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http://journals.tubitak.gov.tr/zoology/

Turkish Journal of Zoology Turk J Zool(2015) 39: 1121-1131© TÜBİTAKdoi:10.3906/zoo-1402-61

New host and new records of Charipinae(Hymenoptera: Cynipoidea: Figitidae) from Iran

Mar FERRER-SUAY1,*, Jesús SELFA2, Ehsan RAKHSHANI3, Elham NADER3, Juli PUJADE-VILLAR11Department of Animal Biology, Faculty of Biology, University of Barcelona, Barcelona, Spain

2Department of Zoology, Faculty of Biological Sciences, University of Valencia, Burjassot-Paterna Campus, Burjassot, Spain3Department of Plant Protection, College of Agriculture, University of Zabol, Zabol, Iran

* Correspondence: mar.ferrer.suay@gmail.com

1. IntroductionMembers of the subfamily Charipinae are very small wasps (0.8–2.0 mm), characterized by having a smooth and shiny body. The morphological features of these small wasps are much reduced, and for this reason the identification at species level is a very difficult task. On the other hand, the Charipinae are a biologically important group, being secondary parasitoids of aphids (Hemiptera: Aphididae) via Aphidiinae (Hymenoptera: Braconidae) and Aphelininae (Hymenoptera: Braconidae), and secondary parasitoids of psyllids (Hemiptera, Psylloidea) via Encyrtidae (Hymenoptera: Chalcidoidea) (Menke and Evenhuis, 1991).

The biological importance of the Charipinae lies in their lifestyle. As secondary parasitoids of aphids, they have an influence on the aphids’ and primary parasitoids’ abundance. According to van Veen et al. (2001), secondary parasitoids can reduce the efficiency of primary parasitoids on their hosts in at least 3 ways: (i) increasing primary parasitoids’ mortality; (ii) increasing the growth rate of the aphid population indirectly; and (iii) increasing the propensity for primary parasitoids to disperse. Charipinae hyperparasitoids influence the effectiveness of the primary parasitoids of aphids by decreasing their abundance and modifying their behavior. As a result, the increase of aphid populations can cause severe yield losses in some crops (Ferrer-Suay et al., 2012a).

The Charipinae fauna from Iran has been recently revised (Ferrer-Suay et al., 2013e). Fifteen species were identified in that work, 9 of which were recorded for the first time from this region. They included Alloxysta arcuata (Kieffer, 1902), A. brevis (Thomson, 1862), A. castanea (Hartig, 1841), A. macrophadna (Hartig, 1841), A. melanogaster (Hartig, 1840), A. pusilla (Kieffer, 1902), A. ruficollis (Cameron, 1883), A. tscheki (Giraud, 1860), and A. ullrichi (Giraud, 1860). Later, A. ramulifera (Thomson, 1862) was also recorded for the first time from Iran (Ferrer-Suay et al., 2013d).

The objective of the present work was to study additional material that was recently collected from various habitats in 4 central and eastern provinces of Iran. Trophic relationships are also recorded for the collected species. Diagnoses, short descriptions, and figures are included for the newly recorded species.

2. Materials and methodsCollection of specimens was performed mainly in Isfahan (central) and North Khorasan (northeastern) provinces by rearing host aphids, and in Khorasan Razavi and Sistan and Baluchestan provinces using a standard sweeping net during 2007–2011. The plant material bearing the aphid colonies was gently cut, directly placed within transparent plastic boxes, and covered with mesh for ventilation. The

Abstract: Charipinae material collected from various localities in Iran was studied. A total of 712 specimens have been determined in 8 species: Alloxysta arcuata (Kieffer, 1902), A. brevis (Thomson, 1862), A. circumscripta (Hartig, 1841), A. citripes (Thomson, 1862), A. darci (Girault, 1933), A. mullensis (Cameron, 1883), A. pusilla (Kieffer, 1902), and Phaenoglyphis villosa (Hartig, 1841). Alloxysta circumscripta and A. darci are cited for the first time from Iran. In total, 91 host associations (Charipinae-Aphidiinae-Aphid-host plant) are recorded, of which 23 are new to science. New trophic relationships are also recorded for Iran for A. brevis, A. mullensis, and P. villosa. Diagnoses, short descriptions, and figures are included for the newly recorded species.

Key words: Figitidae, Charipinae, Alloxysta, Phaenoglyphis, Iran

Received: 27.02.2014 Accepted/Published Online: 05.09.2015 Printed: 30.11.2015

Research Article

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samples were subsequently transferred to the laboratory, where they were subdivided, with part used to obtain voucher aphids for identification and another part held for rearing the parasitoids and hyperparasitoids. The boxes were kept at room temperature (within a range of 22 ± 5 °C) until emergence of the adult parasitoids and hyperparasitoids. The emerged specimens were captured twice per day using an aspirator to avoid further parasitic activities inside the rearing boxes. Captured specimens were directly dropped into 75% ethanol for preservation and identification at later steps.

The external morphology of Charipinae specimens was studied using a stereomicroscope (NIKON SMZ-1) and an environmental scanning electron microscope (FEI Quanta 200 ESEM). The field-emission gun environmental scanning electron microscope was used for high-resolution imaging without gold-coating the specimens. For each new record for Iran, a diagnosis and short description have been added.

