Flexible ovipositor sheaths in parasitoid Hymenoptera (Insecta)

The structure of the ovipositor sheaths is examined in a number of parasitic wasps as well as a few non-parasitic taxa. Parasitic wasps in the families Aulacidae, Braconidae, Ichneumonidae, Megalyridae, and Stephanidae with a very elongate, external ovipositor have the lateral wall of the sheath finely transversely subdivided by narrow furrows. This makes the sheath highly flexible, allowing it to support the ovipositor proper during ovipositing. The taxa having such a flexible sheath all drill into wood to lay their eggs. Support at the tip of the ovipositor is crucial in the initial stages of oviposition. The flexible ovipositor sheath is possibly a very ancient trait, arising prior to the radiation of extant Apocrita. This is corroborated by their presence in fossil Hymenoptera from the Upper Jurassic. The occurrence of short and/or concealed ovipositors with rigid sheaths in many extant apocritan taxa must therefore be assumed to be the result of multiple reversals.     

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.     

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.     

Ovipositor internal microsculpture in the Braconidae (Insecta, Hymenoptera)

Variation in the microsculpture of the ovipositor egg canal, the morphology of the tip of the upper valve, the structure of the valvillus and the apex of the lower valve seals, as revealed by scanning electron microscopy is described for members of 52 genera representing 30 subfamilies of Braconidae. The possible phylogenetic implications of this variation are discussed. Characters supporting monophyly of the cyclostome clade, and of the non-cyclostome subfamilies Agathidinae, Pselaphaninae and Sigalphinae are reported. Functional and phylogenetic implications are discussed and possible convergent evolution of some features associated with endoparasitism are considered in the light of recent molecular systematics findings. © 1998 The Norwegian Academy of Science and Letters     

Global diversity of hymenopterans (Hymenoptera; Insecta) in freshwater

A summary of the known species of aquatic Hymenoptera is presented. In total, 150 species from 11 families are recognized as aquatic (0.13% of the total described species). This number is likely an underestimate, because of the high percentage of undescribed species and a lack of knowledge of host range and behaviour for most species. All aquatic Hymenoptera are parasitoids. Many species have relatively dense pubescence to trap air and elongate, tarsal claws to grip the substrate, when underwater. Most species are known from the Holarctic and Oriental regions, but this is likely an artefact caused by lack of knowledge of other regions of the world. Aquatic behaviour has evolved independently at least 50 times within the order. Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Gross morphology of multiporous plate sensilla in the Hymenoptera (Insecta)

Both external and internal gross morphology of multiporous plate sensilla were investigated throughout the Hymenoptera, with particular reference to nonaculeate families, using SEM. Several scenarios concerning the origin of multiporous plate sensilla are discussed. Shape, attachment, position, and reinforcing structures, were found to vary between the families, and the possible phylogenetic implications of these are discussed.     

Grooming behaviours in the Hymenoptera (Insecta): potential phylogenetic significance

Grooming behaviours from representatives of 36 families of Hymenoptera were video recorded and analysed. Thirty-three distinct types of grooming movements were recognized. The evolutionary pattern of each behaviour is discussed. Some behaviours displayed consistent variation between taxa, and appear to be informative regarding higher level relationships within the order. Putative synapomorphies are reported that support the monophyly of the Apocrita, Cynipoidea, Platygastroidea, Ichneumonoidea, and Chalcidoidea including the Mymaridae.     

New maimetshid wasps in Cretaceous amber from Myanmar (Insecta: Hymenoptera)

The extinct, exclusively Cretaceous wasp family Maimetshidae is newly recorded from Earliest Cenomanian Burmese amber. Two new genera and species are described. Burmaimetsha concava gen. et sp. nov., based on a male and a female, is most similar to Guyotemaimetsha Perrichot, Nel & Néraudeau, from Albian-Cenomanian French amber, but differs in its larger mandibles, distinctly concave face, elongate antennomeres, and forewing with cell [1Rs] smaller and fourth abscissa of Rs shorter. Maimetshasia kachinensis gen. et sp. nov., is based on a male, and is characterized by asymmetric mandibles with two and three teeth, by its forewing venation without cross-vein 2rs-m, with cell [1M] large and trapezoidal, and vein 2Rs + M very short, and by the hind wing without a free apex of Rs. The family was evidently widespread in the Cretaceous, and the new records extend the paleobiogeographical range to the South-East of Eurasia. A discussion about the possible biology of Maimetshidae is provided.     

