Coleoptera: Curculionidae

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Zootaxa 2943: 45–57 (2011) www.mapress.com / zootaxa/ Copyright © 2011 · Magnolia Press

ISSN 1175-5326 (print edition)

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ZOOTAXA ISSN 1175-5334 (online edition)

Study of a part of the A. G. Olivier Lixini collection (Coleoptera: Curculionidae): lectotype designations, new synonymies and nomenclatural acts LEVENT GÜLTEKİN1 & HÉLÈNE PERRIN2 1

Atatürk University, Faculty of Agriculture, Department of Plant Protection, 25240 Erzurum, TURKEY. E-mail: [email protected] Muséum National d’Histoire Naturelle, Département Systématique et Évolution, UMR 7205, MNHN-CNRS, CP 50 (Entomologie), 57 rue Cuvier, F-75231 Paris cedex 05, FRANCE. E-mail: [email protected] 2

Abstract A search for Lixini (Curculionidae: Lixinae) species housed in the Muséum National d’Histoire Naturelle, Paris and the Swedish Natural History Museum, Stockholm allowed the study of a part of the G. -A. Olivier collection. Lectotypes are designated for Larinus centaurii (Olivier, 1807), Larinus cuniculus (Olivier, 1807), Larinus gravidus (Olivier, 1807), Larinus carthami (Olivier, 1807), Lixus cardui Olivier, 1807, Lixus iridis Olivier, 1807, Lixus recurvus Olivier, 1807 and Lixus myagri Olivier, 1807. The following new synonymies and homonymies become evident: Larinus centaurii (Olivier, 1807) [= L. ungulatus Gyllenhal, 1835 syn. nov., L. centaureae Becker, 1864 nom. nud., L. beckeri Petri, 1907 syn. nov.]; Larinus cuniculus (Olivier, 1807) [= L. marki Ter-Minassian, 1982 syn. nov.]; Larinus ursus (Fabricius, 1792) [= L. cuniculus Walker, 1871 hom. nov. nec L. cuniculus (Olivier, 1807) syn. nov.]; Larinus gravidus (Olivier, 1807) [= L. schoenherri Capiomont, 1874 syn. nov.]; Larinus carthami (Olivier, 1807) [= L. crassus Capiomont, 1874 syn. nov., L. griseotessellatus Capiomont, 1874 syn. nov.; L. breviusculus Desbrochers, 1892 syn. nov.]; Lixus recurvus Olivier, 1807 [= L. nordmanni Hochhuth, 1847 syn. nov.]; Lixus myagri Olivier, 1807 [= L. lycophoeus Boheman, 1835 syn. nov. ]. Key words: Lectotypes, new synonymies, Olivier, Lixini, Curculionidae

Introduction The tribe Lixini Schoenherr, 1823 is a rather speciose one, being comprised of around 700 species (Csiki, 1934) with 13 genera worldwide (Alonso-Zarazaga & Lyal, 1999). The highest number of Lixini is to be found in the Palaearctic with over 300 species, closely followed by the Afrotropical region with 200 species. The largest genera are Lixus Fabricius, 1801 (about 200 species) and Larinus Dejean, 1821 (about 150 species). Species within Lixini are strictly phytophagous and economically important; however, they are still taxonomically poorly known, although some recent papers by Gültekin (2006, 2010), Gültekin & Perrin (2006) have started to clarify the status of several species previously incorrectly identified. This paper presents an additional contribution, with the study of the types of some of the species described by Olivier in the first decade of the nineteenth century. Guillaume-Antoine Olivier (1756–1814) was a French naturalist and entomologist. He collected extensively in Europe and made a great journey of six years in the Middle East, then included in the Ottoman Empire, Persia and Egypt. He assembled an extensive collection and returned to France in 1798. He wrote a series of volumes in a span of over 35 years from 1789 to 1825, publishing 10 volumes and 389 colour plates. Volumes dealing with Curculionidae are the fourth (Olivier, 1789) and the fifth (Olivier, 1790) of the “Encyclopédie méthodique”, and the fifth (Olivier, 1807) and the eighth for plates (Olivier, 1808) of the “Entomologie”. His collection is preserved mainly in the Muséum National d'Histoire Naturelle in Paris, a small part in the Royal Museum of Scotland in Edinburgh, the Hunterian Zoology Museum of the University of Glasgow in Glasgow, and a part in the Swedish Natural History Museum in Stockholm (Horn et al. 1990). Olivier (1807) described 70 species under the genus Lixus, 35 of these new. Currently these species are spread among Lixus, Larinus, Rhinocyllus Germar, 1817 and Liocleonus Motschulsky, 1860. The present note deals with type material of eight Lixini species described by Olivier, with some new synonymies and nomenclatural changes. Accepted by R. Anderson: 31 May 2011; published: 7 Jul. 2011

