NORTH-WESTERN JOURNAL OF ZOOLOGY 8 (2): 268-277 ©NwjZ, Oradea, Romania, 2012 Article No.: 121122 http://biozoojournals.3x.ro/nwjz/index.html

Herb gallwasp fauna of Iran (: Cynipidae, Aylacini)

George MELIKA1,* and Younes KARIMPOUR2

1. Pest Diagnostic Department, Plant Protection and Soil Conservation Directorate of County Vas, 9762 Tanakajd, Ambrozy setany 2, Hungary. E-mail: [email protected] 2. Department of Plant Protection, College of Agriculture, Urmia University, Urmia, Iran. E-mail: [email protected] * Corresponding author, G. Melika, Ee-mail: [email protected]

Received: 12. October 2011 / Accepted: 14. April 2012 / Available online: 31. May 2012 / Printed: xxxxxxxxx

Abstract. During a 5-year survey (2006-2010) of the herb gallwasp fauna of Iran a total of 21 gallwasp species (Hymenoptera: Cynipidae, Aylacini) in 6 genera were found, reared from different species of Asteraceae, Lamiaceae and Papaveraceae plants. Ten species, Aulacidea hieracii, A. scorzonerae, A. tragopogonis, Aylax minor, A. papaveris, Barbotinia oraniensis, Isocolus centaureae, I. similis, Phanacis heteropappi, P. varians, are new records for the fauna of Iran. With the exception of Aulacidea irani which was swept on flower heads of Echinops sp., all specimens were reared from galls collected on host plants. Keys to the species identification and detail data on species biology, distribution are given.

Keywords: Aylacini, Cynipidae, distribution, herb gallwasps, Iran.

Introduction (Burks 1979, Melika 2006). This is the first study summarizing all the data The gallwasps (Cynipidae), which comprises on herb gallwasps of Iran. around 1300 species world-wide, are divided into two main trophic groups: the gall inducers and the gall-associated , which together make up Materials and methods six tribes, which from representatives of 4 tribes Different plants, with which the herb gallwasps are are known from Iran: , Aylacini, Dip- known to associate, were collected in different seasons lolepidini and Synergini. Recent studies on from meadows and rangelands in different parts of Iran, cynipid fauna of Iran were focused mainly on the mainly in Azerbaijan-e Gharbi (=West-Azerbaijan) (vicini- oak gallwasps (Cynipini) and their cynipid inqui- ties of Urmia) and Lorestan provinces. Collected plants lines (Synergini) (Melika & Stone 2001, Melika et were kept in glass boxes in the laboratory, they were al. 2004, Azizkhani et al. 2006, Tavakoli et al. 2008, checked weekly and the emerged wasps were preserved in 75% alcohol for further identification. Pénzes et al. 2009, Sadeghi et al. 2006, 2010). Very little is known about the herb gallwasp (Aylacini) fauna of Iran (Melika & Gharaei 2006, Tavakoli & Results Melika 2006, Melika & Karimpour 2008, Karim- pour et al. 2008, Karimpour 2011). During a 5-year survey (2006-2010) a total of 321 The tribe Aylacini comprises a group of ple- herb gallwasp specimens were reared which be- siomorphic genera of cynipid gall-inducers associ- long to 21 species from 6 genera of Aylacini ated with herbaceous plants. Most of the represen- (Cynipidae). With the exception of Aulacidea irani, tatives are associated with species of Asteraceae, which was collected from Echinops sp., all speci- some genera and species with Apiaceae, La- mens were reared from galls collected from host miaceae, Papaveraceae, Rosaceae and Valeriana- plants. Below, keys to genera and species of Ira- ceae (Melika 2006). Galls of Aylacini are not as nian Aylacini are given with data on each species. complex as in Cynipini or Diplolepidini; they are formed mainly in stems, flower heads, fruits and Key to genera of Aylacini in Iran seeds. All Aylacini species are monovoltine, only (all tarsal claws are simple, without basal lobes) sexual generations are known, without heterogeny 1. Mesopleuron reticulate or ruguloso-reticulate; or alternation of generations (Csóka et al. 2004). pronotum with indistinct submedian pronotal The Aylacini has a holarctic distribution, with pits or with transverse narrow impression; head near 130 species in 18 genera, from which ap- never broader than high in anterior view proximately 100 species are known from the Phanacis Palaearctic region and 30 species from the Nearctic Aylacini of Iran 269

-- Mesopleuron with transverse striae; pronotum Europe, three species are known from Tajikistan always with two distinct submedian pronotal and Turkmenistan (Diakontschuk 1984, 1988), one pits, separated by carina; head broader than high species was recently described from the Far East of in anterior view 2 Russia (Melika 2004) and 7 species are listed for 2. R1 and Rs not quite reaching wing margin; ra- America north of Mexico (Burks 1979). A total of dial cell of forewing entirely opened along mar- six species are known from Iran, and of these six gin 3 species two are found nowhere else. -- Rs almost reaching anterior margin of wing, R1 Key to Aulacidea species of Iran touching wing margin or continuing along wing (in all species known from Iran, the forewing mar- margin; radial cell closed, partially closed or ob- gin with a moderately long cilia; the head always soletely closed 4 slightly broader or at least as broad as high in an- 3. Mesopleuron with transverse striae, interspaces terior view; the female antenna with 11-12 flagel- reticulate; head trapezoid in anterior view; galls lomeres) in fruit capsules of Papaver Barbotinia

