The preoral cavity of lower Hymenoptera (Insecta): comparative morphology and phylogenetic significance

The skeletal and muscular morphology of the preoral cavity, including the labrum, hypopharynx and labium, was examined in the imago in representatives of all the ‘symphytan’ families as well as the apocritan families Stephanidae, Megalyridae and Trigonalyidae. Xyelidae have complex modifications for masticating pollen, remarkably similiar to those of primitive Lepidoptera. These modifications, collectively termed the triturating basket complex, include an asymmetrical distal epipharyngeal wall with a microtrichial brush and an enlarged infrabuccal pouch with heavy cuticular armature that interacts with the mandibles during feeding. There were striking structural differences between the two subfamilies of Xyelidae in the ligular region; the reduced glossa and clubshaped paraglossae of Macroxyelinae resembles those of primitive Lepidoptera, while the well developed, flattened glossa and paraglossae in Xyelidae are similiar to those of most other ‘Symphyta’. A putative transformation series, leading from a relatively large labrum with unsclerotised distal epipharyngeal wall lying anterior to the mandibles, as seen in Xyelidae and enthredinoidea, to a small and heavily sclerotised labrum and distal epipharyngeal wall lying posterior to the mandibles, as seen in ‘Siricoidea’, Orussidae and the Apocrita, was revealed. These modifications may be adaptations to enable the adult of the families pupating in wood to emerge from the pupal chamber. The Anaxyelidae, Orussidae and Apocrita have similiar configurations of the glossa and nsertions of the ventral premental adductors. This indicates a close affinity of the Anaxyelidae to Orussidae + Apocrita, a hypothesis that is in conflict with other character systems. The Orussidae and Stephanidae share a unique condition in the development of a pair of large apodemes attached to the labrum; this renders the groundplan state of the labrum in the Apocrita uncertain. Twentyfive characters were defined in an attempt to eludicate the ‘Symphyta’–Apocrita transition. A numerical cladistic analysis of the characters was undertaken, resulting in 522 minimum length trees. The characters are also discussed with reference to a cladogram which resulted from an analysis of the characters derived from the present study and a survey of characters from literature.     

Beyond the wasp‐waist: structural diversity and phylogenetic significance of the mesosoma in apocritan wasps (Insecta: Hymenoptera)

A comprehensive data set of hymenopteran mesosomal anatomy is presented and analysed. Eighty‐nine taxa, including three outgroups, were scored for 273 characters. Analyses were carried out under different weighting conditions (equal and implied weights). Topologies retrieved for the non‐apocritan Hymenoptera were highly congruent with previously published results. Apocrita were always retrieved as monophyletic, as were most superfamilies. Relationships amongst apocritan superfamilies were mostly weakly corroborated. Stephanoidea were almost always the sister group to the remaining Apocrita. Evaniomorpha were usually retrieved, Ceraphronoidea being the sister group to Megalyroidea, and Evanioidea to Trigonaloidea. Aculeata did not always come out as monophyletic, and of the aculeate superfamilies, only Apoidea was retrieved. Ichneumonoidea were always monophyletic and often the sister group of Aculeata. Maamingidae and Mymarommatoidea were usually sister groups; together, they often form the sister group of Chalcidoidea. A large clade comprising Cynipoidea, Platygastroidea, and Proctotrupoidea was usually retrieved, the two former superfamilies being nested within Proctotrupoidea. Cynipoidea were usually closely related to some of the Diapriidae. Platygastroidea were usually the sister group of a clade comprising Heloridae, Pelecinidae, Proctotrupidae, and Vanhorniidae. The mesosomal region proved to be a very substantial source of phylogenetically relevant information. The results of the present analyses indicate that a reclassification, especially of Proctotrupoidea, is required, but this should be carried out after thorough analyses of more comprehensive combined data sets. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 159 , 22–194.     

