. pulchella) avoiding a new combination into Pereskiopsis. The infrageneric classification for Grusonia is discussed and the recognition of only four monophyletic strongly supported series (Bigelovianae, Imbricatae, Leptocaules and Ramosissimae) for Cylindropuntia is presented.     

Phylogenetic systematics of Archembiidae (Embiidina, Insecta)

Abstract. A cladistic analysis of the American genera of Embiidae is presented, using fifty‐seven representative taxa and ninety‐four morphological characters. The results support the elevation (and significant re‐delimitation) of the subfamily Archembiinae to family level; as delimited here, Archembiidae, revised status, includes the genera Ecuadembia n.gen., Calamoclostes Enderlein, Archembia Ross, Embolyntha Davis, Xiphosembia Ross, Ochrembia Ross, Dolonembia Ross, Conicercembia Ross, Neorhagadochir Ross, Pachylembia Ross, Rhagadochir Enderlein, Litosembia Ross, Navasiella Davis, Ambonembia Ross, Malacosembia Ross, Biguembia Szumik, Gibocercus Szumik and Pararhagadochir Davis. The results also indicate that some genera recently proposed are unjustified and therefore they are synonymized: Argocercembia Ross (a junior synonym of Embolyntha), Brachypterembia Ross (Neorhagadochir), Scelembia Ross (Rhagadochir), Ischnosembia Ross (Ambonembia) and Aphanembia Ross (Biguembia); all new synonymy. The new genus Ecuadembia is described (type species Archembia arida Ross). Ischnosembia surinamensis (Ross) is returned to the genus Pararhagadochir. The following species synonymies are established: Archembia lacombea Ross 1971 = Archembia kotzbaueri (Navas 1925), Archembia peruviana Ross 2001 = Archembia batesi (MacLachlan 1877), and Conicercembia septentrionalis (Mariño & Márquez 1988) = Conicercembia tepicensis Ross 1984; all new synonymy. The family Archembiidae, and all its constituent genera, are diagnosed and described. The genus Microembia Ross (originally described as an Embiidae) is transferred to Anisembiidae. Pachylembiinae, Scelembiinae, and Microembiinae proposed by Ross are unsupported by the present cladistic analysis. 1     

Thylacocephala (Arthropoda: Crustacea?) from the Cretaceous of Lebanon and implications for thylacocephalan systematics

An intensive study of a collection of arthropods from the Cretaceous of Lebanon, formerly referred to as stomatopod larvae, reveals that these forms belong to the problematical arthropod class Thylacocephala. The species Protozoea hilgendorfi , P. damesi , and Pseuderichthus cretaceus display defining thylacocephalan characters such as a carapace enclosing the entire body bearing a large anterior optic notch; three pairs of large, raptorial appendages; and a posterior battery of small swimming limbs associated with muscle segments. Unique characters of at least the genus Protozoea are the numerous 'pits' covering the entire carapace, elongated anterior rostral and posterior spines, and a dorsal hinge indicative of a truly bivalved carapace. We note several anatomical curiosities that provide some insight into the anatomy, ecology and phylogenetic relationships of these and other thylacocephalan species. Structures on the carapace surface of thylacocephalans may represent sensory organs comparable to the receptors seen on thecostracan crustaceans. The affinities of the three species described are still not entirely clear, but a phylogenetic analysis of the entire group may resolve this.     

Aligned 18S for Zoraptera (Insecta): phylogenetic position and molecular evolution

The order Zoraptera (angel insects) is one of the least known insect groups, containing only 32 extant species. The phylogenetic position of Zoraptera is poorly understood, but it is generally thought to be closely related to either Paraneoptera (hemipteroid orders: booklice, lice, thrips, and bugs), Dictyoptera (blattoid orders: cockroaches, termites, and mantis), or Embioptera (web spinners). We inferred the phylogenetic position of Zoraptera by analyzing nuclear 18S rDNA sequences, which we aligned according to a secondary structure model. Maximum likelihood and Bayesian analyses both supported a close relationship between Zoraptera and Dictyoptera with relatively high posterior probability. The 18S sequences of Zoraptera exhibited several unusual properties: (1) a dramatically increased substitution rate, which resulted in very long branches; (2) long insertions at helix E23; and (3) modifications of secondary structures at helices 12 and 18.     

