diff --git a/data/0D/B1/E6/0DB1E6C84102FCA6A754C6E48DD838CE.xml b/data/0D/B1/E6/0DB1E6C84102FCA6A754C6E48DD838CE.xml index 61e475762b6..beb36d16f6a 100644 --- a/data/0D/B1/E6/0DB1E6C84102FCA6A754C6E48DD838CE.xml +++ b/data/0D/B1/E6/0DB1E6C84102FCA6A754C6E48DD838CE.xml @@ -1,538 +1,538 @@ - - - -Revision of the European species of Omphale Haliday (Hymenoptera, Chalcidoidea, Eulophidae) + + + +Revision of the European species of Omphale Haliday (Hymenoptera, Chalcidoidea, Eulophidae) - - -Author + + +Author -Hansson, Christer +Hansson, Christer - - -Author + + +Author -Shevtsova, Ekaterina +Shevtsova, Ekaterina -text - - -ZooKeys +text + + +ZooKeys - -2012 - -232 + +2012 + +232 - -1 -157 + +1 +157 - -http://dx.doi.org/10.3897/zookeys.232.3625 + +http://dx.doi.org/10.3897/zookeys.232.3625 -journal article -http://dx.doi.org/10.3897/zookeys.232.3625 -1313-2970-232-1 +journal article +http://dx.doi.org/10.3897/zookeys.232.3625 +1313-2970-232-1 - - - -Omphale Haliday + + + +Omphale Haliday - - -Omphale + + +Omphale Haliday, 1833:339. Type species: -Omphale salicis +Omphale salicis Haliday, by monotypy. - -Smaragdites + +Smaragdites Haliday, 1833:419. Type species: S. admirabilis Haliday, by monotypy. Synonymy by -Graham 1963 +Graham 1963 :240. - -Secodes -Foerster + +Secodes +Foerster , 1856:78, 81. Type species: -Secodes fagi -Foerster +Secodes fagi +Foerster , by monotypy. Synonymy by -Graham 1963 +Graham 1963 :240. - -Holcopelte -Foerster + +Holcopelte +Foerster , 1856:78. Type species: -Elachistus obscurus -Foerster +Elachistus obscurus +Foerster , 1841:40, by original designation. Synonymy by -Hansson 2004 +Hansson 2004 :142. - -Euderomyia + +Euderomyia Girault, 1913:176. Type species: -Euderomyia carlylei +Euderomyia carlylei Girault, by original designation. Synonymy by - -Boucek + +Boucek 1988 :727. - -Chrysocharoideus + +Chrysocharoideus Ashmead, 1904:370. Type species: -Chrysocharis thoracicus +Chrysocharis thoracicus Ashmead, by original designation. Synonymy by -LaSalle and Schauff 1992 +LaSalle and Schauff 1992 :12. - -Chrysocharomyia + +Chrysocharomyia Dodd, in -Girault 1915 +Girault 1915 :207. Type species: -Chrysocharomyia elongata +Chrysocharomyia elongata Girault, by original designation. Synonymy by - -Boucek + +Boucek 1988 :727. - -Paromphale + +Paromphale Girault & Dodd, in -Girault 1915 +Girault 1915 :211-212. Type species: -Paromphale flavicorpus +Paromphale flavicorpus Girault, by original designation. Synonymy by - -Boucek + +Boucek 1988 :727. - -Raphaelonia + +Raphaelonia Girault, 1924:173. Type species: -Raphaelonia sulcatiscutum +Raphaelonia sulcatiscutum Girault, by monotypy. Synonymy by - -Boucek + +Boucek 1988 :727. - -Eugerium + +Eugerium Graham, 1959:202. Type species: -Cirrospilus isander +Cirrospilus isander Walker, by original designation. Synonymy by -Hansson 1996b +Hansson 1996b :5. - -Exodontomphale -Boucek + +Exodontomphale +Boucek , 1984:65. Type species: -Exodontomphale taborskyi -Boucek +Exodontomphale taborskyi +Boucek , by original designation. Synonymy by -Schauff 1991 +Schauff 1991 :61. - -Diagnosis. -Clypeus delimited by grooves at least laterally (e.g. Figs 117, 257, 367, 378); sensilla ampullacea (peglike sensilla) asymmetric; pronotum reduced and usually not visible in dorsal view; occiput without median groove or fold between occipital margin and occipital foramen; male genitalia (Figs 478-501): phallobase with enlarged volsellar setae, paramere with one seta at apex, digitus with two spines (very rarely with one or three spines). The species groups used here are based mainly on the appearance of the male genitalia, and most species have specific characters in this structure. However, since the visualisation of this structure involves slide preparations, characters in the male genitalia have been avoided as much as possible in the identification key, but in a few cases, when this is the only way to separate species, they are used. + +Diagnosis. +Clypeus delimited by grooves at least laterally (e.g. Figs 117, 257, 367, 378); sensilla ampullacea (peglike sensilla) asymmetric; pronotum reduced and usually not visible in dorsal view; occiput without median groove or fold between occipital margin and occipital foramen; male genitalia (Figs 478-501): phallobase with enlarged volsellar setae, paramere with one seta at apex, digitus with two spines (very rarely with one or three spines). The species groups used here are based mainly on the