Recent phylogenetic studies on Nematinae based on DNA sequences have shown extensive incongruencies with current nomenclature of genus-group taxa. Here, we expand previous DNA sequence datasets based on three genes (CoI, Cytb, and EF-1α), to include a fourth (NaK) and more genera. The analyses largely confirm the previous findings, particularly the existence of two well-supported large clades, Euura and Pristiphora, together comprising more than 75% of the species of Nematinae. Basal relationships within these two clades remain poorly resolved, mirroring the difficulties in delimiting genera based on morphology. In addition, a moderately supported small clade, Nematus, is found. The relationships between the Euura, Pristiphora, and Nematus clades are uncertain. Therefore, to stabilize the nomenclature we treat these clades as genera. This taxonomic treatment results in numerous new combinations of species names. The following synonymies are
The sawfly genus Euura of the tenthredinid subfamily Nematinae, in which species level taxonomy has long been regarded as controversial, is particularly species rich in northern parts of the Holarctic. Among a majority of species with more or less free-living larvae, a sizeable minority belongs to a monophyletic lineage whose larvae complete their whole development in galls. We present illustrated keys to the adults and galls of 66 gall-inducing Euura species that occur, or might occur, in northern Europe. The distribution of these species is briefly reviewed, with an emphasis on the fauna of Sweden, where 55 species are now definitely recorded, two of them for the first time (E. bigallae, E. myrtilloidica). The species-level taxonomy of gall-inducing Euura remains partly problematic. Nominal species described on the basis of experimentally tested or assumed host plant specificity, but which cannot be recognised using morphological or genetic characters, are treated as conspecific with currently indistinguishable segregates ("host-plant races") associated with other Salix species. 20 new synonymies are proposed (valid names in parentheses): Eupontania acutifoliae baltica Vikberg & Zinovjev, 2006 and Pontania acutifoliae daphnoides Zinovjev, 1993 (Euura acutifoliae (Zinovjev, 1985)), Euura boreoalpina Kopelke, 2001 (Euura lanatae Malaise, 1921), Euura cinereae Kopelke, 1996 preoccupied and Euura lapponica Kopelke, 1996 preoccupied (Euura auritae Kopelke 2000), Euura gemmacinereae Kopelke, 2001 and E. nigritarsis Cameron, 1885 (Euura mucronata (Hartig, 1837)), Euura phylicifoliae Kopelke, 2001 (Euura myrsinifoliae Kopelke, 2001), Nematus westermanni Boheman, 1852 nomen oblitum (Euura scotaspis (Förster, 1854) nomen protectum), Nematus acerosus Hartig, 1840 (Euura saliciscinereae (Retzius, 1783)), Nematus alienatus Förster, 1854 and Phyllocolpa rolleri Liston, 2005 (Euura leucapsis (Tischbein, 1846)), Nematus angustus Hartig, 1837 (Euura atra (Jurine, 1807)), Nematus erythropygus Förster, 1854 (Euura leucosticta (Hartig, 1837)), Nematus impunctatus Herrich-Schäffer, 1840 (Euura amerinae (Linnaeus, 1758)), Pontania carinifrons Benson, 1940 and Phyllocolpa plicaglauca Kopelke, 2007 (Euura destricta (MacGillivray, 1923)), Pontania obscura Kopelke, 2005 (Euura bridgmanii (Cameron, 1883)), Pontania viminalis var. lugubris Enslin, 1918 and Eupontania collactanea rosmarinifoliae Vikberg & Zinovjev, 2006 (Euura collactanea (Förster, 1854)). Euura weiffenbachiella nom. nov. is proposed as a replacement name for Euura weiffenbachii Ermolenko, 1988; preoccupied in Euura by Pteronidea weiffenbachi Lindqvist, 1958 (Euura piliserra (Thomson, 1863)). Lectotypes are designated for the following 9 taxa: Euura insularis Kincaid, 1900, Euura lanatae Malaise, 1921, Euura lappo Malaise, 1921, Euura lappo var. hastatae Malaise, 1921, Nematus acerosus Hartig, 1840, Nematus leptocerus Förster, 1854, Nematus vallisnierii Hartig, 1837, Pontania megacephala Rohwer, 1908, and Pontania piliserra var. mascula Enslin, 1915. Because of secondary homonymy within Euura, the valid name of the Nearctic species E. arctica MacGillivray, 1919 is E. delicatula (MacGillivray, 1919). The Nearctic Euura megacephala is removed from synonymy with the Holarctic E. destricta and treated as a valid species. 34 species names are newly combined with Euura.