Morphology of the antenna cleaner in the Hymenoptera with particular reference to non-aculeate families (Insecta)

An electron microscopic survey of antenna cleaner morphology, mainly in non-aculcate Hymenoptera, is presented. Modified, scale- or paddle-shaped setae on the fore basitarsus were found to be widely distributed throughout the order, but were particularly well developed in the Xyeloidea. Megalodontoidea, Blasticotomidae, Siricoidca, Orussoidea, Cephoidea and Chalcidoidea. as well as in the aculeate family Formicidae. A comb of fine setae on the fore basitarsus was present in all Apocrita, with the exception of the Trigonalyoidea, Evaniidae, and some families of Chalcidoidea, but among the symphytan families was present only in the Orussidae. Members of the symphytan family, Anaxyelidae have a distinct line of discrete setae in the same position as the fine comb of the Orussidae and apocritans which we term a protocomb; members of the Cephidae also show an indication of a protocomb, in the form of a line of more widely spaced, erect setae, that could form part of the same transition series. Members of the Trigonalyoidea and of the Evaniidae have no comb of fine setae but do possess one or more rows of highly modified, plate-like structures on the fore basitarsi.     

A comparative study of morphological integration in Apis mellifera (Insecta, Hymenoptera)

Morphometric correlation matrices from 11 Mediterranean and European honey bee races have been compared. The degree of integration, measured by the eigenvalue variance of the correlation matrix or by Cheverud's index of integration, varies considerably between races but covaries neither with overall body size nor with the variance of the respective sample. Hence, the degree of morphological integration does not depend on body size or the level of variability The patterns of morphometric correlation are significantly similar among all races, but some minor differences in the patterns could be detected: The similarity between the correlation matrices is independent of the degree of phylogenetic relatedness between the respective races. In all races characters belonin to the same functional and/or developmental unit (leg, wing, abdominal characters) have Eiter correlations than the average. Most of the variation in the pattern of correlation can be expfained as a side effect of variation in the degree of integration. Races with high levels of integration tend to have hiher correlations between the main groups of characters, and races with low levels of integration tave smaller correlations between the main grous of characters, while the basic pattern remains undisturbed. In summary, the comparative analysis of morphometric correlation matrices reveals a picture of stability with respect to the pattern of integration and of variation in the degree of integration, which is random with respect to body size, degree of variability, and phylogeny.     

A new Indo-Australian genus of Braconinae (Insecta, Hymenoptera, Braconidae)

Bicarinibracon gen.n. (type species Atanyco1us tricolor . Szépligeti from Papua New Guinea and Irian Jaya). Bicarinibracon luteus gen. et. 3p.n. from India and Malaya and B. lepicius gen. et. sp.n. from Thailand and the Philippines are described and illustrated. A key to the specics of Bicarinibracon is provided and its systematic placement is discussed.     

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

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

Phylogeny of the Hymenoptera (Insecta): The state of the art

The Hymenoptera form one of the major insect orders in terms of species diversity and they are also of great ecological and economic importance, notably as pollinators and natural enemies of pests. Phylogenies provide an essential framework for systematic, comparative and evolutionary research on hymenopterans. Here, I summarise what we know about the higher-level phylogeny of the Hymenoptera based on presentations given at a recent symposium on hymenopteran phylogenetics. I identify agreement and conflict among morphological and molecular analyses and suggest important priority areas for future research.     

New Early Cretaceous hymenopterous insects (Insecta: Hymenoptera) from Sierra del Montsec (Spain)

Four new species of Hymenoptera are described from the Early Cretaceous (Lower Barremian) lithographic limestones of the Sierra del Montsec (Lérida Province, NE Spain):Prosyntexis montsecensis n. sp. (Sepulcidae),Manlaya lacabrua n. sp. (Gasteruptiidae),Meiagaster cretaceus n. gen., n. sp. (Bethylonymidae),Angarosphex lithographicus n. sp. (Sphecidae). The Spanish assemblage extends our knowledge of the distribution of the Lower Cretaceous hymenopteran fauna and will throw light on its character in a probably dry environment with seasonal rainfall.     