Type material of the species studied is deposited in the following institutions:- BMNH: Natural History Museum (London, United

Kingdom).- MZLU: Lund Museum of Zoology (Lund, Sweden).- NHMA: Natural History Museum (Amiens, France).- QM: Queensland Museum (Brisbane, Australia).- ZSM: Zoologische Staatssammlung Museum (Munich,

Germany). Morphological terms used are taken from Paretas-

Martínez et al. (2007). Measurements and abbreviations include F1–F12, first and subsequent flagellomeres. The width of the forewing radial cell is measured from the margin of the wing to the beginning Rs vein. The transfacial line is measured as the distance between the inner margins of the compound eyes, measured across the face through the antennal toruli, divided by the height of the eye. The malar space is measured by the distance from the lower part of the gena from the mouthparts to the ventral margin of the compound eye, divided by the height of the eye. Females and males of the species shortly described have the same characters except where indicated. Aphid nomenclature follows Remaudiére and Remaudiére (1997). Aphid slides and Aphidiinae specimens are deposited in the insect collection of the Faculty of Agriculture, University of Zabol, Iran (FAUOZ). Charipinae species are deposited in the University of Barcelona (UB) (Pujade-Villar collection). World host range has been included in the host section of each determined species. Host data are listed as follows: HP: host plant; HA: host aphid; HS: host psyllid; HW: primary host parasitoid (wasp); when any of these categories is not known, “unknown” is inserted into the corresponding trophic level.

3. ResultsAmong the 712 examined specimens, there were 8 Charipinae species in association with 19 Aphidiinae parasitoids and 16 aphid species. In total, 91 host associations (Charipinae-Aphidiinae-aphid-host plant) were recorded, of which 23 are new to science. Recorded species are listed below alphabetically. For each species, the list of material examined, including Aphidiinae primary parasitoid, host aphid on host plant, locality date, number of specimens, and name of legator is provided.

Alloxysta arcuata (Kieffer, 1902)Allotria (Allotria) arcuata Kieffer, 1902: 12. Type:

BMNH (examined).Material examined. (64♂ and 68♀). “Lysiphlebus

fabarum – Aphis fabae on Beta vulgaris, Isfahan-Jafarabad, 11-ix-2010”: 4♂ and 2♀; “Diaeretiella rapae, Binodoxys acalephae – Aphis fabae on Phaseolus vulgaris, Isfahan-Flavarjan, 05-xi-2010”: 1♂; “Aphis fabae on Circium vulgare, Isfahan-Fereydonshahr, 04-vi-2011”: 5♂ and 3♀; “Lysiphlebus fabarum – Aphis fabae on Ononis spinosa, Isfahan-Jafarabad, 20-v-2011”: 1♀; “Cucumis sativus, Isfahan-Jafarabad, 11-ix-2010”: 2♂ and 1♀; “Lysiphlebus fabarum – Aphis craccivora Koch on Robinia pseudoacacia, Isfahan-Najvan, 25-iv-2011”: 2♂ and 10♀; “Lysiphlebus fabarum – Aphis craccivora on Sophora mollis, Isfahan-Dourche, 15-iv-2011”: 2♂ and 5♀; “Lysiphlebus fabarum, Aphidius matricariae Haliday – Aphis craccivora on Medicago sativa, Isfahan-Dourche, 15-iv-2011”: 8♂ and 4♀; “Aphidius matricariae, Binodoxys acalephae – Aphis craccivora on Trifolium repens Isfahan-Isfahan”: 5♂; “Binodoxys acalephae – Aphis fabae on Phaseolus vulgaris, Isfahan-Isfahan”: 1♀; “Lysiphlebus fabarum – Brachycaudus tragopogonis (Kaltenbach) on Tragopogon graminfolius, Isfahan-Goldasht,14-iv-2011”: 15♂ and 19♀; “Lysiphlebus fabarum – Aphis craccivora on Robinia pseudoacacia, Isfahan-Dourche, 15-iv-2011”: 2♀; “Lysiphlebus fabarum – Aphis rubiae Narzikulov on Rubia tinctorum, Isfahan-Isfahan”: 3♂ and 1♀; “Praon volucre (Haliday) – Aphis fabae on Phaseolus vulgaris, Isfahan-Najvan, 17-ix-2010”: 10♂ and 4♀; “Binodoxys acalephae – Aphis fabae on Phaseolus vulgaris, Isfahan-Najvan, 10-x-2010”: 1♂ and 4♀; “Lysiphlebus fabarum – Aphis fabae on Phaseolus vulgaris, Isfahan-Filour, 03-ix-2010”: 3♂ and 5♀; “Lysiphlenus fabarum – Aphis fabae on Chenopodium album, Isfahan-Dourche, 21-v-2009”: 2♂ and 4♀, Leg. E. Nader; “Lysiphlebus fabarum – Brachycaudus tragopogonis on Tragopogon gramnifolius, North Khorasan-Shirvan, 25-vi-2007, Leg. S. Kazemzadeh”: 1♂ and 2♀.

Distribution. Cosmopolitan.Hosts. HP: unknown, HA: Myzus cerasi, HW: unknown

(Ionescu, 1969: 268); HP: Ligustrum ovalifolium, HA: Myzus ligustri, HW: unknown (Evenhuis, 1976: 143); HP: unknown, HA: unknown, HW: Ephedrus sp., Ephedrus

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persicae (Evenhuis and Barbotin, 1977: 189). HP: Lilium sp., HA: Macrosiphum sp., HW: unknown; HP: Canna sp., HA: Macrosiphum euphorbiae, HW: unknown; HP: unknown, HA: Aphis fabae, HW: unknown; HP: Zea mays, HA: Rhopalosiphum padi, HW: unknown; HP: Bromus sp., HA: Rhopalosiphum padi, HW: Lysiphlebus testaceipes; HP: Cassia sp., HA: Aphis gossypii, HW: unknown; HP: Zea mays, HA: Rhopalosiphum padi, HW: unknown; HP: Citrus limon, HA: unknown, HW: unknown; HP: unknown, HA: Myzus persicae, HW: unknown; HP: Jacaranda sp., HA: Aphis gossypii, HW: unknown; HP: Populus tremuloides, HA: Chaitophorus sp., HW: unknown; HP: Nerium oleander, HA: Aphis nerii, HW: unknown; HP: Malva sp., HA: Aphis sp., HW: unknown; HP: unknown, HA: Aphis gossypii, HW: Lysiphlebus testaceipes; HP: Iris sp., HA: Dysaphis sp., HW: unknown; HP: unknown, HA: Myzus persicae, HW: unknown; HP: unknown, HA: Mentha piperita var. citrata, HW: unknown; HP: unknown, HA: Mentha pulegium, HW: unknown; HP: unknown, HA: Mentha spicata var. tashkent, HW: unknown (Ferrer-Suay et al., 2013a: 30).