The phylogenetic relationships of the physaloid genera (Solanaceae) based on morphological data

A group of genera, e.g., Chamaesaracha, Leucophysalis, Physaliastrum, Margaranthus, and Withania, in the subfamily Solanoideae (Solanaceae) is centered around the genus Physalis and has been named the physaloid group. It comprises a number of small and often poorly known genera, sometimes seen as united with Physalis and/or each other. A hypothesis of the phylogenetic relationships within this group, based on parsimony analyses of morphological data, is here presented for the first time. The result is discussed in relation to prevailing generic circumsciptions and taxonomic consequences. It is also compared with hypotheses of relationships based on cpDNA data.     

The Labeninae (Hymenoptera: Ichneumonidae): a study in phylogenetic reconstruction and evolutionary biology

The internal phylogenetic relationships of the ichneumonid subfamily Labeninae (Hymenoptera) are re‐assessed using 136 characters for 54 species representing all currently known species‐groups. New character sets are used. These include a new form of seta (named the grossulariform setae) which are present on the inner surface of the hind coxa of some females, and the extensive structural modifications of the terminal segments of the female metasoma, which are described and structural homologies established. The monophyly of the Labeninae and of the four component tribes, Groteini, Poecilocryptini, Xenothyrini and Labenini, is demonstrated. The Groteini is shown to be the sister‐group to the Poecilocryptini + (Xenothyrini + Labenini). Monophyletic genera are established, including three new genera: Ozlabium gen. nov. Gauld & Wahl (type‐species 0. arcadium sp. nov.) in the Groteini from Australia; Torquinsha gen. nov. Gauld & Wahl (type‐species Certonotus invictus Porter) in the Labenini from Chile, and Gauldianus gen. nov. Lanfranco (type‐species Gauldianus chilensis sp. nov. Lanfranco) in the Labenini from Chile. Apechoneura is removed from synonymy and re‐instated as a valid genus for a complex of tropical American species. The biogeographic patterns shown by the Labeninae are discussed. Evidence suggests that the group radiated on Gondwanaland after the separation of Africa/India/Madagascar, but prior to the separation of Australia. Species occurring off the southern continental fragments, in North America, are shown to be members of highly derived lineages, a result consistent with the hypothesis that labenines colonized North America recently through Mesoamerica. Patterns of host utilization and mating behaviour are examined for the group. Members of the more derived lineages show progressively more female specializations for attacking hosts deeply concealed in woody substrates. Males in the more specialized lineages are shown to have a wider array of secondary sexual characteristics and genitalic ornamentations than more plesiomorphic taxa.     

Intestinal coiling pattern in the epilithic algal-feeding cichlids (Pisces, Teleostei) of Lake Tanganyika, and its phylogenetic significance

This paper suggests that, on the basis of ontogenetic change in intestinal coiling pattern, four types and four subtypes could be recognized within 20 species of epilithic algal-feeding cichlids in Lake Tanganyika. The phylogenetically useful anatomical features characterizing each type are as follows. Petrochromis type: torsion of the intestinal tract generally occurs at Stage 5 Intersecting Point (IP), and the haplochromine loop and a two-loop coiling system are observable. Cyathopharynx type: the value of IP becomes 7 before torsion occurs, and the coiling system changes qualitatively from one-loop to two-loop types. Asprotilapia type: torsion occurs at Stage 5 IP, and a one-loop coiling system is retained throughout ontogeny. Telmatochromis type: no torsion occurs, and a one-loop coiling system is retained throughout ontogeny. Within the Petrochromis type, each of the four subtypes is characterized by the following features. Petrochromis subtype: the rostrally directed haplochromine loop has its rostral part situated to the right side of the stomach in the adult stage. Pseudosimochromis subtype: a circular haplochromine loop is present. Eretmodus subtype: the haplochromine loop tends to be prolonged and bends through 90 or 180 further counter-clockwise. Tropheus subtype: torsion occurs at Stage 7 IP.     