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Material The studied type material is deposited in the Muséum National d'Histoire Naturelle, Paris (MNHN) and the Swedish Museum of Natural History, Stockholm (SMNH). For the species treated in this paper, the examined species in Paris came through various sources (Cambefort, 2006: 250). A first part entered into the Muséum National d’Histoire Naturelle, Paris in 1837, and is preserved in the general collection. A second part, kept by Olivier’s family, arrived recently (1995) as a gift of a descendant of Ernest Olivier (1844–1914), entomologist and grandson of G. A. Olivier. To assess the status of the studied material, the original descriptions and plates by Olivier (1807) were used. Lectotype designations are made for taxonomic purposes following the rules of the International Commission on Zoological Nomenclature (1999).

Taxonomy Larinus centaurii (Olivier, 1807) Figs. 1–2, 17. Lixus centaurii Olivier, 1807: 278 ; pl. 21, fig. 276.

Original description. "L. longirostris, ovatus, niger, cinereo pubescens, rostro tenui cylindrico. Corpus ovatum, convexum, nigrum, cinereo pubescens. Rostrum tenue, vix incurvum, cylindricum, longitudine thoracis nec sulcatum neque carinatum. Thorax punctatus dorso linea brevi lateribusque obsolete cinereis. Elytra striatopunctata. Femora inermia". Type locality. “Amadan en Perse”. Types. In the general collection of MNHN, under the name of “L. centaurii Olivier, Perse, M. Olivier” written on yellow paper, a single male, pinned through the right elytron and in good condition, bears the labels “L. centaurii, Perse, Olivier” handwritten on a round paper, “MUSEUM PARIS, Perse, OLIVIER 1834” partly printed. The specimen matches both description and type locality, however Olivier (1807) did not mention on how many specimens the description was based. Accordingly we have designated it as the lectotype with: “LECTOTYPUS ♂, Lixus centaurii Olivier, Gültekin & Perrin des. 2010” and added a white label with “Larinus centaurii (Olivier), Gültekin det. 2010”. Remarks. Larinus centaurii is one of most confused, misspelled and misidentified species with Larinus grisescens Gyllenhal, 1835. The first misspelling “centaurei” was by Chevrolat (1866: 106). Gemminger & Harold (1871: 2408) used another misspelling “centaureae” and Capiomont (1874: 322), Petri (1907: 126), and Winkler (1932: 1506) repeated the same misspelling. A third misspelling, “centaurae”, was made by Angelov (1978: 159). A fourth subsequent misspelling “centaurea” was used by Fremuth (1982: 245) and Lodos et al. (1978: 75). All above subsequent alternative spellings are here rejected. The valid name is Larinus centaurii (Olivier, 1807) as truly spelled by Schoenherr (1835: 139) and Gültekin (2006). Capiomont (1874) stated that the types of L. centaurii (Olivier) were in the Chevrolat collection (SMNH), and gave short morphological remarks above these specimens. The senior author of this note visited the SMNH in 2010, and examined the specimens under the name “Larinus centaurei Oliv., 83, 4, p. 378, f 21, f. 296, seti. 199, Persia, Bagdad” in the Chevrolat collection. There were six specimens representing two different species: a female of L. grisescens bearing the label “Syria”, a teneral male of the true L. centaurii labelled “Chev. centaurea (Ol.)", a male of L. grisescens with the labels “adin”, “Larinus tenuicorpus Chev., longirostris Sch. III. P. 124, var. major”, a male and a female of L. grisescens labelled “Syria”, a female of L. grisescens with the label “Chev. centaurei, bounivieri Stier.”. As seen above, none of these specimens is typical. Thus Capiomont (1874) clearly misidentified the true L. centaurii with Larinus grisescens Gyllenhal, 1835, and his redescription mislead all subsequent authors. In the Schoenherr collection at SMNH, under the name “Ungulatus, Schh.”, there is a female of Larinus centaurii, bearing the handwritten label “Elisabethgrad, D. Steven”, (Elisabethgrad is now Kirovohrad in Ukraine). This sample is a syntype because Gyllenhal (1835) included it in his description mentioning it as the probable other sex of the other(s) type specimen(s), which are said to be in the Faldermann collection (Gyllenhal, 1835: 93). Having the Faldermann collection split among several institutes and museums such as the Zoological Institute of Kiev,

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FIGURES 1–4. Lectotypes. 1–2, Larinus centaurii (Olivier, 1807); 3–4, Larinus cuniculus (Olivier, 1807).