Alloxysta brevis (Thomson, 1862)Allotria brevis Thomson, 1862: 408. Type: MZLU

(examined).Material examined. (68♂ and 80♀). “Lysiphlebus

fabarum – Aphis fabae on Beta vulgaris, Isfahan-Jafarabad, 11-ix-2010”: 1♀; “Diaeretiella rapae, Binodoxys acalephae – Aphis fabae on Phaseolus vulgaris, Isfahan-Flavarjan, 05-xi-2010”: 3♂ and 2♀; “Cucumis sativus, Isfahan-Jafarabad, 11-ix-2010”: 1♀; “Lysiphlebus fabarum, Aphidius matricariae – Aphis craccivora on Medicago sativa, Isfahan-Dourche, 15-iv-2011”: 4♀; “Aphis fabae on Phaseolus vulgaris, Isfahan-Flavarjan, 05-xi-2010”: 1♀; “Aphidius matricariae, Lysiphlebus fabarum – Aphis umbrella on Malva sylvestris, Isfahan-Najafabad, 06-vi-2011”: 1♀; “Adialytus ambiguus – Sipha elegans on Gastridium phleoides, Isfahan-Najvan, 09-v-2011”: 1♀; “Lysiphlebus fabarum – Aphis origani on Mentha longifolia, Isfahan-Varposht, 26-v-2011”: 1♀; “Aphidius matricariae, Binodoxys acalephae – Aphis craccivora on Trifolium repens, Isfahan-Isfahan,”: 1♂ and 5♀; “Aphidius matricariae, Lysiphlebus fabarum – Aphis craccivora on Trifolium campestre, Isfahan-Isfahan”: 3♂ and 1♀; “Adialytus salicaphis (Fitch) – Chaitophorus sp. on Populus alba, Isfahan-Mobarakeh, 13-xi-2010”: 1♂; “Diaeretiella rapae, Aphidius matricariae – Aphis sp. on Brassicaceae host plant, Isfahan-Dourche, 15-iv-2011”: 1♀; “Lysiphlebus fabarum – Aphis rubiae on Rubia tinctorum, Isfahan-Isfahan”: 8♂ and 20♀; “Binodoyxs acalephae – Aphis fabae on Phaseolus vulgaris, Isfahan-Hoye, 15-x-2010”: 1♂; “Aphidius matricariae – Aphis solanella Theobald on Solanum melongena, Isfahan-Najvan, 10-x-2010”: 1♂ and 1♀; “Praon volucre – Aphis fabae on Phaseolus vulgaris, Isfahan-Najvan, 17-ix-2010”:

37♂ and 27♀; “Diaeretiella rapae, Aphidius matricariae – Aphis gossypii on Solanum melongena, Isfahan-Hoye, 18-vii-2009”: 5♂ and 2♀; “Praon sp., Lysiphlebus fabarum – Aphis solanella on Chenopodium album, Isfahan-Dourche, 01-vi-2009”: 5♂ and 4♀; “Binodoxys acalephae – Aphis fabae on Phaseolus vulgaris, Isfahan-Najvan, 10-x-2010”: 1♂ and 1♀; “Lysiphlebus fabarum – Aphis craccivora on Medicago sativa, Isfahan-Najvan, 25-iv-2011”: 1♂ and 3♀; “Lysiphlebus fabarum – Aphis fabae on Phaseolus vulgaris, Isfahan-Filour, 03-ix-2010”: 2♀; “Lysiphlebus fabarum – Aphis fabae on Chenopodium album, Isfahan-Dourche, 21-v-2009”: 1♂ and 1♀, Leg. E. Nader.

Distribution. Cosmopolitan.Hosts. HP: Lycopersicum esculentum, HA: unknown,

HW: unknown (Dalla Torre and Kieffer, 1910: 290). HP: Populus trichocarpa, HA: Chaitophorus populicola Thomas, HW: Aphelinus sp. (Andrews, 1978: 68). HP: unknown, HA: Myzus persicae, HW: Diaeretiella rapae and Aphidius sp. (Horn 1984: 19). HP: unknown, HA: Aphis sp., HW: Lysephedrus sp.; HP: unknown, HA: Myzus cerasi and Dysaphis plantaginea, HW: Ephedrus persicae (Fergusson 1986: 18). HP: unknown, HA: Aphis sp., HW: Lysephedrus sp.; HP: unknown, HA: Myzus cerasi and Dysaphis plantaginea, HW: Ephedrus sp. (Barczak, 1991: 92). HP: unknown, HA: Hyperomyzus lactucae, HW: Praon volucre (Tizado and Nuñez-Perez, 1993: 97). HP: unknown, HA: Eucallipterus tiliae, HW: Trioxys curvicaudus and Trioxys tenuicaudus (Zuparko and Dahlsten, 1995: 730). HP: Citrus sp., HA: Toxoptera citricidus, HW: Lysiphlebus testaceipes (Evans and Stange, 1997: 1). HP: unknown, HA: Capitophorus carduinus and Sitobion sp. (Müller et al., 1999: 346). HP: Euonymus europaea, HA: Aphis fabae, HW: Binodoxys angelicae (Hübner et al., 2002: 508). HP: Solidago altissima, HA: Uroleucon nigrotuberculatum, HW: Aphelinus albipodus (Takada and Nakamura, 2010: 270). HP: Cassia sp., HA: Aphis gossypii, HW: unknown; HP: Yucca sp., HA: Aphis helianthi Monell, HW: unknown (Ferrer-Suay et al., 2013a: 32).