The ovipositor apparatus of basal Hymenoptera (Insecta): phylogenetic implications and functional morphology

The skeleto‐musculature of the ovipositor apparatus and the external sculpture of the 1st and 2nd valvulae was studied in representatives from all ‘symphytan’ families. Nineteen informative characters were coded and scored. The distribution of character states are discussed with reference to recent cladistic treatments of the Hymenoptera. Putative autapomorphies of the Hymenoptera are the presence of cordate apodemes on T9 and basal articulations and associated musculature between the 2nd valvifers and the 2nd valvulae. It is a ground plan feature of the order to have the gonocoxites of abdominal segment 8 fused with the gonangula. The configuration of the musculature of the ovipositor apparatus did not display much variation among the taxa examined, except within the Pamphilioidea. There is considerable variation in the external ovipositor sculpture within the Tenthredinoidea. Putative synapomorphies for the tenthredinoid families except the Blasticotomidae are the presence of alternating strongly and weakly sclerotized zones on the first and/or second valvulae and the presence of serrulae on the sawteeth. The presence of transverse rows of large ctenidia on the 1st valvulae is an autapomorphy of the Diprionidae. Fusion of the 2nd valvifers and the 3rd valvulae is a synapomorphy for the Argidae + Pergidae. The ovipositor apparatus of the Pamphilioidea is highly derived, putative autapomorphies being the close association between T9 and the first valvifers, the reduction of the distal parts of the 1st valvulae, and the fusion of the 2nd valvulae for their entire length. The changes in the ovipositor apparatus of Pamphilioidea are associated with a decrease in the amount of work it has to perform during ovipositing, as the eggs are placed predominantly externally on the substrate. The ovipositor apparatus of the ‘Siricoidea’ is enlarged and modified for ovipositing into wood. Putative synapomorphies of the ‘Siricoidea’ + Orussidae + Apocrita are the presence of sawteeth only distally on the ovipositor and elongation of the cordate apodemes of T9.     

地质时期樟科植物花化石及其系统演化意义

该研究对地质时期樟科植物花化石的主要类群,即Androglandula、Lauranthus、Mauldinia、Neusenia、Perseanthus和Potomacanthus属分别从属和种的形态特征、分布、地层以及系统意义进行了论述,并结合现代樟科植物从地层和分布、花序类型、花部形态特征和显微构造特征进行了分析。结果表明:(1)樟科植物在中晚白垩纪期,已经起源于劳亚古陆的中纬度区域。(2)樟科植物的花序类型为Mauldinia属的侧生花序类型和假伞形花序类型。(3)花为3基数的两性花,花被片6枚排列为2轮,雄蕊12或6,排列为4、3或2轮,最内轮雄蕊不育,第三轮雄蕊基部常见一对附属腺体,雄蕊药室瓣裂,4或2药室,雌蕊为单心皮。(4)花被片上常有大量的油细胞、并列型气孔器和单细胞毛。该研究结果中樟科花化石的发现,为樟科植物的系统演化提供了古生物学的证据和资料。     

Plastid phylogenomics improve phylogenetic resolution in the Lauraceae

Abstract The family Lauraceae is a major component of tropical and subtropical forests worldwide, and includes some commercially important timber trees and medicinal plants. However, phylogenetic relationships within Lauraceae have long been problematic due to low sequence divergence in commonly used markers, even between morphologically distinct taxa within the family. Here we present phylogenetic analyses of 43 newly generated Lauraceae plastomes together with 77 plastomes obtained from GenBank, representing 24 genera of Lauraceae and 17 related families of angiosperms, plus nine barcodes from 19 additional species in 18 genera of Lauraceae, in order to reconstruct highly supported relationships for the Lauraceae. Our phylogeny supports the relationships: sisterhood of the Lauraceae and a clade containing Hernandiaceae and Monimiaceae, with Atherospermataceae and Gomortegaceae being the next sister groups, followed by Calycanthaceae. Our results highlight a monophyletic Lauraceae, with nine well‐supported clades as follows: Hypodaphnis clade, Beilschmiedia–Cryptocarya clade, Cassytha clade, Neocinnamomum clade, Caryodaphnopsis clade, Chlorocardium–Mezilaurus clade, Machilus–Persea clade, Cinnamomum–Ocotea clade, and Laurus–Neolitsea clade. The topology recovered here is consistent with the patterns of plastome structural evolution and morphological synapomorphies reported previously. More specifically, flower sex, living type, inflorescence type, ovary position, anther locus number, leaf arrangement, leaf venation, lateral vein number, tree height, and inflorescence location all represent morphological synapomorphies of different lineages. Our findings have taxonomic implications and two new tribes, Caryodaphnopsideae and Neocinnamomeae, are described, and the composition of four other tribes is updated. The phylogeny recovered here provides a robust phylogenetic framework through which to address the evolutionary history of the Magnoliids, the third‐largest group of Mesangiospermae.     