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the Zoological Institute of St. Petersburg and the Zoological Museum of Moscow (Horn et al. 1990), the other types could not be traced, and based on the above female syntype the following new synonymy become evident: Larinus centaurii (Olivier, 1807) [= L. ungulatus Gyllenhal, 1835, syn. nov.]. Capiomont (1874) also gave a short redescription of Larinus ungulatus Gyllenhal, 1835 and placed the name of L. centaureae Becker, 1864 under synonymy of L. ungulatus. This is correct because Becker (1864: 483) used the name of Larinus centaureae without giving a description of it, so this is evidently a nomen nudum. Petri (1907), considering the Capiomont L. ungulatus as different from the species described under this name by Gyllenhal, and from the true L. centaurii, replaced L. ungulatus sensu Capiomont with L. beckeri, but this replacement name is unnecessary because Capiomont (1874) quoted Schoenherr (1835) writing “Larinus ungulatus Gyll. in Sch., 133” in his redescription. Thus, another new synonymy must be established: Larinus centaurii (Olivier, 1807) [= L. centaureae Becker, 1864 nom. nud. = L. beckeri Petri, 1907, syn. nov.].

Larinus cuniculus (Olivier, 1807) Figs. 3–4, 18. Lixus cuniculus Olivier, 1807: 282 ; pl. 30, fig. 458.

Original description. "L. albo pubescens, elytris striatis fusco alboque variis, thorace scabro. Lixo Jaceae paulo minor. Corpus nigrum albo pubescens. Rostrum cylindricum vix longitudine thoracis. Thorax scaber. Elytra striata, fusco alboque varia versus apicem gibbere parvo notata. Pedes corpore concolores. Femora mutica". Type locality. “Nord de l’Arabie, près de l’Euphrate”. Types. In the general collection of MNHN, under the name of “L. cuniculus Olivier, arabie, M. Olivier” written on yellow paper, there is a single male pinned through the left elytron by a headless and curved pin, elytra divaricated and hind left leg missing. It bears the labels “L. cuniculus, arabie” handwritten on a round paper, “MUSÉUM PARIS, Arabie” partly printed. The specimen agrees with both description and type locality and is here designated as the lectotype with: “LECTOTYPUS ♂, Lixus cuniculus Olivier, Gültekin & Perrin des. 2010” and identified as “Larinus cuniculus (Olivier), Gültekin det. 2010”. Remarks. The name of this species was first misspelled as “cuniculinus” by Gemminger & Harold (1871: 2408), a spelling used also by Marseul (1888: 418), Winkler (1932: 1504), and Csiki (1934: 75), the last authors both considering L. cuniculus as a questionable synonym of L. onopordi (Fabricius, 1787). Study of the types clearly revealed that L. cuniculus is a good species different from L. onopordi, and that in addition L. marki TerMinassian, 1982 is conspecific with it, according to the examination of types of both species by the senior author. Consequently Larinus cuniculus (Olivier, 1807), synonym revised (= L. marki Ter-Minassian, 1982, syn. nov.). Moreover, Walker (1871) described a species from Arabia with the name of Larinus cuniculus Walker, 1871, hom. nov. of Larinus cuniculus (Olivier, 1807) and mentioned that this Arabian species was different from Olivier’s one. However his description is that of a Larinus ursus (Fabricus, 1792), so another synonymy must be established: Larinus ursus (Fabricius, 1792) [= L. cuniculus Walker, 1871, nec L. cuniculus (Olivier, 1807), syn. nov.].

Larinus gravidus (Olivier, 1807) Figs. 5–6, 19. Lixus gravidus Olivier, 1807: 275 ; pl. 30, fig. 450.