Comments. Alloxysta brevis is here recorded for the first time in association with Binodoxys acalephae and Aphidius matricariae.

Alloxysta circumscripta (Hartig, 1841) (Figures 1a–1d)Xystus circumscriptus Hartig, 1841: 352. Type: ZSM

(examined). Material examined: (17♂ and 18♀). “Praon yomenae,

Ephedrus niger – Uroleucon sp. on Acroptilon repens, Isfahan-Filour, 09-iv-2011”: 17♂ and 18♀; “Praon yomenae, Aphidius persicus – Uroleucon sonchi (L.) on Sonchus oleraceus, Isfahan-Najafabad, 29-x-2010”: 1♂; “Uroleucon sonchi on Sonchus oleraceus, Isfahan-Najafabad, 07-vii-2010”: 1♀, Leg. E. Nader.

Diagnosis. Alloxysta circumscripta is mainly characterized by having a closed radial cell being 2.3 times

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as long as wide (Figure 1a), pronotal carinae present (Figure 1c), propodeal carinae absent (Figure 1d), female antennae with the beginning of rhinaria in F5, F2 shorter than F3, F3 shorter than F4 (Figure 1b), male antennae with the beginning of rhinaria in F4, F2 longer than F3, F3 shorter than F4. It is similar to A. consobrina (Zetterstedt, 1838), but they can be differentiated by the proportions between flagellomeres: F1 subequal to F2, F2 shorter or subequal to F3 in A. circumscripta (Figure 1b) while F1 longer than F2, F2 subequal to F3 in A. consobrina (Zetterstedt, 1838); size of radial cell: 2.3 times as long as wide in A. circumscripta (Figure 1a) but 2.7 times as long as wide in A. consobrina.

Short description. Head, mesosoma, and metasoma dark brown; scape, pedicel, F1, and F2 yellow, F3–F12 yellowish brown; legs yellow and veins yellowish brown. Female antennae 13-segmented; F1–F2 smooth, thinner

than remaining flagellomeres, F3–F11 with rhinaria and club-shaped; F1 longer than pedicel and F2, F2 shorter then F3, F3 shorter than F4 (Figure 1b). Male antennae 14-segmented; F1–F3 smooth and thinner than remaining flagellomeres, F4–F12 with rhinaria and club-shaped; F1–F3 straight; F1 longer than pedicel and F2, F2 longer than F3, and F3 shorter than F4. Pronotum covered by many setae with 2 carinae clearly present under the pubescence (Figure 1c). Propodeum also covered by abundant setae, without carinae present (Figure 1d). Forewing longer than body; radial cell partially open, 2.3 times as long as wide (Figure 1a).

Distribution. Andorra (Ferrer-Suay et al., 2011a: 354); Austria (Giraud, 1860: 127; Hellén, 1963: 17); England (Dalla Torre and Kieffer, 1910: 278; Müller et al., 1999: 346); Finland (Hellén, 1963: 17); France (De Gaulle, 1908:

Figure 1. Diagnostic features of Alloxysta circumscripta: a) radial cell; b) antenna; c) pronotum; d) propodeum.

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26); Germany (Hartig, 1841: 352); Poland (Kierych, 1979: 15); Scotland (Cameron, 1886: 86). New record from Iran.

Host. HP: Raphanus sp., HA: Aphis sp., HW: unknown (Giraud, 1877: 416). HP: Fennel sp., HA: Aphis sp., HW: unknown; HP: Chaerophyllum sylvestre, HA: Cinara pini (L.), Cryptomyzus ribis (L.), and Aphis sambuci (L.) (De Gaulle, 1908: 26; Dalla Torre and Kieffer, 1910: 277). HP: unknown, HA: Uroleucon sp., HW: Praon dorsale (Haliday) auct. (Evenhuis, 1982: 22). HP: unknown, HA: Acyrthosiphon pisum (Harris), HW: Praon sp. (Müller et al., 1999: 352).

Comments. Alloxysta circumscripta is here recorded for the first time in association with Praon yomenae, Ephedrus niger, and Aphidius persicus.

Alloxysta citripes (Thomson, 1862)Allotria citripes Thomson, 1862: 410. Type: MZLU

(examined).Material examined. (18♂ and 20♀). “Trioxys pallidus

– Chromaphis juglandicola on Juglans regia, Isfahan-Najafabad, 02-v-2011”: 1♂ and 2♀; “Trioxys pallidus – Chromaphis juglandicola on Juglans regia, Isfahan-Mohamadiye, 20-v-2011”: 1♂ and 3♀; “Trioxys pallidus – Chromaphis juglandicola on Juglans regia, Isfahan-Hoye, 15-v-2011”: 11♂ and 9♀; “Trioxys pallidus – Chromaphis juglandicola on Juglans regia, Isfahan-Varposht, 28-v-2011”: 4♂ and 4♀; “Trioxys pallidus – Chromaphis juglandicola on Juglans regia, Isfahan-Najafabad, 29-v-2011”: 1♂ and 2♀, Leg. E. Nader.

Distribution. Holarctic, Neotropic, and African (Ferrer-Suay et al., 2012a).

Host. HP: unknown, HA: Eucallipterus tiliae, HW: unknown (De Gaulle, 1908: 26). HP: unknown, HA: Dysaphis mali, HW: unknown (Belizin, 1966: 6). HP: Quercus sp., HA: Tuberculoides annulatus, HW: unknown; HP: Alnus glutinosa, HA: Pterocallis alni, HW: unknown; HP: Corylus avellana, HA: Myzocallis coryli, HW: unknown; HP: unknown, HA: unknown, HW: Trioxys pallidus (Evenhuis, 1976: 140). HP: unknown, HA: Tuberculoides sp., Pterocallis sp., and Myzocallis sp., HW: Trioxys sp.; HP: unknown, HA: Drepanosiphum sp., HW: Aphelinus sp. (Fergusson, 1986: 18). HP: unknown, HA: Eucalipterus tiliae, HW: unknown (Evenhuis and Barbotin, 1987: 214). HP: Tilia cordata, HA: Eucallipterus tiliae, HW: Trioxys pallidus (Hübner et al., 2002: 508). HP: Juglans regia, HA: Chromaphis juglandicola, HW: unknown (Ferrer-Suay et al., 2012c: 10).