The intramandibular gland,a novel exocrine structure in ants (Insecta,Hymenoptera)

Summary A novel glandular structure is described within the mandibles of a series of representatives of the Formicidae. It consists of a variable number of secretory cells with their accompanying duct cells, located in the vicinity of the mandibular cuticle. Because of its localization inside the mandibles, we propose to designate this structure as the intramandibular gland of ants. Ultrastructural observations reveal the development of smooth endoplasmic reticulum in the glandular cells. Several electron-lucid and electron-dense inclusions occur in the cytoplasm. Until now, the function of this widespread gland among the Formicidae remains unknown.     

Ichneumon wasps of the Khasurty locality in Transbaikalia (Insecta, Hymenoptera, Ichneumonidae)

One new genus and four new species of ichneumon wasps are described from the Lower Cretaceous of Buryatia (Khasurty locality): Amplicella mininae, sp. nov., A. shcherbakovi, sp. nov., A. bashkuyevi, sp. nov., Khasurtella buriatica, gen. et sp. nov. The species Tanychora sinensis Zhang, 1991 is transferred to the new genus Khasurtella. Similarity of the ichneumonid faunas of Khasurty and the Chinese localities Yixian and Laiyang indicates the age similarity between these localities. Identification key to genera and species of Tanychorinae is given.     

Diversity of Tiphiidae (Insecta: Hymenoptera) in the fragmented Brazilian semi-deciduous Atlantic Forest

The Atlantic Forest is one of the most important areas of biodiversity in the world, but it has been largely replaced with agropastoral areas and at the present only 12.5 % of the original cover remains. Despite the ecological importance of insects, few studies have been used in conservation approaches for the Atlantic Forest, mainly due to a great taxonomic impediment. A group quite ecologically important but deeply neglected includes parasitoid wasps that control a great number of invertebrates, like tiphiid wasps that are parasitoids of underground coleopteran larvae. The present study aimed to estimate Tiphiidae species richness and diversity in 15 patches of a highly fragmented Atlantic Forest region, using factors that drive the diversity pool from a metacommunity, such as immigration and speciation probabilities. The parameters were estimated using the Neutral Biodiversity Theory, which is based on the total ecological equivalence of species at the same trophic level. Diversity values were molded to the area size, the immigration probabilities, and/or the speciation probability. Eight genera and 460 individuals of Thynninae, Myzininae and Tiphiinae were collected. Variation in species richness, estimated by both rarefaction and first-order jackknife methods, was explained by patch size and by immigration and speciation probabilities. These variables also explained the variation in Shannon diversity and species evenness. Variations in species richness and diversity of Tiphiidae are strongly associated with neutral processes, but they are also influenced by forest fragmentation and intensive agricultural activities.     

Phylogeny and classification of the extant basal lineages of the Hymenoptera (Insecta)

The phylogeny of the basal hymenopteran lineages, including representatives of all ‘symphytan’ families, is anal; In total, 236 morphological characters were scored for 44 exemplars, including six outgroup, two xyelic tenthredinoid, five pamphilioid, three cephoid, five ‘siricoid’, one orussid, and six apocritan taxa. The datas analysed with parsimony under equal weights and under implied weights. The monophyly of the Hymenopte strongly supported but the sistergroup of the Hymenoptera cannot be identified with confidence. The relations of the ‘symphytan’ lineages are Xyeloidea +(Tenthredinoidea+ (Pamphilioidea + (Cephoidea + (Ariaxyelic (Siricidae + (Xiphydriidae +(Orussoidea+Apocrita))))))). Many of the relationships between the superfamilies, especially in the basal branching pattern, are only weakly corroborated. The monophyly of most superfamilies is supported, and all may be monophyletic except the ‘Siricoidea’, which is clearly paraphyletic. It is difficult to di whether the Siricidae or the Anaxyelidae are the closest relatives of Xiphydriidae + (Orussoidea + Apocrita). support for the sistergroup relationship between the Orussoidea and the Apocrita is substantial, putative apomorphies being provided by most character systems. There is also good evidence in favour of the monophj the Apocrita. The internal phylogeny of the Tenthredinoidea differs considerably from the results of earlier anal The Blasticotomidae are the sistergroup of the Tenthredinoidea s.s. Relationships at the base of the Tenthredini s.s. are weakly supported. It is uncertain whether the Tenthredinidae are monophyletic or comprise a 1 paraphyletic grade within the Tenthredinoidea s.s. The Diprionidae may be the sistergroup to Cimbicidae +(Argidae+ Pergidae). Most relationships within the Cimbicidae + (Argidae + Pergidae) clade are corroborated, with the exception of the monophyly of the Argidae. It is proposed to elevate the Anaxyelidae the Xiphydriida both to superfamily status. The family‐level classification of the Tenthredinoidea will probably have to be changed, but this must await further clarification of the phylogeny of this superfamily.     