-- Mesopleuron with transverse striae, interspaces 1. F2 in female antenna nearly 2.0 times as long as smooth, shiny; head rounded in anterior view, F1; metascutellum medially 3.0-4.0 times as high never trapezoid; mesoscutum with more or less as height of ventral impressed area; 2nd me- distinct transverse striae; galls on Centaurea and tasomal tergite with posterior band of punctures, other Asteraceae Isocolus occupying its half length, without distinct patch 4. Radial cell opened along margin, R1 usually not of dense setae anterolaterally; male antenna with reaching wing margin or if so, than continuing 11 flagellomeres; galls in stems of Acroptilon re- on short distance along margin; pronotum dor- pens acroptilonica sally relatively short, submedian pronotal pits -- F2 in female antenna nearly equal, shorter or strongly transverse, narrowly separated; scutel- slightly longer than F1; metascutellum medially lar foveae small, coriaceous; galls on Papaver nearly as high as height of ventral impressed Aylax area; 2nd metasomal tergite without posterior -- Radial cell closed or almost closed, R1 always band of punctures, with distinct patch of dense more or less continuing along wing margin; setae anterolaterally; male antenna with 12 flag- pronotum dorsally always quite long; submedial ellomeres; galls in other plants 2 pronotal pits and scutellar foveae different; galls 2. 3rd metasomal tergite entirely punctuate 3 on Asteraceae and Salvia (Lamiaceae) 5 -- 3rd metasomal tergite with posterior band of 5. Mesopleuron uniformly and entirely trans- punctures which usually extending to at most nd versely striate; 2 metasomal tergite with dis- half length of tergite 5 tinct patch of white setae anterolaterally; female 3. Scutellar foveae transversely ovate, always antenna with 11 flagellomeres; notauli usually broader than high, smooth, shiny; punctures on complete, reaching pronotum; wing margin with 3rd metasomal tergite uniformly dense and dis- moderately long cilia; galls mainly on Asteraceae tinct; hypopygium large; galls in stems on Scor- Aulacidea zonera scorzonerae -- Mesopleuron partially transversely striate, par- -- Scutellar foveae elongated, longer than broad, nd tially alutaceous; 2 metasomal tergite without with wrinkles; punctures on 3rd metasomal ter- patch of white setae anterolaterally; female an- gite more sparse and weak anteriorly; hypopy- tenna with 10 flagellomeres; notauli incomplete, gium small; galls on Tragopogon 4 wing margin without cilia; galls on Salvia 4. POL 1.7 times as long as OOL and 3.7 times Hedickiana longer than diameter of lateral ocellus; clypeus quadrangular, nearly as high as broad; pedicel in Genus Aulacidea Ashmead, 1897 female antenna subglobose, only 1.25 times Type species: Aulax mulgediicola Ashmead, 1896 (= longer than broad; lateral propodeal carinae Aulax harringtoni Ashmead, 1897). The genus can converging outwards in posterior 1/3; F1 in be easily distinguished from other Aylacini by the male antenna longer than F2, F2 shorter than F3; closed radial cell of the forewing and the uni- galls always at the base of Tragopogon flowers, formly transverse striation of the mesopleuron. never in stems tavakolii Holarctic genus, with 29 world-wide known spe- -- POL 1.2 times as long as OOL and 4.5–4.7 times cies (Melika 2006): 17 species are known from as long as diameter of lateral ocellus; clypeus 270 G. Melika & Y. Karimpour

rectangular, at least 2.0 times as broad as high; Aulacidea hieracii (Linnaeus, 1758) pedicel in female antenna elongate, 1.5 times as Synonyms. Aulax graminis Cameron, 1875, Aulax long as broad; lateral propodeal carinae nearly artemisiae Thomson, 1877, Aulax foveigera Thom- straight, slightly converging inwards posteriorly; son, 1877, Aulax crassinervis Thomson, 1877, Aylax F1 in male antenna slightly shorter than F2, F2 sabaudi Hartig, 1840, Aulacidea cacaliae Belizin, equal F3; galls in Tragopogon stems, often merge 1959. into large conspicuous lignified, hard swellings Gall. In stems, multilocular, differently up to 40 mm in length, with rough surface shaped, large, hard and fuzzy galls; the stem is tragopogonis malformed. The gall develops on the stem or at 5. Frons and interocellar area uniformly finely the base of the leaf rosette, 40 mm long, 10-15 mm coriaceous, without punctures; lower face and in diameter, usually spindle-like, sometimes gena with distinct radiating striae; mesoscutum rounded. During the development, leaves are without piliferous punctures; galls in stems of coming out from it. The surface of the gall is cov- Hieracium spp. hieracii ered with hairs. The gall loosing its hairs after -- Frons and interocellar area uniformly finely overwintering and the bases of hairs are visible coriaceous, with distinct piliferous punctures, only on a glabrous epidermis. The parenchyma of lower face and gena with indistinct very minute the gall soft, spongy, with rounded larval cells. radiating striae; mesoscutum with piliferous The number of larval cells is decreasing towards punctures; gall unknown irani the center of the gall. The gall is green when young and growing, turns brown when mature. Aulacidea acroptilonica Tyurebaev, 1972 The shoot above the gall is malformed, abnormally Gall. Spindle-like, lignified formations on the short. Sometimes galls develop on the leaves stem, 25-50 mm long and 20 mm in diameter. The which are lying on the ground. gall size depends on the number of larval cham- Host plant. Hieracium murorum L., H. umbella- bers (from 1 to 30). A single larval chamber nearly tum L., H. verruculatum Link, H. virosum Pall., H. rounded, 3.0 mm in diameter, with smooth sur- vulgatum L. (Asteraceae). face. Galls develop on all parts of the stem, from Biology. Galls mature in autumn, larvae the ground-surface up to the flower (Kovalev & overwinter in the gall, adults emerge next spring, Diakontschuk 1986). in May (Zerova et al. 1988). Host plant. Acroptilon repens (L.) DC. (As- Distribution. Iran, Lorestan (Ghelaei or Ghara- teraceae) is the only known host plant; probably ei) and Azerbaijan-e Gharbi provinces (vicinity of can induce galls on a closely related species, A. Kelïsä Kandï village, 37º29′N, 45º01′E, 1600 m australe Iljin. (Kovalev & Diakontschuk 1986). a.s.l., 2.5 km SW Urmia) -- a new record for Iran. Biology. Majority of larvae overwintering in Europe (Melika 2006), Far East of Russia (Kovalev the gall, pupate in March-April; adults emerge by 1965), Transcaucasus (Zerova et al. 1988). the end of April (Middle Asia) – beginning of May (Kazakhstan and Ukraine) (Zerova, Diakontschuk Aulacidea irani Melika & Gharaei, 2006 & Ermolenko 1988); part of adults in Iran emerge Gall. Unknown. in early summer. Host plant. Unknown.. Distribution. Iran, Azerbaijan-e Gharbi prov- Biology. Adult wasps emerge from mid-May ince (vicinities of Urmia) (Melika & Karimpour (Melika & Gharaei 2006) till beginning of June. 2008), Lorestan (Zalian) (Karimpour et al. 2008). Distribution. Iran, Ilam (Melika & Gharaei Ukraine (southern part only), European part of 2006) and Azerbaijan-e Gharbi provinces (vicinity Russia, Kazakhstan, Kyrgyzstan, Turkmenistan, of Marmishü lake: 37º40′N, E44º48′E, 1570 m a.s.l., Tajikistan, Uzbekistan (Zerova, Diakontschuk & 42 km W Urmia) -- a new record for the province.