Alloxysta darci (Girault, 1933) (Figures 2a–2d)Allotria d’arci Girault, 1933: 2. Type: QM (examined).Material examined: (6♂ and 2♀). “Swept on Medicago

sativa, Khorasan Razavi-Sabzevar-Joghatay, 08-ix-2011”: 5♂ and 1♀, Leg. Kh. Fathabadi; “Swept on Medicago sativa, Sistan & Baluchestan-Zabol, 09-vi-2011”: 1♂ and 1♀, Leg.: F. Goli-Mahmoudi.

Diagnosis. Alloxysta darci is mainly characterized by having small closed radial cell, being 2.4 times as long as wide (Figure 2a), pronotal carinae absent (Figure 2b), propodeal plate present (Figure 2d), rhinaria and club shape begin in F4, F1 shorter than pedicel, and antennae longer than body length in both male and female (Figure 2c). This species is similar to A. brevis, but they can be differentiated by the antennae length: longer than body in A. darci, while shorter in A. brevis; forewing with marginal setae longer in A. darci, while they are shorter in A. brevis.

Short description. Head yellowish brown, mesosoma and metasoma brown; scape, pedicel, F1–F3 yellow, rest of flagellomeres brown; legs yellowish; veins yellowish brown. Female antennae 13-segmented; F1–F3 smooth and thinner than remaining ones; F4–F11 with rhinaria and club shape; F1 shorter than pedicel, F1–F3 subequal, F3 shorter than F4 (Figure 2c). Male antennae 14-segmented; F1–F3 smooth and thinner than remaining ones; F4–F12 with rhinaria and club shape, with the same proportions between flagellomeres as female. Pronotum covered with setae with few setae on distolateral corners, without carinae present (Figure 2b). Propodeum covered by abundant pubescence; 2 wide carinae present forming a plate, separated by a few setae on first third and forming a plate on last two-thirds; sides subparallel anteriorly and very slightly curved posteriorly (Figure 2d). Forewing longer than body, radial cell closed, 2.4 times as long as wide in both sexes (Figure 2a).

Distribution. Species known from the Palearctic and the Australian region (Ferrer-Suay et al., 2012a). Spain (Ferrer-Suay et al., 2013c: 323). New record from Iran.

Host. HP: unknown, HA: Aphis craccivora and Hyalopterus pruni (Geoffroy), HW: Aphelinus flaviventris Kurdjumov; HP: unknown, HA: Macrosiphum euphorbiae, HW: Aphelinus gossypii Timberlake; HP: unknown, HA: Myzus persicae, HW: Aphelinus gossypii; HP: unknown, HA: unknown, HW: Aphelinus varipes (Förster) (Carver, 1992: 770). HP: Sorghum bicolor, HA: Rhopalosiphum maidis (Fitch), HW: Aphelinus varipes.

Alloxysta mullensis (Cameron, 1883)Allotria mullensis Cameron, 1883: 366. Type: BMNH

(examined).Material examined. (3♂ and 16♀). “Lysiphlebus

fabarum – Aphis fabae on Circium vulgare, Isfahan-Fereydonshahr, 04-vi-2011”: 4♀; “Lysiphlebus fabarum – Aphis fabae on Ononis spinosa, Isfahan-Jafarabad, 20-v-2011”: 2♀; “Aphidius matricariae, Lysiphlebus fabarum – Aphis umbrella on Malva sylvestris, Isfahan-Najafabad, 06-vi-2011”: 1♀; “Aphidius matricariae, Binodoxys acalephae – Aphis craccivora on Trifolium repens, Isfahan-Isfahan,”: 1♂ and 1♀; “Trioxys pallidus – Chromaphis juglandicola on Juglans regia, Isfahan-Varposht, 28-v-2011”: 1♂ and 1♀; “Lysiphlebus fabarum – Aphis fabae and Brachycaudus

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cardui (L.) on Circium arvense, Isfahan-Varposht, 28-v-2011”: 1♂; “Lysiphlebus fabarum – Aphis craccivora on Robinia pseudoacacia, Isfahan-Dourche, 15-iv-2011”: 3♀; “Aphidius matricariae, Lysiphlebus fabarum – Aphis craccivora on Trifolium campestre, Isfahan-Isfahan”: 1♀; “Diaeretiella rapae, Aphidius matricariae – Aphis sp. on Brassicaceae host plant, Isfahan-Dourche, 15-iv-2011”: 1♀; “Lysiphlebus fabarum – Aphis fabae on Phaseolus vulgaris, Isfahan-Filour, 03-ix-2010”: 2♀.

Distribution. Cosmopolitan, except for Australia and Oriental regions (Ferrer-Suay et al., 2012a).

Host. HP: Urtica dioica, HA: Aphis urticata, HW: Lysiphlebus fabarum; HP: Philadelphus coronarius, HA: Aphis fabae, HW: Lysiphlebus fabarum (Evenhuis, 1978: 173). HP: unknown, HA: unknown, HW: Praon sp. and

Ephedrus sp. (Bokina, 1997: 435). HP: Brassica oleracea, HA: unknown, HW: unknown (Ferrer-Suay et al., 2013a: 37).

Comments. Alloxysta mullensis is here recorded for the first time in association with Binodoxys acalephae.

Alloxysta pusilla (Kieffer, 1902)Allotria (Allotria) pusilla Kieffer, 1902: 13. Type:

NHMA. Material examined. (1♂). “Lysiphlebus fabarum –

Aphis fabae on Ononis spinosa, Isfahan-Jafarabad, 20-v-2011”: 1♂, Leg. E. Nader.