Comparative morphology of terminalinstar larvae of Cynipoidea: phylogenetic implications. -Zoologica Scripta , 34 , 15-36. We describe the external morphology of the terminal-instar larvae of 30 species of Cynipoidea (Hymenoptera), with special reference to the head capsule and mouthparts. Twenty-five of the species belong to the Cynipidae and are gall inducers or phytophagous inquilines (guests) in galls, while five represent different insect-parasitic lineages of the Cynipoidea. Although we find only limited variation in body shape, the head sclerites and mandibles offer many characters of potential phylogenetic value. For instance, the mandibles of the parasitoids have one large pointed tooth, with several smaller dents along the inner margin in core figitids, whereas the phytophagous gall inducers and inquilines have mandibles with two or three blunt teeth of subequal size. The mandibles of inquiline larvae are unique in being covered by vertical striations and in having a dominating, broad second tooth. We summarize the qualitative variation among the studied terminal-instar larvae in terms of 33 morphological characters and one life-history trait and examine the phylogenetic implications of these data by running parsimony analyses under uniform character weights and under implied weights (Goloboff weights). The analysis under uniform weights is poorly resolved but the relationships suggested by the implied-weights analysis are largely congruent with previous analyses of adult morphology and molecular data. The larval data support inclusion of the genus Liposthenes in the Neaylax -Isocolus clade, in agreement with the molecular data but in weak conflict with adult morphology. However, the larval data agree with adult morphology and conflict with the molecular data in supporting monophyly of the inquilines.
We describe gross egg morphology and provide the first data on eggshell ultrastructure in cynipoids (Hymenoptera) based on species representing three distinctly different life histories: internal parasitoids of endopterygote larvae, gall inducers and phytophagous inquilines (guests in galls). We then use existing phylogenetic hypotheses to identify putative changes in egg structure associated with evolutionary life-history transitions. We find four major structural changes associated with the shift from parasitoids laying their eggs inside a host larva to gall inducers laying their eggs in or on plants: (1) from a narrow and gradually tapering gross form to a distinct division into a stout body and a long and thin stalk; (2) from a thin to a thick eggshell; (3) from a flexible to a rigid endochorion; and (4) from crystal bundles with shifting orientation in the exochorion to layers of parallel crystal rods. By contrast, we find no major changes in egg structure associated with the transition from gall inducers to inquilines. Comparison between preand post-oviposition eggs of one gall inducer and one inquiline suggests that mechanical stress during the passage through the egg canal gives rise to numerous tiny stress fractures in the boundary separating the exo-and endochorion. In one of the gall inducers, Diplolepis rosae , that end of the egg, which is inserted into the plant, has a specialized and apparently porous shell that may permit chemical exchange between the embryo and the plant. Other structures that could facilitate chemical communication with the host plant through the eggshell were, however, not observed in the eggs of gall inhabitants.
Vårdal, H., Bjørlo, A. & Sæther, O. A. (2002). Afrotropical Polypedilum subgenus Tripodura, with a review of the subgenus (Diptera: Chironomidae). —Zoologica Scripta, 31, 331–402. A subgeneric diagnosis for all stages of the subgenus Tripodura Townes, 1945 of the genus Polypedilum Kieffer, 1912 is given. Nine new Afrotropical species of Tripodura are described: P.(T.)chelum Vårdal sp. n., P.(T.)amplificatus Bjørlo sp. n., P.(T.)patulum Bjørlo sp. n., P.(T.)spinalveum Vårdal sp. n., P.(T.)ewei Bjørlo sp. n., P.(T.)ogoouense Bjørlo sp. n., P.(T.)akani Bjørlo sp. n., P.(T.)dagombae Bjørlo sp. n., and P.(T.)amputatum Bjørlo sp. n.; all as male imagines only. P.(T.)alboguttatum Kieffer, P.(T.)albosignatum Kieffer, P.(T.)tropicum Kieffer, P.(T.)pruina Freeman, P.(T.)quinqueguttatum Kieffer, P.(T.) aegyptium Kieffer, P.(T.) tridens Freeman, P.(T.)allansoni Freeman, P.(T.)longicrus Kieffer, P.(T.)annulatipes Kieffer and P.(T.)abyssiniae Kieffer are re‐described as male and female imagines, while P.(T.)majiis Lehmann, P.(T.)subovatum Freeman, P.(T.)griseoguttatum Kieffer, P.(T.)aferum Lehmann and P.(T.)kijabense Freeman are re‐described as male imagines only. Keys to the male and the known female imagines of the 30 Afrotropical species in the subgenus are presented. A phylogenetic analysis based on all available information on Tripodura from all over the world (135 species) is presented and discussed. The monophyly of the subgenus Tripodura is confirmed. The subgenus can be divided into 20 groups with the acifer group forming the sister group of two larger assemblages of groups in the order acifer (titicacae (ginzansecundum ((aferum (ewei (malickianum (floridense (halterale, pullum)))))) (subovatum (labeculosum ((parascalaenum (allansoni (apfelbecki (udominatum, parvum))))) ((((alboguttatum, aegyptium) quinqueguttatum) annulatipes)). Only in the titicacae, halterale, pullum and apfelbecki groups are the larvae of more than one species described, while one larva is known in each of the subovatum, parascalaenum, aegyptium and quinqueguttatum groups. Three or more pupae are known only from the halterale, pullum, apfelbecki and aegyptium groups. Thus, the tentative nature of the group divisions is obvious. Geographical co‐evolutionary analyses (Brooks parsimony analyses) of the subgenus as a whole and of the major groups are performed and the areas most likely to be part of the original areas estimated. Most probably, eastern South America and Africa were part of the ancestral area. There are multiple sister‐group relationships and generalized tracks between South and East Asia and Africa, between Africa and the Palaearctic region, between South and East Asia, between tropical Brazil and Africa, between East Asia and North America across a former Beringian land bridge, and between the Indo‐West Pacific region and New Zealand, but no evidence for transantarctic relationships.
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