Ermolenko 1988, Kovalev & Diakontschuk 1986). Aulacidea scorzonerae (Giraud, 1859) Taxonomic comments. Middle Asian Gall. A stem swelling-like gall, usually located populations of A. acroptilonica differ from those in under the flower head, sometimes in the lower Iran by possessing much lighter flagellomeres, part of the stem; spindle –like or cylindrical stem legs and light brown metasoma. Specimens enlargement, 30–60 mm long, 10 mm in diameter; collected in southern part of Ukraine characterized the surface with longitudinal ribs, like on the stem, by dark brown F1–F2, F1 and proximal flagello- sometimes the ribs are waved and distance be- meres usually slightly longer (Melika 2006). tween them larger, depending on the diameter of Aylacini of Iran 271 the gall. In a longitudinal dissection, single 2 mm Quercus brantii. Wasps emerge from the end of large larval cells are visible, which are located one January untill March - April. near another. The number of larval cells is larger Distribution. Iran, Lorestan, Shorab (Karim- towards the periphery of the gall. The stem above pour et al. 2008), and Azerbaijan-e Gharbi prov- the gall is shortened and the flower stops devel- ince (Qasemlü valley, vicinity of Shïrü Kandï vil- opment (Melika 2006). lage ), 37º18′N, 45º07′E, 1420 m a.s.l., 29 km SE Host plant. Scorzonera austriaca Willd. (As- Urmia) -- a new record for the province. Turkey teraceae) (Ambrus 1974). Here we give a new host (Katilmiş & Kiyak 2011). plant record, S. calyculata Boiss., which from the species was recently reared in Iran. Genus Aylax Hartig, 1840 Biology. Adults overwinter in the mature galls Type species: Cynips rhoeadis Bouché, 1834 (= Ay- and emerge in April. lax papaveris (Perris, 1840). The genus Aylax com- Distribution. Iran, Azerbaijan-e Gharbi prov- prises four species; two are associated with Pa- ince (vicinity of Kelïsä Kandï village: 37º 29′ N, paver, one species, A. hypecoi, with Hypecoum (Pa- 45º01′E, 1600 m a.s.l., 2.5 km SW Urmia) - a new paveraceae). Three species are known from record for Iran. Austria, Russia, Hungary, Ukraine Europe: A. papaveris (Perris), A. minor Hartig and (Melika 2006). A. hypecoi Trotter; one species, Aylax quinque- costatus (Provancher) was described from Canada Aulacidea tragopogonis (Thomson, 1877) (Ontario); however, the status of this species is Synonyms. Aulax pigeoti Kieffer, 1898. questionable (Burks 1979). Galls on Papaver and Gall. Galls are in stems, small, egg-shaped Hypecoum. with smooth surface; the stem is not malformed externally, however, very often galls merge into a Key to Aylax species of Iran large conspicuous lignified, hard swelling up to 40 1. Body with variable extension of reddish brown mm in length, with rough surface (Melika 2006). colour; female antenna with 11-12 flagellomeres; Host plant. Tragopogon dubius Scop., T. maior Rs not reaching margin of wing; 2nd metasomal Jacq. (Kierych 1979), T. majus Jacq., T. orientale L., tergite with patch of dense white setae anter- T. porrifolius L., T. pratense L. (Asteraceae) (Kierych olaterally; hypopygium very small in lateral 1979, Melika 2006). view. Galls on fruits of Hypecoum hypecoi Biology. Adults overwinter in the gall and -- Body black; female antenna with 12 flagel- emerge next spring, in April. lomeres; Rs reaching margin of forewing; 2nd me- Distribution. Iran, Azerbaijan-e Gharbi prov- tasomal tergite with few short white setae anter- ince (vicinity of Täzehkand-e Qäterchï village, olaterally; hypopygium large in lateral view. 37º39′N, 44º58′E, 1335 m a.s.l., 13 km SW Urmia Galls in Papaver capsules 2 and vicinity of Rashkän village, 37º19′N, 45º18′E, 2. Notauli indistinct in anterior 1/3; mesoscutum 1270 m a.s.l., 31.5 km SE Urmia) – a new record for coriaceous; fruit capsule not enlarged and swol- Iran. Sweden, Spain, Romania, Poland, Great Brit- len, with multiple individual galls minor ain, Denmark, Austria, France, Russia, Ukraine -- Notauli distinct in anterior 1/3; mesoscutum co- (Melika 2006). riaceous-rugose; multilocular galls in hypertro-