Distribution. Palearctic and Neotropic (Ferrer-Suay et al., 2012a).

Host. HP: Chrysanthemum leucanthemum, Salix viminalis and Populus alba, HA: Aphis sp., HW: unknown

Figure 2. Diagnostic features of Alloxysta darci: a) fore wing; b) pronotum; c) antenna; d) propodeum.

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(Kieffer, 1902: 13). HP: Chrysanthemum sp., Populus sp. and Salix sp., HA: Aphis sp., HW: unknown (De Gaulle, 1908: 26). HP: Cirsium arvense, HA: Aphis fabae cirsiiacanthoidis Scopoli, HW: Lysiphlebus cardui (Marshall) (Hübner et al., 2002: 509).

Phaenoglyphis villosa (Hartig, 1841)Xystus villosus Hartig, 1841: 353. Type: ZSM

(examined).Material examined. (118♂ and 152♀). “Lysiphlebus

fabarum – Aphis craccivora on Amaranthus retroflexus, Isfahan-Najvan, 17-ix-2010”: 2♀; “Diaeretiella rapae, Binodoxys acalephae – Aphis fabae on Phaseolus vulgaris, Isfahan-Flavarjan, 05-xi-2010”: 1♂; “Lysiphlebus fabarum – Nearctaphis bakeri on Trifolium campestre, Isfahan-Hoye, 15-v-2011”: 1♂ and 4♀; “Diaeretiella rapae – Lipaphis pseudobrassicae on Brassica napus, Isfahan-Goldasht, 14-iv-2011”: 2♂ and 1♀; “Diaeretiella rapae, Binodoxys acalephae – Aphis fabae on Phaseolus vulgaris, Isfahan-Flavarjan, 05-xi-2010”: 5♂ and 3♀; “Lipaphis pseudobrassicae on Capsella bursa-pastoris, Isfahan-Dourche, 15-iv-2011”: 1♀; “Lysiphlebus fabarum – Aphis affinis on Mentha longifolia, Isfahan-Dourche, 15-iv-2011”: 3♂ and 4♀; “Aphidius matricariae, Praon abjectum – Aphis solanella on Solanum melongena, Isfahan-Hoye, 15-x-2010”: 4♂and 1♀; “Adialytus veronicaecola – Aphis gossypii on Cucurbita pepo, Isfahan-Ghahderijan, 05-xi-2010”: 5♂ and 3♀; “Lysiphlebus fabarum – Aphis craccivora on Robinia pseudoacacia, Isfahan-Najvan, 25-iv-2011”: 2♀; “Lysiphlebus fabarum – Aphis craccivora on Sophora mollis, Isfahan-Dourche, 15-iv-2011”: 7♂ and 6♀; “Lysiphlebus fabarum, Aphidius matricariae – Aphis craccivora on Medicago sativa, Isfahan-Dourche, 15-iv-2011”: 11♂ and 12♀; “Aphidius matricariae, Diaeretiella rapae – Myzus persicae on Brassica napus, Isfahan-Hoye, 15-x-2010”: 5♂and 7♀; “Lysiphlebus fabarum – Aphis fabae on Beta vulgaris, Isfahan-Flavarjan, 05-xi-2010”: 2♀; “Aphidius transcaspicus – Hyalopterus pruni on Phragmites australis, Isfahan-Mobarakeh, 13-xi-2010”: 2♀; “Praon volucre, Aphidius matricariae, Lysiphlebus fabarum, Diaeretiella rapae – Schizaphis graminum on Hordeum vulgare, Isfahan-Goldasht, 14-ix-2011”: 3♂ and 1♀; “Aphidius sp. and Lysiphlebus fabarum on Viola tricola, Isfahan-Najafabad, 06-vi-2011”: 2♀; “Adialytus veronicaecola, Binodoxys sp. – Aphis rubiae on Rubia tinctorum, Isfahan-Mobarakeh, 3-xi-2010”: 27♂ and 54♀; “Aphidius transcaspicus – Hyalopterus pruni on Phragmites australis, Isfahan-Mobarakeh, 03-xi-2011”: 3♂ and 2♀; “Lysiphlebus fabarum – Aphis craccivora on Robinia pseudoacacia, Isfahan-Dourche, 15-iv-2011”: 1♂ and 1♀; “Aphidius matricariae, Lysiphlebus fabarum – Aphis craccivora on Trifolium campestre, Isfahan-Isfahan”: 2♀; “Adialytus salicaphis – Chaitophorus sp. on Populus alba, Isfahan-Mobarakeh, 13-xi-2010”: 19♂ and 21♀;

“Diaeretiella rapae, Aphidius matricariae – Aphis sp. on Brassicaceae host plant, Isfahan-Dourche, 15-iv-2011”: 1♀; “Diaeretiella rapae, Lysiphlebus fabarum – Aphis craccivora on Cardaria draba, Isfahan-Dourche, 15-iv-2011”: 7♂ and 1♀; “Praon yomenae, Aphidius persicus – Uroleucon sonchi on Sonchus oleraceus, Isfahan-Najafabad, 29-x-2010”: 1♂; “Praon rosaecola, Aphidius rosae – Macrosiphum rosae on Rosa sp. Isfahan-Najvan, 10-x-2010”: 4♀; “Aphidius matricariae – Aphis gossypii on Solanum melongena, Isfahan-Najvan, 10-x-2010”: 1♀; “Praon volucre – Aphis fabae on Phaseolus vulgaris, Isfahan-Najvan, 17-ix-2010”: 2♂ and 2♀; “Binodoxys acalephae – Aphis fabae on Phaseolus vulgaris, Isfahan-Najvan, 10-x-2010”: 1♂ and 1♀; “Lysiphlebus fabarum – Aphis craccivora on Medicago sativa, Isfahan-Najvan, 25-iv-2011”: 5♂ and 2♀; “Praon barbatum, Aphidius smithi – Acyrtosiphon pisum on Medicago sativa, Isfahan-Hoye, 27-vii-2010: 1♂ and 2♀; “Lysiphlebus fabarum – Aphis fabae on Phaseolus vulgaris, Isfahan-Filour, 03-ix-2010”: 4♂ and 3♀; “Lysiphlebus fabarum – Aphis origani on Mentha longifolia, Isfahan-Najafabad, 27-iv-2011”: 2♀, Leg. E. Nader.