Original description. "L. longirostris fuscus, thoracis lateribus albis, elytris striatis albo pulverulentis. Similis L. Scolymi. Antennae nigrae apice cinereae. Rostrum incurvum cylindricum nigrum basi cinereo pubescens, a basi usque ad insertionem antennarum linea vix elevata; inter antennas foveola oblonga. Thorax punctatus fuscus lateribus albo pubescentibus. Elytra striato-punctata fusca pube alba irrorata. Corpus subtus pedesque fusca cinereo pubescentia. Femora mutica." Type locality. “Amérique septentrionale”. Types. In a box in the MNHN recently given by Olivier’s family to the Museum, there is a female bearing the labels “COLLECTION OLIVIER TYPE” printed on a green round paper, “Rhinobatus, Gravidus. Ol., C. Olivier,

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FIGURES 5–8. Lectotypes. 5–6, Larinus gravidus (Olivier, 1807); 7–8, Larinus carthami (Olivier, 1807).

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Amer. bor.” handwritten on green paper, and “MUSEUM PARIS, Famille Olivier, leg. 1995”, “MNHN EC 1940” printed. The specimen, agreeing with both description and type locality is here designated as the lectotype with: “LECTOTYPUS ♀, Lixus gravidus Olivier, Gültekin & Perrin des. 2010”, identified as “Larinus gravidus (Olivier), Gültekin det. 2010”. The lectotype is pinned through the right elytron. Remarks. Description and label data of the type specimen are in accordance with the original description (Olivier 1807); however, there is no further record about the finding of this species in America (O’Brien & Wibmer, 1982; Wibmer & O'Brien, 1986). Members of the genus Larinus are not distributed in the Nearctic, Neotropical and Australian regions (Csiki, 1934), except for species introduced into the United States of America such as L. minutus Gyllenhal, 1835, L. obtusus Gyllenhal, 1835 and L. curtus Hochhuth, 1851 (Lang et al. 1997; Sobhian & Fornasari, 1994), and into the Australian region, like L. latus (Herbst, 1783) (Woodburn & Briese, 1996), for biological control of weeds. Recently an accidental introduction into the United States of L. carlinae (Olivier, 1807) (McClay, 1990) [= Larinus planus sensu Germar, 1824, (not Fabricius, 1792)] (see Gültekin, 2006) was also reported. Consequently, we considered the type locality “Amérique septentrionale” as erroneus, also because L. gravidus (Olivier, 1807) turns out to be conspecific with L. schoenherri Capiomont, 1874 which is distributed in Spain, Portugal and Algeria (Csiki, 1934; Alonso-Zarazaga, 2002). Consequently: Larinus gravidus (Olivier, 1807) [= L. schoenherri Capiomont, 1874, syn. nov.].

Larinus carthami (Olivier, 1807) Figs. 7–8, 20. Lixus carthami Olivier, 1807: 278; pl. 21, fig. 277.

Original description. "L. longirostris, ovatus niger, cinereo pubescens, rostro nigro carinato. Corpus ovatum, convexum, nigrum, pube cinereo-rufescente plus minusve tectum. Rostrum nigrum, longitudine thoracis, incurvum, a basi ad apicem vix carinatum. Thorax punctulatus basi congeneribus paulo latior. Elytra striatopunctata. Femora inermia." Type locality. “Egypte”. Types. In the Chevrolat collection in SMNH under the label “Larinus carthami Ol. type, Ent. V. 83, p. 278, p21, f277; Capiom. Leprieur 204/47, Aegyptus, ex. mus. Olivier” handwritten on blue paper, there is a female bearing the handwritten label “Chevrolat, Carthami, type”. This sample, matching both description and type locality, is here designated as the lectotype with: “LECTOTYPUS ♀, Lixus carthami Olivier, Gültekin des. 2010”, and identified as “Larinus carthami (Olivier), Gültekin det. 2010”. The lectotype was pinned through the right elytron, but there is also a pin hole on the left one, and its fore right claw segment, right middle tarsus and left middle leg are lacking. Remarks. The study of the type of L. carthami and that of several other Larinus types recently made by the first author made evident the following new synonymies: Larinus carthami (Olivier, 1807) [= L. crassus Capiomont, 1874, syn. nov., L. griseotessellatus Capiomont, 1874, syn. nov., L. breviusculus Desbrochers, 1892, syn. nov.].

Lixus iridis Olivier, 1807 Figs. 9–10, 21. Lixus iridis Olivier, 1807: 239 ; pl. 9, fig. 106.