Simultaneous analysis of the basal lineages of Hymenoptera (Insecta) using sensitivity analysis

The first simultaneous analysis of molecular and morphological data of basal hymenopterans that includes exemplars from all families is presented. DNA sequences (of approximately 2000–2700 bp for each taxon) from the nuclear genes 18S and 28S and the mitochondrial genes 16S and CO1 have been sequenced for 39 taxa (four outgroup taxa, 29 symphytans, and six apocritans). These DNA sequences and 236 morphological characters from Vihelmsen [Zool. J. Linnean Soc. 131 (2001) 393] were analyzed separately as well as simultaneously. All analyses were performed on unaligned sequences, using the optimization alignment (= direct optimization) method. Sensitivity analysis sensu Wheeler [Syst. Biol. 44 (1995) 321] was applied by analyzing the data under nine different combinations of analysis parameter values. The superfamily level relationships of basal hymenopterans as proposed by Vilhelmsen [Zool. J. Linnean Soc. 131 (2001) 393] and Ronquist et al. [Zool. Scr. 28 (1999) 13] are mostly confirmed, except that Pamphilioidea is the sister group to Tenthredinoidea s.l. and that Anaxyelidae (i.e., Syntexis libocedrii) and Siricidae are supported as a monophyletic group, partly reestablishing the traditional concept of Siricoidea. The resulting hypothesis that best represents the combined evidence from morphology and DNA sequences is (Xyeloidea (Tenthredinoidea s.l. Pamphilioidea) (Cephoidea (Siricoidea (Xiphydrioidea (Orussidae Apocrita))))), with Siricoidea = Anaxyelidae +Siricidae. The phylogenetic system within Tenthredinoidea s.l., derived from the combined evidence, is (Blasticotomidae (Tenthredinidae including Diprionidae (Cimbicidae (Argidae Pergidae)))).     

New fossils from China elucidating the phylogeny of Praesiricidae (Insecta: Hymenoptera)

A new subfamily of Praesiricidae (Pamphilioidea), Decorisiricinae subfam.n. , is erected based on three new genera: Decorisiricius gen.n. , Limbisiricius gen.n. and Brevisiricius gen.n. Two new species – Decorisiricius patulus gen. et sp.n. and D. longus sp.n. – from the Lower Cretaceous Yixian Formation and three species –Limbisiricius aequalis gen. et sp.n. , Limbisiricius complanatus sp.n. and Brevisiricius partialis gen. et sp.n. – from the Middle Jurassic Jiulongshan Formation, are described. Based on these well‐preserved new fossil specimens and previously published data, the nonmonophyly of Praesiricidae is confirmed and the phylogenetic relationships of species of Praesiricidae are analysed for the first time. Two main clades within Praesiricidae are recognized from the cladistic analysis: Decorisiricinae subfam.n. forms a monophyletic lineage, with the remaining members of Praesiricidae plus Megalodontes (Megalodontesidae) forming its sister group. The two subfamilies Archoxyelydinae and Praesiricinae are discarded with no strong supported synapomorphic characters based on phylogenetic research. A key to all genera of Praesiricidae is provided. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:38D703ED‐127A‐4DB0‐8153‐8D78AF4AC212 .