appearance of the male genitalia, and most species have specific characters in this structure. However, since the visualisation of this structure involves slide preparations, characters in the male genitalia have been avoided as much as possible in the identification key, but in a few cases, when this is the only way to separate species, they are used. - -Identification. - + +Identification. + There are no taxonomically or nomenclaturally updated keys to European genera of -Eulophidae +Eulophidae . The most recent keys are in -Graham (1959) +Graham (1959) and -Peck et al. (1964) +Peck et al. (1964) . They are useful for identification by keeping in mind that -Holcopelte +Holcopelte and -Eugerium +Eugerium come out as separate genera in the keys. Another option is to use the key for Nearctic genera of -Chalcidoidea +Chalcidoidea ( -Schauff et al. 1997 +Schauff et al. 1997 ), but -Holcopelte +Holcopelte is treated as a genus separate from -Omphale +Omphale . - - -Description + + +Description . - + Flagellum with 2-3 small and discoid anelli; five flagellomeres, free or with apical 2-3 flagellomeres more or less fused, occasionally with a distinct club with apical flagellomeres fused and wider than proximal flagellomeres (degree of fusion of apical flagellomeres is in many cases difficult to assess, and varies intraspecifically). Sensilla ampullacea (peglike sensilla) asymmetric, long or short. Ventral sense area of male scape reaching along almost entire length of scape. Males with verticillate or scattered arrangement of setae. Mandibles usually endodont with two large teeth at apex and with one to several smaller teeth above large teeth. A few Nearctic species have exodont mandibles, but this feature has not been found in European species. Cly- peus entirely delimited, quadrangular to semicircular or triangular in shape. Lower clypeal margin usually arcuately protruding, in a few species straight. Lower frons usually with a cross-ridge reaching almost from eye to eye (see -Hansson 1996b +Hansson 1996b , fig. 1), cross-ridge is protruding and is not an edge resulting from the collapse of frons above the antennal toruli, cross-ridge missing in some species. Frontal suture between lower and upper frons present. Antennal scrobes either more or less broadly separated and never joining, or joining at or below frontal suture. Surface between antennal scrobes more or less raised to form an interscrobal ridge. Occipital margin rounded off, rarely with an edge or a carina. Without a weak median groove or fold from occipital margin down to occipital foramen. - + Pronotum reduced and hardly visible in dorsal view. Mesoscutum usually with weak delimited notauli; midlobe of mesoscutum with two pair of setae, some non-European species either with one pair or without setae. Scutellum usually without grooves but occasionally with a median groove with variable strength and length, always with one pair of setae usually situated in posterior half. Dorsellum short, smooth, convex or flat, occasionally with strong sculpture. Transepimeral sulcus curved or straight. Propodeum usually smooth and shiny, without anteromedian pit or raised carinae, with or without a narrow transverse proximal groove; propodeal callus with two setae; petiolar foramen semicircular to triangular, usually wide to fit the wide petiole. Forewing with a narrow and bare costal cell; speculum open or closed below; radial cell bare or hairy; with or without one hair line from stigmal vein; postmarginal vein 0.2 --2.1x +-2.1x as long as stigmal vein in European species. Legs usually long and slender. -Petiole short and wide, occasionally as long as wide, pale or dark. Gaster usually subsessile; ovate to lanceolate in female. The male genitalia display important characters useful in classification of the species groups, but also in the separation of species. +Petiole short and wide, occasionally as long as wide, pale or dark. Gaster usually subsessile; ovate to lanceolate in female. The male genitalia display important characters useful in classification of the species groups, but also in the separation of species. - -The species-groups. - -Graham (1963) + +The species-groups. + +Graham (1963) initially divided the European -Omphale +Omphale into four groups with two unplaced species. However, he did so without any motivation and the groups were not diagnosed. The species groups presented here largely follow the groups presented by