Original description. "L. longirostris fuscus, cylindricus, elytris mucronatis lineis plurimis obsoletis cinereis. Magnitudo et statura L. anguini. Antennae fuscae articulo primo piceo, clava cinerascente. Rostrum elongatum, tenue, cylindricum vix arcuatum nigrum. Thorax rugosus antice linea brevi elevata, fuscus vitta laterali obsoleta cinerea. Elytra mucronata punctato-striata fusea lineis plurimis obsoletis cinereis. Corpus fuscum". Type locality. “Midi de la France”. Types. In a box in the MNHN recently given by Olivier’s family to the Museum, are a female and a male. The female, pinned through the right elytron, bears the labels “COLLECTION OLIVIER TYPE” printed on green

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FIGURES 9–12. Lectotypes. 9–10, Lixus iridis Olivier, 1807; 11–12, Lixus recurvus Olivier, 1807.

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round paper, “Lixus, Iridis. Ol., Pulverulentus. Dahl., C. Olivier, Gal. mer” handwritten and “MUSEUM PARIS, Famille Olivier, leg. 1995”, “MNHN EC 1941” printed. The specimen, agreeing with both description and type locality is here designated as the lectotype with: “LECTOTYPUS ♀, Lixus iridis Olivier, Gültekin & Perrin des. 2010”. The male, also pinned through the right elytron, bearing the label “COLLECTION OLIVIER TYPE” printed on green round paper and “MUSEUM PARIS, Famille Olivier, leg. 1995”, and matching the description, is here designated as the paralectotype with: “PARALECTOTYPUS ♂, Lixus iridis Olivier, Gültekin & Perrin des. 2010”. Remarks. Gültekin (2006) considered Lixus iridis Olivier, 1807 as a nomen protectum, and Lixus pulverulentus Fabricius, 1801 a nomen oblitum, leaving the latter into synonymy of the former. This is a well known Palaearctic species related to L. recurvus Olivier, 1807 (see below).

Lixus recurvus Olivier, 1807 Figs. 11–12, 22. Lixus recurvus Olivier, 1807: 237 ; pl. 21, fig. 287.

Original description. "L. longirostris, niger, elytris striatis acutis flavo pulverulentis. Lixo paraplectico major et crassior. Antennae tenues, nigrae. Rostrum cylindricum, thorace longius, nigrum. Thorax rugosus, niger, immaculatus. Elytra vix punctato-striata, flavo pulverulenta, apice acuto reflexo. Pedes flavo pubescentes femoribus muticis". Type locality. “Georgie”. Types. In a box in the MNHN recently given by Olivier’s family to the Museum, a female bears the labels “COLLECTION OLIVIER TYPE” printed on green round paper, and “Lixus, Recurvus. Ol., C. Olivier, Georg. Asia.” handwritten, “MUSEUM PARIS, Famille Olivier, leg. 1995”and “MNHN EC 1944” printed. The specimen, agreeing with both description and type locality is here designated as the lectotype with: “LECTOTYPUS ♀, Lixus recurvus Olivier, Gültekin & Perrin des. 2010”. This female is pinned through the right elytron causing the elytra to divaricate, and its right funicle, left funicle beyond the first segment, left fore claw segment, right middle tarsus beyond segment 2 are missing. Remarks. This name is listed as a questionable synonym of Lixus furcatus Olivier, 1807 in Csiki (1934). Lixus recurvus Olivier, 1807 is removed from synonymy and placed in the Lixus iridis group by Gültekin & Korotyaev (in press). The above lectotype designation confirmed that Lixus recurvus Olivier, 1807 is the same species to Lixus nordmanni Hochhuth, 1847, and since the younger name does not fully meet the requirements of article 23.9.1.2 of the Code (ICZN 1999), the following synonymy is here established: Lixus recurvus Olivier, 1807 [= L. nordmanni Hochhuth, 1847, syn. nov.]. Lixus recurvus is closely related to L. iridis, but its rostrum is longer, and the apex of its aedeagus is round instead of triangular.

Lixus myagri Olivier, 1807 Figs. 13–14, 23. Lixus myagri Olivier, 1807: 249 ; pl. 30, fig. 453.

Original description. “L. rostro elongato carinato, cinereus, thorace vittis tribus fuscis intermedia obsoleta. Lixo Ascanii paulo major et crassior. Antenna cinerascentes. Rostrum inflexum, fere longitudine thoracis linea dorsali elevata. Corpus cinereum. Thorax vix rugosus postice foveola impressa dorso fusco lineis duabus lateribusque albicantibus. Elytra striata cinerea fusco punctata marginibus albicantibus. Corpus subtus pedesque cinerea femoribus muticis". Type locality. “Paris”. Types. In a box in the MNHN recently given by Olivier’s family to the Museum are two females. The first, pinned through the right elytron, bears the labels “COLLECTION OLIVIER TYPE” printed on green round paper, “Cleonis, myagri. Ol., C. Olivier, [P.]” handwritten and “MUSEUM PARIS, Famille Olivier, leg. 1995”, “MNHN EC 1942” printed. The letter [P.] on the label is surely an abbreviation of Paris which is the type locality, so that the

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FIGURES 13–16. Lectotypes. 13–14, Lixus myagri Olivier, 1807; 15–16, Lixus cardui Olivier, 1807.