Distribution. Cosmopolitan (Ferrer-Suay et al., 2012a).Host. HP: Sonchus sp., HA: Aphis sp., HW: unknown;

HP: Sinapis sp., HA: Aphis sp., HW: unknown; HP: Aegopodium sp., Alisma sp., and Platanus sp., HA: Aphis sp., HW: unknown (De Gaulle, 1908: 26). HP: unknown, HA: Aphis ambrosia, HW: unknown (Dalla Torre and Kieffer, 1910: 289). HP: Sinapis alba, HA: Aphis sp., HW: unknown (Dalla Torre and Kieffer, 1910: 269). HP: Sonchus asper, HA: Aphis sp., HW: unknown (Dalla Torre and Kieffer, 1910: 268). HP: Platanus sp., Alisma plantago, and Aegopodium podagraria, HA: Aphis sp., HW: unknown (Dalla Torre and Kieffer, 1910: 278). HP: Aulacorthum pelargonii, HA: Myzus ornatus and Rhopalodiphoum padi, HW: unknown (Valentine, 1975: 60). HP: Quercus sp., HA: Tuberculoides annulatus, HW:  Trioxys pallidus; HP: Berberis sp., HA: Liosomaphis berberidis, HW: Aphidius hortensis (Vasileva-Sumnalieva, 1975: 24). In the Netherlands, the following combinations were established by Evenhuis: HP: Quercus robur, HA: Tuberculoides annulatus, HW: Praon flavinode; HP: Acer campestre, HA: Periphyllus sp., HW: Trioxys falcatus Mackauer; HP: Tilia sp., HA: Eucallipterus tiliae, HW: an unidentified species of Aphelinidae; HP: Malus sylvestris, HA: Rhopalosiphum insertum, HW: Monoctonus cerasi; HP: Prunus sp., HA: unknown, HW: unknown; HP: Phragmites australis, HA: Hyalopterus pruni, HW: Praon volucre; HP: Cirsium arvense, HA: Aphis sp., HW: Lysiphlebus sp; HP: Malus sylvestris, HA: Dysaphis plantaginea, HW: Ephedrus persicae; HP: Sinapis arvensis, HA: Brevicoryne brassicae, HW: Diaeretiella rapae; HP: Berberis vulgaris, HA: Liosomaphis berberidis, HW: Trioxys hortorum; HP: Sonchus asper and Sonchus oleraceus, HA: Hyperomyzus lactucae, HW: Aphidius sonchi Marshall; HP: Galeopsis

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tetrahit, HA: Cryptomyzus galeopsidis, HW: Aphidius ribis; HP: Rosa sp., HA: Macrosiphum rosae, HW: Aphidius rosae; HP: Campanula sp., HA: Uroleucon campanulae, HW: Binodoxys centaureae; HP: Urtica dioica, HA: Microlophium urticae, HW: Aphidius ervi; HP: Poaceae, HA: Metopolophium dirhodum, HW: Aphidius uzbekistanicus. In France, the combinations established by Barbotin and identified by Dr. G. Remaudière are: HP: Poa annua, HA: Rhopalosiphum padi, HW: Ephedrus plagiator, Trioxys auctus, and Aphidius sp.; HP: Mahonia aquifolium, HA: Liosomaphis berberidis, HW: Aphelinus sp.; HP: Leontodon sp., HA: Nasonovia ribisnigri, HW: Aphidius hieraciorum Starý; HP: Hordeum murinum, HA: Sitobion avenae, HW: Aphidius sp.; HP: Statice sp., HA: Staticobium sp., HW: Praon sp.; HP: Cyperaceae, HA: Rhopalosiphum insertum, HW: unknown; HP: Lapsana communis, HA: Myzus ornatus, HW: unknown; HP: Gomphocarpus fruticosus, HA: Aphis nerii, HW: unknown (Evenhuis and Barbotin, 1977: 185). HP: Artemisia douglasiana, HA: Uroleucon ambrosiae, HW: Lysiphlebus testaceipes (Andrews, 1978: 32). HP: unknown, HA: Myzus persicae, HW: Diaeretiella rapae and Aphidius sp. (Hor, 1984: 19). HP: unknown, HA: Therioaphis trifolii, HW: Trioxys complanatus Quilis (Wilson and Swincer, 1984: 47). HP: unknown, HA: Aphis fabae, HW: Ephedrus persicae, Binodoxys angelicae, and Lysiphlebus confusus (Al-Jassani and Al-Adil, 1986: 59). HP: unknown, HA: Acyrthosiphon kondoi, HW: Aphidius ervi and Ephedrus plagiator; HP: unknown, HA: Aphis craccivora, HW: Aphidius colemani; HP: unknown, HA: Aphis gossypii, HW: Aphidius colemani; HP: unknown, HA: Aphis nerii, HW: Aphidius colemani; HP: unknown, HA: Aphis spiraecola, HW: Aphidius colemani; HP: unknown, HA: Brachycaudus helichrysi, HW: Aphidius colemani; HP: unknown, HA: Brachycaudus persicae, HW: Aphidius colemani; HP: unknown, HA: Cavariella aegopodii, HW: Aphidius salicis; HP: unknown, HA: Hyadaphis foeniculi, HW: Aphidius colemani; HP: unknown, HA: Hyperomyzus lactucae, HW: Aphidius sonchi; HP: unknown, HA: Myzus cerasi, HW: Aphidius colemani; HP: Prunus persicae, HA: Myzus persicae, HW: Aphidius colemani; HP: unknown, HA: Myzus persicae, HW: Diaeretiella rapae; HP: unknown, HA: Myzus persicae, HW: Ephedrus persicae; HP: unknown, HA: Rhopalosiphum maidis, HW: Aphidius colemani; HP: unknown, HA: Rhopalosiphum padi, HW: Aphidius colemani; HP: unknown, HA: Rhopalosiphum padi, HW: Aphidius similis; HP: unknown, HA: Therioaphis trifolli, HW: Trioxys complanatus; HP: unknown, HA: Toxoptera citricidus, HW: Aphidius colemani (Carver, 1992: 770). HP: unknown, HA: unknown, HW: Aphidius sp., Ephedrus sp., Praon sp., and Trioxys sp. (Höller et al., 1993: 15). HP: unknown, HA: unknown, HW: Aphidius sp., Praon sp., and Ephedrus sp. (Bokina, 1997: 435). HP: Rosa sp., HA: Macrosiphum rosae, HW: unknown; HP: Asparagus