On the head morphology of Phyllium and the phylogenetic relationships of Phasmatodea (Insecta)

Friedemann K., Wipfler B., Bradler S. and Beutel R.G. 2011 . On the head morphology of Phyllium and the phylogenetic relationships of Phasmatodea (Insecta). —Acta Zoologica (Stockholm) 00 : 1–16. External and internal head structures of Phyllium siccifolium are described in detail. The findings are compared with conditions found in other phasmatodeans and members of other neopteran lineages. The compiled 125 characters were analysed cladistically. A clade Eukinolabia (Phasmatodea + Embioptera) was confirmed. Synapomorphies of these two taxa are the shift of the origin of M. tentorioparaglossalis to the hind margin of the prementum, the presence of M. tentorioscapalis medialis, and antennal muscles that originate exclusively on the anterior tentorial arms. Within Eukinolabia, the position of Timema remains somewhat ambiguous because of missing anatomical data. However, it was confirmed as sister group of Euphasmatodea in a monophyletic Phasmatodea. Apomorphic groundplan features of Euphasmatodea are salivary ducts with separate external openings, apically rounded glossae, the presence of the galealobulus, and the reduction of the antennifer. The monophyly of Neophasmatidae was confirmed. Autapomorphies are the loss of M. frontobuccalis posterior, the anteriorly or dorsally directed maxillary palps, and the reduction of the mandibular incisivi. The analysis of characters of the head yielded three new autapomorphies of Phylliinae, the presence of a protuberance on the attachment site of the dorsal tentorial arms, dorsoventrally flattened maxillary‐ and labial palps, and possibly the narrow and U‐shaped field of trichomes on the apical part of the galea.     

The female postabdomen of the enigmatic Nannochoristidae (Insecta: Mecopterida) and its phylogenetic significance

Hünefeld, F. and Beutel, R.G. 2011. The female postabdomen of the enigmatic Nannochoristidae (Insecta: Mecopterida) and its phylogenetic significance. —Acta Zoologica (Stockholm) 00: 1–8. External and internal features of the female postabdomen of Nannochorista neotropica are described in detail. The conditions found in females of Nannochoristidae come closest to the ground plan of Mecopterida. This lineage is characterised by telescoping postabdominal segments, a presumptive autapomorphic feature that is modified in some antliophoran groups, but displayed by the nannochoristid species in a typical manner. More potential autapomorphies of Mecopterida, all present in Nannochoristidae, are the neo‐formation of an intersegmental muscle, a transverse muscle spanning between the genital appendages of segment VIII, a muscle connecting these appendages and the genital chamber and the loss of an intersegmental muscle. Plesiomorphic features of Nannochoristidae are the presence of paired genital appendages on segments VIII and IX. Information on the egg‐depositing substrates of the females is not available. The telescoping postabdomen is suitable for oviposition in soft substrates such as moist soil, or rotten plant materials in the riparian zone, and this is possibly a ground‐plan feature of Mecopterida. The results of recent phylogenetic analyses based on morphological data support a placement of Nannochoristidae in Antliophora, whereas the exact position of the group remains ambiguous. No characters of the female postabdomen were found supporting the monophyly of Mecoptera as conventionally circumscribed, that is Nannochoristidae + Boreidae + Pistillifera.     