Aulacidea tavakolii Melika, 2008 phised capsules papaveris

Gall. The gall is always formed at the base of Aylax hypecoi Trotter, 1913 flowers, bluntly conical; 10–15 mm in height and Synonyms. Aylax spirorhynchusii Diakontschuk, 9–13 mm in diameter. The gall is multilocular, con- 1990. taining 15–22 larval chambers (cells). The young Gall. Galls in fruits form a conspicuous swell- growing gall is green, later yellow and turns to ing. Fruits of Hypecoum species are typically grayish-black when mature. The walls of the gall nodose, divided by transverse septa, thus galled are semi-hard, even after the gall totally dries out. fruits affect one or more seeded sections which are Host plant. Tragopogon longirostris and T. mar- inflate considerably. The shape is oval or globular, ginatus Boiss. (Asteraceae) (Karimpour et al. 2008, slightly more elongated along the longitudinal Katilmiş & Kiyak 2011). axis. Each gall is 0.4-0.5 mm wide and 0.6-0.7 long. Biology. Galls are visible on the host by the In each fruit 1-3 galls develop. Each individual gall end of spring and beginning of summer. The usual has a single larval chamber separated from out habitat is dry grassland under oak trees, mainly 272 G. Melika & Y. Karimpour side by a thick wall. shorter and ripening occurs earlier, the poppy Host plant. Hypecoum geslini L., H. grandiflo- seeds turn brown and shrivel (Melika 2006). rum L., H. imberbe Sibth and Sm., H. pendulum L. Host plant. Papaver dubium L., P. rhoeas L., P. (Papaveraceae) (Nieves-Aldrey & Melika 2005, somniferum L. (Papaveraceae). For the first time re- Karimpour 2011, Katilmiş & Kiyak 2011). corded from P. argemone L. in Iran. Biology. Males are very rare. Adults emerge Biology. Adults emerge next year in May. In from galls from the end of the winter in North Af- Iran adults emerge in April. rica to early spring in Europe, at the time when the Distribution. Iran, Azerbaijan-e Gharbi prov- host plant starts flowering. Galls develop and ma- ince (Qasemlü valley, vicinity of Shïrü Kandï vil- ture in spring. Larvae overwinter inside the galls lage, 37º18′N, 45º07′E, 1420 m a.s.l., 29 km SE Ur- and pupate during the next winter or spring. The mia) -- a new record for Iran. Europe and Israel pupal stage of A. hypecoi starts in late September (Melika 2006). and continuing until the beginning of November. The imago is formed in October -- November, Genus Barbotinia Nieves-Aldrey, 1994 overwinter in the gall and emerges in March-April Type-species: Aylax oraniensis Barbotin, 1964. Bar- next year (Stojanova & Draganov 2008, Karimpour botinia closely related to Aylax; in both genera the 2011). pronotum relatively short medially, submedian Distribution. Iran, Azerbaijan-e Gharbi prov- pronotal pits transverse and indistinctly sepa- ince (vicinity of Täzehkand-e Qäterchï village, rated. Barbotinia differs from Aylax in that the head 37º39′N, 44º58′E, 1335 m a.s.l., 13 km SW Urmia) is trapezoid, the malar space relatively long; (Karimpour 2011). Turkey (Katilmiş & Kiyak scutellar foveae are large and rounded, nearly as 2011). The species was originally described from long as broad; the mesopleuron with short irregu- Tripoli (North Africa) and recorded from Algeria lar striae, reticulate in between them; the radial and Greece (Trotter 1913). After the synonymiza- cell completely opened along the margin; R1 tion of A. spirorhynchusii the distribution of A. hardly reaching the wing margin; 2nd metasomal hypecoi extends to Transcaucases (Armenia) and tergite without antero-lateral patch of setae. The Middle Asia (Turkmenistan) (Nieves-Aldrey & genus comprises a single species, Barbotinia Melika 2005). Presumably occurs in southern part oraniensis (Barbotin), known to induce galls in fruit of Ukraine also, particularly in Crimea (Melika capsules of Papaver species (Papaveraceae). 2006). Barbotinia oraniensis (Barbotin, 1964) Aylax minor Hartig, 1840 Gall. Galls in fruit capsules, irregularly spherical, Gall. The gall is rounded, 2 mm in diameter, whit- 2.0–3.0 mm in diameter. Usually 2–3 galls devel- ish yellow, attached to septa of the fruit capsule. oping in one fruit capsule, rarely only one or more Galls are often located so near to one another that than 3. The infested capsule is swollen, malformed they are malformed. Sometimes galls cover all externally, caused by the hypertrophy of plant tis- septa in the capsule, however, the latter never sues. malformed and swollen externally. Host plant. Papaver dubium L., P. hybridum L., Host plant. Papaver argemone L., P. rhoeas L., P. rhoeas L. (Nieves-Aldrey 2001, Diakontschuk Papaver dubium L., P. hybridum L. (Papaveraceae) 2003). For the first time recorded also from P. (Ambrus 1974, Kierych 1979, Diakontschuk 2003). argemone L. in Iran. Biology. Galls can be found from July, adults Biology. Galls start to develop in April, ma- emerge next year in May. ture in June; adults emerge next spring. Parnips ni- Distribution. Iran, Lorestan, Shorab -- new re- gripes (Barbotin, 1964), a figitid parasitoid, recently cord for Iran. Europe (Diakontschuk 2003, Melika placed in a new subfamily Parnipinae (Figitidae), 2006) and Turkey (Katilmiş & Kiyak 2011). is frequently associated with B. oraniensis in Europe (Ronquist & Nieves-Aldrey 2001) and here Aylax papaveris (Perris, 1839) for the first time is recorded for Romania and Iran Synonyms. Cynips rhoeadis Bouché, 1834. (authors). Gall. Mainly in fruit capsules, occasionally in Distribution. This is the first record from Iran, the stem. The individual galls are like in A. minor, Azerbaijan-e Gharbi province (Qasemlü valley, vi- but they never form a conglomerate of gall and the cinity of Shïrü Kandï village, 37º18′N, 45º07′E, 1420 infested fruit capsule is never elongated, swollen. m a.s.l., 29 km SE Urmia). Mediterranean region, As a result of galling, the duration of plant life is Aylacini of Iran 273 mentioned also from Algerie (Barbotin 1964). in flower heads of Centaurea species, while others Spain, France and Italy (Nieves-Aldrey 2001a), induce stem galls (Melika 2006). Six species are Ukraine (Melika 2006), Romania (new data, au- represented in the Aylacini fauna of Iran, which thors). from 3 species were described from Iran and cur- rently known only from this country. Genus Hedickiana Nives-Aldrey, 1994 Type species: Aulacidea levantina Hedicke, 1928. Key to Isocolus species of Iran