Graham, but now defined by morphological characters, and due to the addition of species two groups have been added. Characters in the male genitalia are used extensively to group species and to separate these groups. Graham was not aware of such characters and some species have therefore been transferred from their original placement. The European species are here separated into six groups, and with six unplaced species. The unplaced species comprise species that do -not +not fit in any of the species groups, and that do not share features with each other to form species groups on their own. - -Biology. - + +Biology. + Hosts or host plant/fungi associations are known for 16 of the 37 European species (Table 1), and in all cases gall midges ( -Diptera +Diptera : -Cecidomyiidae +Cecidomyiidae ) are the target group. Host records from North America ( -Hansson 1996b +Hansson 1996b ) and Central America ( -Hansson 2004 +Hansson 2004 ) support the findings for European species, i.e. that -Omphale +Omphale species are exclusively parasitoids on gall midges. -Dziurzynski (1961) +Dziurzynski (1961) investigated the biology of -Omphale lugens +Omphale lugens in Poland (given as -Secodes coactus +Secodes coactus in the Dziurzynski publication). -Omphale lugens +Omphale lugens is a koinobiont primary endoparasitoid and the female oviposits in the second instar larvae of its gall midge host, -Mikiola fagi +Mikiola fagi , which induces galls on the upper surface of leaves of beech ( -Fagus sylvatica +Fagus sylvatica ). This parasitoid is solitary or gregarious with up to ten individuals per host. If more than one larva is present per host either the same female lays more than one egg per host, or more than one female oviposits in the same host larva. However, because of the high mortality among gregarious larvae eventually only one parasitoid per host larva will complete its development. The parasitoid speeds up the development of its host, indicated by a faster growth rate in parasitized gall midge larvae, as compared to non-parasitized larvae. There does not seem to be additional, abnormal, instars in the host, just an acceleration of growth. The gall midge larva is killed before pupation by the feeding of the last instar parasitoid larva. -Omphale lugens +Omphale lugens has four larval instars and pupates inside the empty larval skin of its host, still remaining inside the gall. - -Table 1. + +Table 1. - - - -
-Omphale aethiops + + + + - - - + - - - + - - - + - - - + - - - + - - - + - - - + - - - + - - - + - - - + - - - + - - - + - - - + - - - + - - - + -
+Omphale aethiops -Dasineura epilobii -Diptera -Cecidomyiidae -Chamaenerion angustifolium -Gijswijt 1976 -Silene dioica -Askew 2003 -Dasineura traili -Ranunculus + +Dasineura epilobii +Diptera +Cecidomyiidae +Chamaenerion angustifolium +Gijswijt 1976 +Silene dioica +Askew 2003 +Dasineura traili +Ranunculus
-Omphale brevis +
+Omphale brevis -Cystiphora taraxaci -Diptera -Cecidomyiidae -Gijswijt 1976 -Cystiphora sonchi -Sonchus palustris -Vidal 1993 -Cystiphora sanguinea -Hieracium sabaudum -Askew 2003 + +Cystiphora taraxaci +Diptera +Cecidomyiidae +Gijswijt 1976 +Cystiphora sonchi +Sonchus palustris +Vidal 1993 +Cystiphora sanguinea +Hieracium sabaudum +Askew 2003
-Omphale chryseis +
+Omphale chryseis -Contarinia medicaginis -Diptera -Cecidomyiidae - -Boucek + +Contarinia medicaginis +Diptera +Cecidomyiidae + +Boucek and Askew 1968 - -Kralovic + +Kralovic 1964
-Omphale clymene +
+Omphale clymene -Dasineura pyri -Diptera -Cecidomyiidae + +Dasineura pyri +Diptera +Cecidomyiidae
-Omphale clypealis +
+Omphale clypealis -Dasineura brassicae -Diptera -Cecidomyiidae -Gijswijt 1976 + +Dasineura brassicae +Diptera +Cecidomyiidae +Gijswijt 1976
-Omphale erginnus +
+Omphale erginnus -Cecidomyiidae -Diptera -Hansson 1996b + +Cecidomyiidae +Diptera +Hansson 1996b
-Omphale grahami +
+Omphale grahami -Dasineura trifolii -Dasineura glechomae -Diptera -Cecidomyiidae -Gijswijt 1976 + +Dasineura trifolii +Dasineura glechomae +Diptera +Cecidomyiidae +Gijswijt 1976
-Omphale isander +
+Omphale isander -Mycodiplosis -Diptera -Cecidomyiidae -Populus -Kamijo 1986 + +Mycodiplosis +Diptera +Cecidomyiidae +Populus +Kamijo 1986
-Omphale lugens +
+Omphale lugens -Mikiola fagi -Dziurzynski 1961 -Contarinia tiliarum -Dasyneura alni -Gijswijt 1976 -Placochela nigripes -Diptera -Cecidomyiidae + +Mikiola fagi +Dziurzynski 1961 +Contarinia tiliarum +Dasyneura alni +Gijswijt 1976 +Placochela nigripes +Diptera +Cecidomyiidae