The function and phylogenetic implications of the tentorium in adult Neuroptera (Insecta)

Despite several recent analyses on the phylogeny of Neuroptera some questions still remain to be answered. In the present analysis we address these questions by exploring a hitherto unexplored character complex: the tentorium, the internal cuticular support structure of the insect head. We described in detail the tentoria of representatives of all extant neuropteran families and the muscles originating on the tentorium using 3D microCT images and analyzed differences in combination with a large published matrix based on larval characters. We find that the tentorium and associated musculature are a source of phylogenetically informative characters. The addition of the tentorial characters to the larval matrix causes a basad shift of the Sisyridae and clearly supports a clade of all Neuroptera except Sisyridae and Nevrorthidae. A sister group relationship of Coniopterygidae and the dilarid clade is further corroborated. A general trend toward a reduction of the dorsal tentorial arms and the development of laminatentoria is observed. In addition to the phylogenetic analysis, a correlation among the feeding habits, the development of the maxillary muscles, and the laminatentoria is demonstrated.     

Preliminary phylogenetic analysis of the Protanisoptera (Insecta: Odonatoptera)

The Permian suborder Protanisoptera (Insecta: Odonatoptera) is revised and a new phylogenetic hypothesis proposed after analyses based on wing venation and different outgroups. After our study the families Camptotaxineuridae and Kaltanoneuridae are excluded from the Protanisoptera. After a new phylogenetic analysis, the family Permaeschnidae is redefined and the families Pholidoptilidae, Polytaxineuridae, Callimokaltaniidae and Hemizygopteridae are restored, as already proposed for the latter three families by Bechly (1996). The new genus Proditaxineura is described. The genus Gondvanoptilon RÖSLER et al., 1981 is excluded from the Meganisoptera: Erasipteridae and re-included in the Permaeschnidae, as already proposed by Bechly (1998). Permaeschna proxima MARTYNOV, 1931 is considered as a junior synonym of Permaeschna dolloi MARTYNOV, 1931. Pholidoptilon camense ZALESSKY, 1931 is excluded from Permaeschna MARTYNOV, 1931 and the genus Pholidoptilon [Zalessky, 1931a] and [Zalessky, 1931b] is restored. Ditaxineurella stigmalisMARTYNOV, 1940 is excluded from the Hemizygopteridae and considered as a Protanisoptera Incertae sedis.     

A phylogenetic analysis of Myriapoda (Arthropoda) using two nuclear protein-encoding genes

Nucleotide and inferred amino acid sequences from two nuclear protein-encoding genes, elongation factor-aα and RNA polymerase II, were obtained from 34 myriapods and 14 other arthropods to determine phylogenetic relationships among and within the myriapod classes. Phylogenetic analyses using maximum parsimony and maximum likelihood methods recovered all three represented myriapod classes (Chilopoda, Diplopoda, Symphyla) and all multiply sampled chilopod and diplopod orders, often with high node support. In contrast, relationships between classes and between orders were recovered less consistently and node support was typically lower. The temporal structure of phylogenetic diversification in Myriapoda may explain this apparent pattern of the phylogenetic recovery.     

Cephalic morphology of Hymenopus coronatus (Insecta: Mantodea) and its phylogenetic implications

External and internal head structures of the mantodean Hymenopus coronatus are examined and described in detail. The results are elaborately compared with the literature. Strong crests on the anterior tentorial arms that articulate with the subantennal suture, a parietal suture and glossae and paraglossae with anteriorly bent tips are proposed as new potential apomorphies for Mantodea while a head capsule being wider than long, enlarged compound eyes, the presence of a frontal shield or scutellum, lateral lobes in the anterior tentorial arms, the presence of a transverse and an interantennal suture and the reduction of the mentum are confirmed as apomorphies, As potential apomorphies for Dictyoptera the reduction of Musculus tentoriobuccalis lateralis (M. 49) is newly presented and a “perforate” tentorium, lacinial incisivi that are located in a galeal pouch and the presence of a postmola are confirmed. The present study shows the value of cephalic morphology for phylogenetic analysis but also points out that further studies including evolutionary key taxa are essential for resolving the evolutionary adaptations among dictyopterans.     