Quantitative Aspects of Antenna Grooming in Bees (Apoidea: Hymenoptera)

Antenna grooming in more than 100 species of bees (Apoidea), representing 34 genera of the 7 major bee families is recorded and quantitatively analysed. Most species of bees fall into one of two groups with respect to repetitive antenna cleaning: “Uniscrapers” predominantly clean their antennae with one stroke, “biscrapers” mostly with two subsequent strokes. Uniscrapers are more consequent in their behaviour than biscrapers. Most biscrapers occasionally clean their antenna with one or three strokes. Individual variation in the ratio of stroke repetition is considerably larger in bi- than in uniscrapers. In several species males and females differ with respect to their antenna cleaning behaviour, females tend to be more uniscraping. Most species of the families Colletidae, Halictidae, and Andrenidae, as well as the species of the genera Ceratina and Nomada (Anthophoridae) are biscrapers. Almost all species of Melittidae, Megachilidae, Apidae, and Anthophoridae (except Ceratina and Nomada) are uniscrapers. Bees with an antenna cleaner with ancestral (plesiomorphic) morphology are mostly, but not always, biscrapers, those with a derived antenna cleaner are always uniscrapers. Bees with a derived antenna cleaner perform on average less cleaning actions and strokes than those with an ancestral antenna cleaner. Uniscrapers with an ancestral strigilis do on average more cleaning actions per minute than biscrapers, thus they compensate partly for the fewer number of strokes. But nevertheless the uniscrapers do fewer strokes than the biscrapers (both with an ancestral strigilis). Females clean their antennae on average more often than males. It is interpreted that the behaviour of uniscraping and a derived morphology of the antenna cleaner result in greater efficiency than the status which is ancestral for Apoidea (biscraping and a plesiomorphic antenna cleaner).     

Comparison of sequence-based and structure-based phylogenetic trees of homologous proteins: Inferences on protein evolution

Several studies based on the known three-dimensional (3-D) structures of proteins show that two homologous proteins with insignificant sequence similarity could adopt a common fold and may perform same or similar biochemical functions. Hence, it is appropriate to use similarities in 3-D structure of proteins rather than the amino acid sequence similarities in modelling evolution of distantly related proteins. Here we present an assessment of using 3-D structures in modelling evolution of homologous proteins. Using a dataset of 108 protein domain families of known structures with at least 10 members per family we present a comparison of extent of structural and sequence dissimilarities among pairs of proteins which are inputs into the construction of phylogenetic trees. We find that correlation between the structure-based dissimilarity measures and the sequence-based dissimilarity measures is usually good if the sequence similarity among the homologues is about 30% or more. For protein families with low sequence similarity among the members, the correlation coefficient between the sequence-based and the structure-based dissimilarities are poor. In these cases the structure-based dendrogram clusters proteins with most similar biochemical functional properties better than the sequence-similarity based dendrogram. In multi-domain protein families and disulphide-rich protein families the correlation coefficient for the match of sequence-based and structure-based dissimilarity (SDM) measures can be poor though the sequence identity could be higher than 30%. Hence it is suggested that protein evolution is best modelled using 3-D structures if the sequence similarities (SSM) of the homologues are very low.     

The old wasp and the tree: fossils, phylogeny and biogeography in the Orussidae (Insecta, Hymenoptera)

The phylogenetic relationships of the fossil orussid taxa Mesorussus taimyrensis and Minyorussus luzzi are examined by analysing them together with a large data set compiled previously for the extant Orussidae. The fossils are placed in an unresolved trichotomy with the extant Orussidae. The phylogeny is used for evaluating the hypothesis that the ancestors of the family had reduced body size; the results of this analysis are inconclusive. The biogeographical history of the Orussidae is explored. The common ancestor of the family was probably widespread, the initial splitting events taking place prior to or coinciding with the separation of Laurasia from Gondwana. Later putative vicariance events can be correlated with the gradual breakup of Gondwana. However, the biogeographical history of the Orussidae is dominated by speciation within regions and dispersal. The minimum age of the common ancestor of the Orussidae is >180 Myr when estimated from the biogeographical pattern, >95 Myr when estimated from the phylogenetic position of the fossils; the earlier date is considered to be the most likely.  © 2004 The Linnean Society of London , Biological Journal of the Linnean Society , 2004, 82 , 139–160.     