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FIGURES 17–20. Labels of lectotypes. 17. Larinus centaurii (Olivier, 1807); 18. Larinus cuniculus (Olivier, 1807); 19. Larinus gravidus (Olivier, 1807); 20. Larinus carthami (Olivier, 1807).

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FIGURES 21–24. Labels of lectotypes. 21. Lixus iridis Olivier, 1807; 22. Lixus recurvus Olivier, 1807; 23. Lixus myagri Olivier, 1807; 24. Lixus cardui Olivier, 1807.

specimen, agreeing with both description and type locality is here designated as the lectotype with: “LECTOTYPUS ♀, Lixus myagri Olivier, Gültekin & Perrin des. 2010”. The second female pinned through the right elytron bears the labels “COLLECTION OLIVIER TYPE” printed on green round paper and “MUSEUM PARIS, Famille Olivier, leg. 1995”, and is here designated as the paralectotype with: “PARALECTOTYPUS ♀, Lixus myagri Olivier, Gültekin & Perrin des. 2010”. Remarks. This species was always recognized by subsequent authors. Lixus lycophoeus Boheman, 1835, misspelled as “lycophorus” by Gemminger & Harold (1871: 2416) firstly, Desbrochers (1904: 102) and Csiki (1934: 120 and 144) repeated this misspelling, and Csiki (1934) placed this name as a synonym of Lixus meles Boheman, 1835. However, examination of type material of L. lycophoeus revealed that it is the same species to L. myagri, and the following synonymy must be established: Lixus myagri Olivier, 1807 [= L. lycophoeus Boheman, 1835, syn. nov.].

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Lixus cardui Olivier, 1807 Figs. 15–16, 24. Lixus cardui Olivier, 1807: 250 ; pl. 30, fig. 454.

Original description. "L. longirostris, cinereo pubescens, thorace lineis quatuor obsoletis albis, rostro laevi incurvo. Similis L. Myagri, at differt rostro tenuiori, longiori, haud carinato. Corpus fuscum cinereo pubescens. Thorax linea laterali lineisque duabus dorsalibus obsoletis albicantibus. Elytra vix striata apice rotundato". Type locality. “Aux environs de Constantinople”. Types. In a box in the MNHN recently given by Olivier’s family to the Museum, a female pinned through the right elytron and with another pin hole on the left one bears the labels “COLLECTION OLIVIER TYPE” printed on green round paper, “Lixus, Cardui. Ol., Tigrinus? 11109, Constantinople., C. Olivier”, handwritten, and “MUSEUM PARIS, Famille Olivier, leg. 1995”, “MNHN EC 1943” printed. The specimen, agreeing with both description and type locality is here designated as the lectotype with: “LECTOTYPUS ♀, Lixus cardui Olivier, Gültekin & Perrin des. 2010”. Remarks. This species is an important biological control agent against Onopordum weeds, already introduced into Australian region (Briese, 1996), always correctly recognized by subsequent authors.

Acknowledgements We thank Dr. Kjell Arne Johanson and Bert Viklund (Swedish Museum of Natural History, Stockholm), Dr. Lutz Behne (Deutsches Entomologisches Institute, Müncheberg), Dr. Massimo Cristofaro (Biotechnology and Biological Control Agency, Rome) and Dr. Lincoln Smith (USDA-ARS, Western Regional Research Center, Albany) for their contributions and support. We would like to express our special thanks to Dr. Enzo Colonnelli (Rome, Italy), Dr. Boris A. Korotyaev (Zoological Institute, Russian Academy of Sciences, St. Petersburg) and Dr. Robert Anderson (Canadian Museum of Nature, Ottawa) for their valuable review of the manuscript. The study was supported by grants from the European Commission’s (FP 7) Integrated Infrastructure Initiative programme SYNTHESYS (SE-TAF-477) for visit SMNH Stockholm and a BBCA grant for visit MNHN Paris.

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