The genus was erected for a single species, H. le- 1. 2nd metasomal tergite with anterolateral patch of vantina (Hedicke), originally described within Au- white dense setae; galls in stems of Serratula lacidea. In common with Aulacidea, Hedickiana ex- karimpouri hibits a closed (or partially closed) radial cell, but -- 2nd metasomal tergite without anterolateral it can be distinguished from Aulacidea as follows: patch of setae, without or with only few scat- the mesopleuron not clearly transversely striate tered setae 2 and partially irregularly reticulate; the mesoscu- 2. 2nd metasomal tergite smooth, without punc- tum with distinct large piliferous punctures; 2nd tures or with only very indistinct superficial metasomal tergite without a patch of setae anter- sparse micropunctures in posterior 1/3, occupy- olaterally, induces galls on Salvia only. ing 1/3 of metasoma length in dorsal view 3 -- 2nd metasomal tergite with distinct punctures in Hedickiana levantina (Hedicke, 1928) posterior 1/2 to 2/3 4 Gall. The gall is rounded, elongated, spindle 3. Mesoscutum delicately coriaceous-striate, striae shaped or ovoid, usually on stems and branches in in posterior 1/2-1/3 not transversely orientated, a form of a stem swelling, encircling the stem, 10– not raised over surface; lower face with sparse 45 mm long, 8–17 mm in diameter. The surface of white setae and strong irradiating striae; median the young light-green or yellowish-green growing mesoscutal line absent; scutellar foveae rounded, gall is felt-like, with very dense short hairs, with indistinctly delimited posteriorly; pronotum and some weak longitudinal ribs. Mature galls turn mesoscutellum dark brown; galls in flowerheads yellowish brown to grayish, hard, lignified the of Carthamus tinctorius and C. oxycanthus spongy parenchyma also turn hard, like a cork. tinctorius Larval chambers usually nested along the periph- -- Mesoscutum with distinct transverse strong ery of the gall, more or less perpendicularly to the striae, raised above surface; posterior half of in- axis of the stem. Multilocular (Melika 2006). ternotauli area with very weak transverse inter- Host plants. Salvia syriaca L. and S. triloba L. rupted striae, distance between them nearly (Lamiaceae). equal to width of a stria; median mesoscutal line Biology. Larvae overwintering in the gall and absent or in a form of short triangle; pronotum adults emerge next spring. In Iran adults emerge and mesoscutellum black; galls in flowerheads of in April. Centaurea spp. centaureae Distribution. Iran, Azerbaijan-e Gharbi prov- 4. R1 continuing prolong wing margin, 1/3 of ra- ince (vicinity of Kelïsä Kandï village, 37º29′N, dial cell closed; galls in flowerheads of Centaurea 45º01′E, 1600 m a.s.l., 2.5 km SW Urmia), Eastern behen beheni Azarbaijan (Karimpour et al. 2008). Armenia, Cre- -- R1 not reaching wing margin, radial cell opened; te, Israel, Jordan, Syria, Ukraine (Melika 2006, galls in other species of Centaurea and Cirsium Nieves-Aldrey & Massa 2006). 5

5. Internotauli area and parapsides in posterior Genus Isocolus Förster, 1869 2/3 with strong, raised shiny striae, interspaces Type species: Diastrophus scabiosae Giraud, 1859. finely coriaceous, 2.0 or more times longer than The genus Isocolus is defined mainly on the basis width of striae; areolet absent or very indistinct; of the forewing characters: R1 and Rs are not galls in flowerheads of Cirsium and Centaurea reaching the anterior margin, the radial cell clearly cirsii opened or partially closed; cilia on wing margin -- Mesoscutum with more delicate, interrupted absent or very short. Currently 23 palaearctic spe- and less raised transverse striae, distinct in pos- cies of Isocolus are known. Without exception, all terior 1/2-2/3, especially in internotauli area; known Isocolus species induces galls on Asteraceae areolet distinct; galls in stems of Centaurea (Compositae), although the majority induce galls similis 274 G. Melika & Y. Karimpour