-Omphale lugubris +
+Omphale lugubris -Picea -Gijswijt 1976 + +Picea +Gijswijt 1976
-Omphale obscura +
+Omphale obscura -Dasineura viciae -Diptera -Cecidomyiidae -De Stefani 1905 -Galium mollugo - -Boucek + +Dasineura viciae +Diptera +Cecidomyiidae +De Stefani 1905 +Galium mollugo + +Boucek and Askew 1968
-Omphale phruron +
+Omphale phruron -Dasineura pyri -Diptera -Cecidomyiidae -Gijswijt 1976 + +Dasineura pyri +Diptera +Cecidomyiidae +Gijswijt 1976
-Omphale rubigus +
+Omphale rubigus -Trigonodiplosis -Diptera -Cecidomyiidae -Vicia cracca - -Boucek + +Trigonodiplosis +Diptera +Cecidomyiidae +Vicia cracca + +Boucek and Askew 1968
-Omphale salicis +
+Omphale salicis -Contarinia lentis -Diptera -Cecidomyiidae - -Szelenyi + +Contarinia lentis +Diptera +Cecidomyiidae + +Szelenyi 1944 -Contarinia loti -Gijswijt 1976 -Contarinia vincetoxici +Contarinia loti +Gijswijt 1976 +Contarinia vincetoxici
-Omphale incognita +
+Omphale incognita -Geocrypta galii -Diptera -Cecidomyiidae -Galium + +Geocrypta galii +Diptera +Cecidomyiidae +Galium
-Omphale euphorbiae +
+Omphale euphorbiae -Bayeria capitigena -Diptera -Cecidomyiidae -Euphorbia esula + +Bayeria capitigena +Diptera +Cecidomyiidae +Euphorbia esula
- + The only other -Omphale +Omphale species for which more detailed biological information is available is -Omphale clypealis +Omphale clypealis . This species is an important biological control agent against the brassica pod midge ( -Dasineura brassicae +Dasineura brassicae ) on rape ( -Brassica napus +Brassica napus ), and has thus been the subject of biological investigation ( -Williams 2003 +Williams 2003 ). Similar to -Omphale lugens +Omphale lugens , -Omphale clypealis +Omphale clypealis is a koinobiont endoparasitoid. Females oviposit into mature gall midge larvae in their pod gall, and the parasitoid larva is inside the host when it leaves the gall and burrows into the ground for pupation. The pupa is subsequently killed by the parasitoid. The sex ratio for -Omphale clypealis +Omphale clypealis is strongly female biased, and -Murchie (1996) +Murchie (1996) reared 97-100% females from samples of the brassica pod midge in the U.K. Material from this species that has been available for this investigation, 147 females and three males, suggests the same female bias. This skewed sex ratio indicates that -Omphale clypealis +Omphale clypealis is thelytokous, as are possibly some other -Omphale +Omphale species, such as -Omphale rubigus +Omphale rubigus where males have never been found, and -Omphale theana +Omphale theana in which very few males are recorded. - + If data from these two species are transferrable, -Omphale +Omphale species are koinobiont endoparasitoids, possibly solitary - at least with only one surviving parasitoid per host. Little is known about the host specificity of -Omphale +Omphale species. However, known host records suggest host specialization because there are no overlap between the species. The host record for -Omphale phruron +Omphale phruron , -Dasineura pyri +Dasineura pyri - same host as in -Omphale clymene +Omphale clymene , is possibly based on a misidentification of the parasitoid. The specimens of -Omphale phruron +Omphale phruron from -Dasineura pyri +Dasineura pyri have not been available for this investigation and as the species in this group are difficult to identify misidentification cannot be ruled out. - -Distribution. - + +Distribution. + Even though several parts of the World are very poorly investigated or not investigated at all, existing records show that -Omphale +Omphale is a cosmopolitan genus, -known +known from all zoogeographical regions ( -Noyes 2012 +Noyes 2012 ). As can be seen from previous investigations ( -Hansson 1996b +Hansson 1996b , -1997 +1997 , -2004 +2004 ), and this investigation (Figs 502-538), the species have a very large distribution. Many European species are distributed throughout Europe, and the more limited distribution in some species is possibly due to lack of distributional data. Five European species, -Omphale acamas +Omphale acamas , -Omphale erginnus +Omphale erginnus , -Omphale salicis +Omphale salicis , -Omphale theana +Omphale theana , -Omphale versicolor +Omphale versicolor , are also found in North America ( -Hansson 1996b +Hansson 1996b ), and one of these ( -Omphale erginnus +Omphale erginnus ) is distributed south to Central America ( -Hansson 2004 +Hansson 2004 ).