Johnston's organ in Neodiprion sertifer (Insecta: Hymenoptera)

The fine structure of Johnston's organ in the pine sawfly, Neodiprion sertifer, was studied by electron microscopy to determine if there exists a dimorphism in this organ corresponding to the sexual dimorphism in antennal shape and surface area. The organ is made up of scolopidia that are ultrastructurally similar to those of other insects. The scolopidia, identical in both sexes, comprise three sensory cells bearing two types of sensory processes: Two are shorter and smaller in diameter than the third, which extends into the cuticle of the membrane connecting pedicel and flagellum and terminates at an epicuticular invagination. The dendrites and sensory processes are surrounded by two types of enveloping (glial) cells-a scolopale cell and an attachment cell. Other enveloping cells occur at different levels of the scolopidium. Sexual dimorphism is evident only in the numbers of scolopidial groups: Males have more groups with fewer scolopidia, but both sexes possess about the same total number of scolopidia.     

Isozyme variation and phylogenetic relationships in Vicia subgenus Cracca (Fabaceae)

BACKGROUND AND AIMS: The phylogenetic relationships among 27 vetch species belonging to the subgenus Cracca of the genus Vicia were studied in comparison with three species of Lathyrus section Lathyrus on the basis of isozyme variation. METHODS: Isozymes encoded by 15 putative loci of ten enzymes were resolved by polyacrylamide gel electrophoresis and isozyme variation was analysed by using parsimony and neighbour-joining methods. KEY RESULTS: The analyses revealed 63 parsimony-informative and 36 species-specific orthozymes. Of the latter, 23 are monomophic and are suitable for identification of V. benghalensis, V. palaestina, V. dumetorum, V. pisiformis, V. sylvatica, V. onobrychioides, V. cappadocica, V. cretica, V. articulata, V. tetrasperma, V. ervilia, V. hirsuta and V. loiseleurii. Polymorphism with heterozygous and homozygous isozyme genotypes was found for V. cracca, V. tenuifolia, V. ochroleuca, V. villosa, V. sylvatica, V. cassubica, V. sparsiflora, V. megalotropis, V. altissima, V. onobrychioides, V. cassia, V. cretica and L. heterophyllus, reflecting outcrossing in these species. By contrast, V. benghalensis, V. palaestina, V. disperma, V. dumetorum, V. pisiformis, V. orobus, V. pauciflora, V. tetrasperma and V. loiseleurii had only homozygous isozyme genotypes at polymorphic loci. Isozyme-based phylogenetic trees are presented. CONCLUSIONS: Sections Cracca, Ervum, Pedunculatae and Lenticula of traditional taxonomy are monophyletic groups, whereas sections Oroboideae (= Vicilla) and Panduratae appear polyphyletic and section Cassubicae is split into two species-couples linked at a low level of support. Treatment of ervoid species in a separate subgenus Ervum is not supported because of its polyphyly.     

Sperm structure and ultrastructure in the Hymenoptera (Insecta)

A light and electron microscopical survey of spermatozoan gross morphology and ultrastructure in the Hymenoptera is presented. Details are provided for the first time for members of the families Xyelidae, Argidae, Tenthredinidae, Diprionidae, Cephidae, Figitidae, Proctotrupidae, Diaprii- dae, Heloridae, Eurytomidae, Leucospidae, Perilampidae, Torymidae, Braconidae, Dryinidae, Sphecidae, Pompilidae and Vespidae. Spermatozoan length ranged from 8 μ m in some Braconidae to 500 μm in one chalcidoid. Considerable variation in gross morphology and ultrastructure were observed between taxa. Several phylogenetically informative characters were noted. Very small spermatozoa characterized most of the non-cyclostome subfamilies of Braconidae; spirally twisted axoneme and mitochondrial derivatives occur in the Eulophidae, Eurytomidae and Pteromalidae; spermatozoa with virtually indistinguishable head (nucleus and acrosome) regions characterized the Vespinae and Polistinae. The presence of well-developed spermatodesmata in the vas deferens and seminal vesicle characterize the Symphyta and were largely absent from other groups though they are occasionally present in some bees.     

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.