Isocolus beheni Melika & Karimpour, 2008 late April until June-August (Diakontschuk 1987). Gall. Galls are scattered between scales at the base Distribution. Iran, Azerbaijan-e Gharbi prov- of the flower head, grayish, monolocular, lignified, ince (vicinity of Kelïsä Kandï village, 37º29′N, 2.2–3.0 mm long and 1.5–2.0 mm in diameter; el- 45º01′E, 1600 m a.s.l., 2.5 km SW Urmia), Lorestan liptical, with slightly rough surface, thin-walled, (Karimpour et al. 2008, Melika & Karimpour 2008). scattered at the base of the flower head. The Ukraine (Melika 2006). flower head is not malformed externally. Usually Isocolus karimpouri Melika & Tavakoli, 2008 more than one larval cell develops within one Gall. Galls are in stems. Galled stems are without flower head (Karimpour et al. 2008). external deformation. Inside the stem slightly Host plants. Centaurea behen L. (Asteraceae). elongated (up to 3.5–4.0 mm) galls are formed, Biology. Galls were collected in early August, with a thin wall which is not detachable from the adult wasps emerged in laboratory conditions by inner tissues of the stem; larval chambers are lo- the end of August. It is possible that in the nature cated along the axis of the stem, never across; in the adult wasps overwinter in the gall and emerge the cavity only one larva is located. only next spring. Host plant. Serratula cerinthifolia (Sm.) Boiss. Distribution. Iran, Azerbaijan-e Gharbi prov- (Asteraceae). ince (Qasemlü valley, vicinity of Shïrü Kandï vil- Biology. Adults emerge in late April, suppos- lage, 37º18′N, 45º07′E, 1420 m a.s.l., 29 km SE Ur- edly overwintering in the larval cell in the plant. mia) (Melika & Karimpour 2008). Comments. There are no other Isocolus species, Isocolus centaureae Diakontschuk, 1982 known to induce such a galls. Some Aylacini spe- Synonyms. Isocolus minutus Diakontschuk, 1982. cies, Panteliella fedtschenkoi (Rübsaamen, 1896), Gall. Galls are between scales in the flower Aulacidea phlomica Belizin, 1959, and Phanacis head, grayish, monolocular, lignified, elliptical, phlomidis Belizin, 1959 induce such kind of galls in and narrowed till the apex, with slightly rough stems of Phlomis tuberosa (Lamiaceae), however, surface, thin-walled, 2.0×1.0 mm, galls when ma- the larval cells (chambers) in the dry stems are ture, freely falling out; the flower head is not mal- free-rolling; its outer wall is separated from the formed externally (Melika 2006). tissues of the stem, even after the gall is mature Host plant. Centaurea diffusa Lam., C. pseudo- (Karimpour et al. 2008). maculosa Dobrocz, C. squarrosa (Boiss.) (As- Distribution. Iran, Azerbaijan-e Gharbi prov- teraceae) (Melika 2006). In Iran for the first time ince (Qasemlü valley, vicinity of Shïrü Kandï vil- was recorded from Centaurea solstitialis L. lage, 37º18′N, 45º07′E, 1420 m a.s.l., 29 km SE Ur- Biology. Adults emerge from early May till mia) (Karimpour et al. 2008). late June (Diakontschuk 1982). In Iran adults Isocolus similis Diakontschuk, 1982 emerge in April. Galls. Galls are in a form of swellings at the base Distribution. Iran, Azerbaijan-e Gharbi prov- of flower heads. ince (vicinity of Arablü village, 37º24′N, 45º15′E, Host plants. Centaurea breviceps Iljin (As- 1280 m a.s.l., 20.5 km SE Urmia, a new record for teraceae). In Iran for the first time was reared from Iran). Ukraine, Turkmenistan, Kazakhstan (Melika C. pseudoscabiosa Boiss. 2006). Biology. Adults emerge in mid May (Dia- Isocolus cirsii Diakontschuk, 1987 kontschuk 1982). Gall. Galls are located at the base of flower head, Distribution. Iran, Azerbaijan-e Gharbi prov- monolocular, with lignified hard walls, 3.0 mm in ince (vicinity of Kelïsä Kandï village, 37º29′N, diameter; up to 28 galls can develop in one flower 45º01′E, 1600 m a.s.l., 2.5 km SW Urmia). Ukraine head. (only in the steppe zone) (Diakontschuk 1982). Host plant. Cirsium arvense (L.) Scop., C. ukrai- nicum Bess., C. vulgare (Savi.) Ten., Onopordum sp., Isocolus tinctorius Melika & Gharaei, 2006 Centaurea behen L. (Asteraceae) (Melika 2006, Me- Gall. Galls are scattered at the base of the flower lika & Karimpour 2008). For the first time re- head. Two types of galls’ location were found. corded also from Centaurea pseudoscabiosa Boiss. & Few galls are formed in the ovary, which is de- Buhse. formed, after the larva hatched from the egg and Biology. In Iran adults emerge in May; for gradually modifying it into a cylindrical whitish Ukraine a longer emerging period was given, from gall, with thick wall and rounded tip and after- Aylacini of Iran 275 wards the growing of the ovary is stopped. As the scribed from this country and are only known to larva grows, the width of the gall increases and exist in Iran. becomes nearly similar to a healthy seed. The gall, Key to Phanacis species of Iran when mature becomes dark brown to blackish. (all Phanacis species known from Iran, the female This type of the gall is very rare. Majority of galls antenna with 11-12 flagellomeres, the male an- are on brackets. Each flower head contains only tenna with 11-13 flagellomeres; the radial cell of one gall, very rarely two. Each bracket has only the forewing closed or partially closed) one gall. The location of galls on brackets varies – from the base to the tip, but more frequently they 1. Pronotum measured along dorsomedian line are formed on the upper part of brackets. Galls at 2.0–2.2 times as short as greatest length the base of brackets are elongate, elliptical and measured on outer margin; mesoscutellum in green when young, with very hard and smooth lateral view strongly humped anteriorly, arised wall, 3.0–6.0 mm in length and 1.2–1.9mm in di- above mesoscutum 2 ameter. After maturing, similarly to the flower -- Pronotum measured along dorsomedian line head, they are become yellow. Galls on free parts usually 3.0 times as short as greatest length of brackets are rounded, spherical, 1.0–2.0 mm in measured on outer margin; mesoscutellum in diameter, or somewhat elongated (2.1–3.5 mm lateral view without hump anteriorly, not long and 1.5–2.0 mm in diameter); green, with elevated over mesoscutum 3 smooth, very hard and thick wall (Melika & 2. R1 going along wing margin and reach at least Gharaei 2006). to 3/4 of radial cell length or radial cell closed; Host plant. Carthamus tinctorius L. (Asteraceae) central elevated area of lower face without deep (Melika & Gharaei 2006). For the first time re- punctures; female metasoma longer than high in corded from C. oxycanthus M. Bieb. lateral view, longer than head+mesosoma; F1 of Biology. Wasps emerge in late May–June (Me- male antenna without placodeal sensilla; stem lika & Gharaei 2006). swelling-like galls on Lactuca lorestanicus Distribution. Iran, Ilam province (Melika and -- R1 reach to half length of radial cell; central Gharaei 2006), Azerbaijan-e Gharbi province (vi- elevated area of lower face with distinct deep cinity of Täzehkand-e Qäterchï, 37º39′N, 44º58′E, punctures; female metasoma nearly as long as 1335 m a.s.l., 13 km SW Urmia; a new record for high in lateral view, shorter than head plus the province). Distributed in Iraq as well (Abdul- mesosoma; F1 of male antenna with placodeal Rassoul 1980). sensilla; stem swelling-like galls on Lactuca irani Genus Phanacis Förster, 1860 3. Pedicel nearly equal in length to F1; F2 shorter Type species: Phanacis centaureae Förster. Syno- than F1; forewing margin with very short or nyms: Gillettea Ashmead, 1897, Timaspis Mayr, without cilia, radial cell closed at most to half 1881, Aylacopsis Hedicke, 1923, Parapanteliella Dia- length of radial cell; galls in stems of Centaurea kontschuk, 1981, Pseudophanacis Diakontschuk, varians 1981. Phanacis is easily distinguishable from other -- Pedicel at least 1.3 or more times longer than F1; genera of Aylacini by the absence of submedian forewing margin with long, distinct cilia; galls in pronotal pits and the reticulate sculpture of the stems of Centaurea heteropappi mesopleuron. The genus has a palaearctic distri- bution with two species, Phanacis hypochoeridis and Phanacis heteropappi Diakontschuk, 1988 P. taraxaci, accidentally introduced to North Amer- Gall. Larval chambers nested in the stem, without ica. The first species was also introduced to South visible external swelling or malformation. Africa. Twenty five species of Phanacis are known Host plant. Heteropappus canescens (Nees) No- from the Palaearctic. Only the sexual generation is vopokr. (Asteraceae) (Diakontschuk 1988). For the known, all species are monovoltine. All Phanacis first time recorded from Centaurea solstitialis L. in species, with a few exceptions, are associated with Iran. Asteraceae, usually galls are in a form of small Biology. In Iran adults emerge in April, in larval cells, hidden in the plant stem, without Turkmenistan emergence is in May (Diakontschuk causing any deformation externally (Melika 2006). 1988) Currently 4 species are known from Iran, two of Distribution. Iran, Azerbaijan-e Gharbi prov- these species (P. irani and P. lorestanicus) were de- ince (vicinity of Arablü village, 37º24′N, 45º15′E, 276 G. Melika & Y. Karimpour