Comparative spermatozoal ultrastructure and molecular analysis in dromiid crabs and their phylogenetic implications for Dromiidae and Podotremata (Decapoda: Brachyura)

We described the spermatozoal ultrastructure and conducted a molecular analysis of Dromiidae Hypoconcha parasitica, Hypoconcha arcuata, Moreiradromia antillensis and Dromia erythropus. To elucidate the relationship between the different species of this brachyuran group, we also compared the spermatozoal morphologies and phylogenetic positioning among species of Dromiidae, Dromioidea and Podotremata. Specimens were collected from the northern coast of São Paulo, Brazil and were fixed and processed followed by transmission electron microscopy and molecular analysis routines. The Dromiidae spermatozoa studied are characterized by a discoidal acrosome, with three or four concentric zones, which are centrally separated by a bilaterally capitate perforatorial chamber, with a “mushroom”-shaped apex in the Hypoconchinae and a “T-shape” in Dromiinae. Above the perforatorial chamber, there is an apical protuberance, continuous with the subopercular region and the operculum, which forms a low, centrally perforated dome. Under differential interference contrast microscopy, the spermatozoa show 3 to 4 radial arms. The spermatozoal characters in Hypoconchinae and Dromiinae do not separate these subfamilies from the Dromiidae and Dromioidea. Ultrastructural differentiation was only found between representative Dromioidea and other Podotremata. Thus, the spermiotaxonomy of these Hypoconcha, Moreiradromia and Dromia species corroborated previous morphological and molecular studies, supporting the monophyly of Dromiidae and Dynomenidae in relation to Homolidae and Latreilliidae.     

Molecular phylogenetic analysis of the Pneumoroidea (Orthoptera, Caelifera): molecular data resolve morphological character conflicts in the basal acridomorpha

A key transition in the evolution of the insect suborder Caelifera (Orthoptera; Insecta) was from predominantly non-angiosperm-feeding basal lineages to the modern acridomorph fauna (grasshoppers and related insects). However, because of conflicts in the distribution of several complex morphological characters, the relationships of the presumed intermediates, and in particular of the superfamily Pneumoroidea, are presently unclear. We undertook a phylogenetic study of representatives of all of the transitional acridomorph families using mitochondrial and nuclear DNA sequences. No support for pneumoroid monophyly was obtained from nonparametric bootstrap analysis. Furthermore, adopting a maximum-likelihood approach, specific hypotheses of relationships within the Pneumoroidea were firmly rejected using parametric bootstrapping and Kishino-Hasegawa tests. The results indicate that the Pneumoroidea are at best a grade. This distinction implies that the evolution of the proposed pneumoroid synapomorphies, femoro-abdominal stridulation and simple male genital structure, might previously have been misinterpreted as cases of single character gains or losses within lineages. Reconstructions of character states for the femoro-abdominal stridulation indicate that, in fact, multiple losses or gains are equally likely. An important implication of our findings is that, in grasshoppers, auditory tympana may have evolved before stridulation, supporting the argument that the original function of tympana may have been related not to conspecific communication but to predator detection. Overall, the results of this study emphasize the high information content of these minor groups (in this case, the four intermediate families under consideration contain only 0.2% of extant orthopteran species diversity). Our analyses also demonstrate the advantages of model-based methods in analyzing systematic problems and, in particular, of the importance of testing specific phylogenetic hypotheses when a priori support for groupings (e.g., from nonparametric bootstrapping) is marginal.     

A combined morphological and molecular phylogenetic analysis of Parthenocissus (Vitaceae) and taxonomic implications

Parthenocissus (the Virginia creeper genus, Vitaceae) consists of 13 species and shows a disjunct distribution between Asia and North America. We investigated the inflorescence structure, calyx morphology, appendages on the inner side of petals, leaf epidermis, pollen and seed characters throughout the genus. A combined phylogenetic analysis with 27 morphological and 4137 molecular characters was conducted and the result was largely congruent with that of the previous molecular work, but with higher resolution. The combined analysis identified two clades corresponding to the Asian and North American taxa. Parthenocissus feddei was resolved as closely related to the clade containing P. cuspidifera, P. heterophylla and P. semicordata. The four species share synapomorphies of having conspicuously raised veinlets, an obscurely five‐ (to eight‐) lobed calyx, appendages on the inner side of petals covering the entire length of anthers and foveolate pollen exine ornamentation. Within the Old World clade, the pentafoliolate species were weakly supported as more closely related to species with both simple and trifoliolate leaves. Furthermore, the ancestral states of tendril apices, inflorescence structure, appendages on the inner side of petals and exine ornamentation of pollen grains were reconstructed on the molecular strict consensus tree. The appendages on the inner side of petals and exine ornamentation of pollen grains were suggested to be important characters in the taxonomy of Parthenocissus, especially for species with trifoliolate leaves. Finally, the previous classifications of Parthenocissus were evaluated within the phylogenetic framework. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, ?? , ??–??.