1280 m a.s.l., 20.5 km SE Urmia; a new record for Host plant. Centaurea adpressa L. and C. orien- Iran). Earlier was known only from Turkmenistan talis L. (Asteraceae) (Melika 2006). For the first (Diakontschuk 1988). time recorded from C. pseudoscabiosa Boiss. & Bu- hse in Iran. Phanacis irani (Tavakoli & Melika, 2006) Biology. Adults emerge in May. Gall. Galls usually occur on branches, located at Distribution. Iran, Azerbaijan-e Gharbi prov- an acute angle to the main stem, mainly on the ince (vicinity of Kelïsä Kandï village, 37º29′N, upper shoots. Usually the galls are situated on the 45º01′E, 1600 m a.s.l., 2.5 km SW Urmia; the first end of a branch. The gall is a spindle-like, globular record for Iran). Ukraine (Melika 2006). or ovate, elongate lignified stem-swelling, with average 25–30 mm in length, the largest diameter 12–16 mm; multilocular, green when young, turn Discussion to yellow or greyish white when mature, after dry- ing the colour is white. The parenchyma is Currently 21 species of herb gallwasps from 6 spongy, however, outside the gall is hard. genera (Aulacidea, Aylax, Barbotinia, Hedickiana, Iso- Host plant. Lactuca orientalis Boiss. (= Scariola colus and Phanacis) are known from Iran. The oc- orientalis (Boiss.) (Asteraceae) (Tavakoli & Melika currence of 10 species of herb gallwasps in Iran, 2006). Aulacidea hieracii, A. scorzonerae, A. tragopogonis, Biology. The gall develops very rapidly, start- Aylax minor, A. papaveris, Barbotinia oraniensis, Iso- ing to grow in late spring, mature in June and colus centaureae, I. similis, Phanacis heteropappi, and adults emerge in late June and can be found until P. varians is given for the first time. No doubt, fur- August (Tavakoli & Melika, 2006). ther sampling will increase the number of known Distribution. Iran (Lorestan, Brujerd, Zalian, herb gallwasp species in Iran. Ghelaie; Hamedan Province, Gamasiab; slopes of Alborz Mountains, Damavand, around Tehran in the Lar Valley (Tavakoli & Melika, 2006); Azerbai- References jan-e Gharbi province (vicinity of Kelïsä Kandï vil- lage, 37º29′N, 45º01′E, 1600 m a.s.l., 2.5 km SW Abdul-Rassoul, M.S. (1980): New species of Sycophila Walker from Urmia.). The host plant is distributed also in Syria, Iraq (Hymenoptera, Eurytomidae). Annales Historico-Naturales Musei Nationalis Hungarici 72: 281–284. Israel and Jordan. So, it is quite possible that this Ambrus, B. (1974): Cynipida-Gubacsok-Cecidia Cynipidarum. gallwasp species can be find there as well. Further Fauna Hungaricae 116, XII, Hymenoptera II. fuz.1/a: 1-120 [In sampling in similar habitats is required to Hungarian]. Azizkhani, E., Rasoulian, G.R., Kharazi-Pardel, A., Tavakoli, M., establish its true distribution. Sadeghi, S.E., Melika, G., Stone, G.N., Atkinson, R. (2006): New

species of oak gall wasps from Zagross Mountains of Iran Phanacis lorestanicus (Tavakoli & Melika, 2006) (Hymenoptera: Cynipidae: Cynipini). Folia Entomologica Gall. This species induces galls which are hard to Hungarica 67: 161-197. distinguish from those of P. irani (see the gall de- Barbotin, F. (1964): Sur quelques nouveaux cycles, nouvelles galles, nouvelles especes. Marcellia 37(5): 539-551. scription above). Burks, B.D. (1979): Superfamily Cynipoidea. pp. 1045-1059. In: Host plant. Lactuca orientalis Boiss. (Astera- Krombein, K.V., Hurd, P.D., Jr., Smith, D.R. & Burks, B.D. (eds.), ceae). Catalog of Hymenoptera in America North of Mexico. Symphyta and (Parasitica). Vol. 1. Smithsonian Biology. The same as in P. irani. Both species Institution Press, Washington, D.C. were reared from galls collected on the same Csóka, G., Stone, G.N., Melika, G. (2004): Biology, Ecology and plants. Evolution of gall-inducing Cynipidae. pp. 569-636. In: Raman, A., Schaefer, C.W. & Withers, T.M. (eds) Biology, ecology and Distribution. Iran (Lorestan, Brujerd, Zalian, evolution of gall-inducing . Science Publishers, Inc. Ghelaie) (Tavakoli & Melika 2006). Enfield, New Hampshire, USA. Diakontschuk, L.A. (1982): New cynipid species of the genus Phanacis varians Diakontschuk, 1980 Isocolus Förster (Hymenoptera, Cynipidae). Entomologicheskoye Obozreniye 61(2): 382–391. [In Russian] Synonyms. Phanacis (Pseudophanacis) culmicola Diakontschuk, L.A. (1984): New species of Cynipidae (Cynipidae, Diakontschuk, 1981, Phanacis (Pseudophanacis) o- Hymenoptera) from Georgia. Vestnik Zoologii 3: 74-77. [In rientalis Diakontschuk, 1981, Phanacis (Pseudopha- Russian] Diakontschuk, L.A. (1987): A new species of the genus Isocolus nacis) stepicola Diakontschuk, 1981. Foerster (Hym.: Cynipidae) from the Ukraine. pp. 56-58 in Fauna Gall. In a form of typical small larval cells in- and biocoenotic relationships of of Ukraine. Naukova Dumka, side the stem, without external deformations, as it Kiev. [In Russian] Diakontschuk, L.A. (1988): New and little known gall wasps of the is typical for many Phanacis species (Melika 2006). subfamily Cynipinae (Hymenoptera, Cynipidae) from Central Aylacini of Iran 277

Asia. Entomologicheskoye Obozreniye 67(1): 166–181. [In Nieves-Aldrey, J.L., Massa, B. (2006): Contribution to the Russian] knowledge of the Cynipidae (Hymenoptera) of Jordan. Zoology Diakontschuk, L.A. (2003): New and little known species of in the Middle East 37: 73-82. cynipids (Hymenoptera, Cynipidae) from Ukraine. Vestnik Nieves-Aldrey, J.L., Melika, G. (2005): Aylax hypecoi Trotter Zoologii 37(2): 9-16. [In Russian] (Hymenoptera, Cynipidae) in Europe: Redescription, with Karimpour, Y. (2011): First report of Aylax hypecoi (Hym., taxonomic and biological notes. Journal of Natural History Cynipidae: Aylacini) from Iran with some notes on its biology 39(27): 2525-2535. and associated parasitoids. Journal of Entomological Society of Pénzes, Zs., Melika, G., Bozsóki, Z., Bihari, P., Mikó, I., Tavakoli, Iran 2011: in press [In Persian with English summary]. M., Pujade-Villar, J., Fehér, B., Fülöp, D., Szabó, K., Bozsó, M., Karimpour, Y., Tavakoli, M., Melika, G. (2008): New species of herb Sipos, B., Somogy, K., Stone, G.N. (2009): Systematic re- gallwasps from the Middle East (Hymenoptera, Cynipidae, appraisal of the gall-usurping wasp genus Synophrus Hartig, Aylacini). Zootaxa 1854: 16–32. 1843 (Hymenoptera: Cynipidae: Synergini). Systematic Katilmiş, Y., Kiyak, S. (2011): New records of herb gallwasps Entomology 34: 688-711. (Hymenoptera, Cynipidae, Aylacini) from Turkey. North- Ronquist, F., Nieves-Aldrey, J.L. (2001): A new subfamily of Western Journal of Zoology 7(1): 17-19. Figitidae (Hymenoptera, Cynipoidea). Zoological Journal of the Kierych, E. (1979): Galasowkowate. Cynipoidea. Catalogus faunae Linnean Society 133: 483–494. Poloniae. Panstwowe Wydawnictwo Naukowe, Warszawa Sadeghi, S.E., Assareh, M.H., Tavakoli, M. (2010): Oak gall wasps of 26(2), 104 pp. [In Polish] Iran. Published by Research Institute of Forests and Rangelands, Kovalev, O.V., Diakontschuk, L.A. (1986): A redescription of Tehran. Aulacidea acroptilonica (Cynipidae, Hymenoptera). Vestnik Sadeghi, S.E., Melika, G., Pujade-Villar, J., Pénzes, Zs., Ács, Z., Zoologii 2: 16-19. [In Russian] Bechtold, M., Assareh, M.H., Tavakoli, M., Yarmand, H., Kovalev, O.V. (1965): Gall wasps (Hymenoptera, Cynipidae) from Askary, H., Stone, G.N., Azizkhani, E., Zargaran, M.R., the south of the Soviet Far East. Revue d’Entomologie l,URSS Aligolizade, D., Barimani, H., Dordaei, A.A. (2006): Oak cynipid 44(1/2): 46-73. [In Russian] gall inquilines of Iran (Hym.: Cynipidae: Synergini), with Melika, G. (2004): A new species of from Russian Far East description of new species. Journal of the Entomological Society (Hymenoptera: Cynipiadae, Aylacini). Proceedings of the of Iran 25(2): 15-50. Russian Entomological Society, St. Petersburg 75(1): 214-218. Stojanova, A.M., Draganov, M.M. (2008): Life cycle of Aylax hypecoi Melika, G. (2006): Gall Wasps of Ukraine. Cynipidae. Vestnik (Insecta: Hymenoptera: Cynipidae), a gall inducer on Hypecoum Zoologii, Supplement 21(1): 1–300 and Supplement 21(2): 301– ssp. (Papaveraceae). Central European Journal of Biology 3(2): 644. 199-204. Melika, G., Gharaei, B. (2006): New species of aylacine gall wasps Tavakoli, M., Melika, G. (2006): Two new species of Timaspis Mayr, from Iran (Hymenoptera: Cynipidae: Aylacini). Acta Zoologica 1881 – aylacine gall wasps from Iran (Hymenoptera: Cynipidae: Academiae Scientiarum Hungaricae 52(4): 385–399. Aylacini). Folia Entomologica Hungarica 67: 75–87. Melika, G., Karimpour, Y. (2008): The first report of Aulacidea Tavakoli, M., Melika, G., Sadeghi, S.E., Pénzes, Z., Assareh, M.A., acroptilonica and Isocolus cirsii (Hym., Cynipidae) from Iran. Atkinson, R., Bechtold, M., Mikó, I., Zargaran, M.R., Journal of Entomological Society of Iran 27(2): 19–20. Aligolizade, D., Barimani, H., Bihari, P., Pirozi, F., Fülöp, D., Melika, G., Stone, G.N. (2001): A new species of Cynipid gall wasp Somogyi, K., Challis, R., Preuss, S., Nicholls, J., Stone, G.N. from Turkey (Hymenoptera: Cynipidae). Folia Entomologica (2008): New species of oak gallwaps from Iran (Hymenoptera: Hungarica 62: 127-131. Cynipidae: Cynipini). Zootaxa 1699: 1-64. Melika, G., Stone, G.N., Sadeghi, S.E., Pujade-Villar, J. (2004): New Trotter, A. (1913): Contributo alla conoscenza delle galle della species of cynipid gall wasps from Iran and Turkey Tripoliania. Marcellia 11: 210-219. (Hymenoptera: Cynipidae: Cynipini). Acta Zoologica Zerova, M.D., Diakontschuk, L.A., Ermolenko, V.M. (1988): Academiae Scientiarum Hungaricae 50(2): 139-151. Gallmaker insects of the European part of the USSR, I. Naukova Nieves-Aldrey, J.L. (2001a): Hymenoptera, Cynipidae. In: Fauna Dumka, Kiev, 159 pp. [In Russian] Ibérica, vol. 16. Ramos, M.A. et al. (eds), Museo Nacional de Ciencias Naturales. CSIC. Madrid. 636 pp. Nieves-Aldrey, J.L. (2001b): Nuevos datos faunísticos, corológicos y biológiocos sobre los cinípidos del ámbito Ibero-Balear (Hymenoptera, Cynipidae). Graellsia 57(1): 39-72.