maritimus, HA: Myzus persicae, HW: unknown; HP: Sonchus tenerrimus, HA: unknown, HW: unknown (Suay et al., 1998: 106). HP: unknown, HA: Capitophorus carduinis (sic), HW: unknown; HP: unknown, HA: Sitobion sp., HW: Aphidius sp., Ephedrus sp., and Aphelinus sp. (Müller et al., 1999: 352). HP: Salix caprea, HA: Aphis farinosa Gmelin, HW: Lysiphlebus confusus (Hübner et al., 2002: 509). HP: Acer pseudoplatanus, HA: Drepanosiphum platanoides, HW: Trioxys cirsii; HP: Rapistrum rugosum, HA: Lipaphis pseudobrassicae, HW: Diaeretiella rapae; HP: Quercus canariensis, HA: Myzocallis castanicola, HW: Trioxys tenuicaudus; HP: Triticum aestivum, HA: Rhopalosiphum padi, HW: Aphidius colemani (Carver, 2004: 2). HP: Solidago altissima, HA: Uroleucon nigrotuberculatum, HW: Ephedrus plagiator (Takada and Nakamura, 2010: 270). HP: unknown, HA: Acyrthosiphon pisum, HW: unknown; HP: Nothofagus sp., HA: unknown, HW: unknown; HP: kiwifruit orchard, HA: unknown, HW: unknown (Ferrer-Suay et al., 2012b: 238). HP: Lilium sp., HA: Macrosiphum euphorbiae, HW: unknown; HP: Bromus sp., HA: Rhopalosiphum padi, HW: Lysiphlebus testaceipes; HP: Bromus sp., HA: Metopolophium dirhodum, HW: unknown; HP: Bromus sp., HA: Sitobion avenae, HW: unknown; HP: Bromus sp., HA: Rhopalosiphum padi, HW: unknown; HP: Canna sp., HA: unknown, HW: Aphidius sp.; HP: Alfalfa, HA: unknown, HW: Lysiphlebus testaceipes (Ferrer-Suay et al., 2013a: 40).

Host remarks. For the following trophic relationships, HP: Berberis vulgaris, HA: Liosomaphis berberidis, HW: Trioxys hortorum (Evenhuis and Barbotin, 1977), the parasitoid species is most probably Aphidius hortensis, which has been incorrectly written as Trioxys hortorum.

Comments. Phaenoglyphis villosa is here recorded for the first time in the following associations: Lysiphlebus fabarum – Aphis affinis, Aphis craccivora, and Nearctaphis bakeri; Binodoxys acalephae – Nearctaphis bakeri; Praon abjectum – Aphis solanella; Aphidius transcaspicus – Hyalopterus pruni; Adialytus salicaphis – Chaitophorus sp., Praon yomenae; Aphidius persicus – Uroleucon sonchi; and Praon rosaecola – Macrosiphum rosae.

4. DiscussionThe early evidence about Charipinae from Iran included only 5 species of Alloxysta and 1 of Phaenoglyphis (Rakhshani et al., 2001; Lotfalizadeh, 2002; Lotfalizadeh and van Veen, 2004; Rakhshani et al., 2004; Pujade-Villar et al., 2007; Khayrandish-Koshkooei et al., 2013). They were Phaenoglyphis villosa, Alloxysta citripes, Alloxysta erythrothorax, Alloxysta consobrina, Alloxysta mullensis, and Alloxysta victrix. The most recent work by Ferrer-Suay et al. (2013e) significantly increased knowledge about the Charipinae from Iran. In this work, 15 Charipinae species were determined, of which 9 species were cited for the

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first time from Iran. Many new host records were also established, in this way broadening the knowledge about the Charipinae in this area.

Alloxysta consobrina was cited in Iran as Alloxysta fuscicornis (Hartig 1841), a well-known cosmopolitan species. These 2 have been recently synonymized (Ferrer-Suay et al., 2013b). Despite our reservation in establishing this synonymy, because the name of A. fuscicornis has been used frequently in the bibliography, according to the nomenclature rules, the valid name must be A. consobrina.

In the present study, 8 species were determined within 91 host associations from different regions including A. arcuata, A. brevis, A. circumscripta, A. citripes, A. darci, A. mullensis, A. pusilla, and P. villosa; among them, Alloxysta circumscripta and A. darci are cited for the first time from Iran. Additionally, the important contribution of this work is that for the already known Charipinae species present in Iran, new data about their trophic relationships are also given here (see the comments in Section 3).

Currently, some studies aiming to analyze the specificity of the Alloxysta species are in progress (Ferrer-Suay et al., unpublished data). With this study, we are trying to elucidate possible host selection by the Charipinae based on qualitative data. Until now, A. citripes has been found only on the genus Trioxys; the host association Alloxysta citripes – Trioxys pallidus – Chromaphis juglandicola on Juglans regia seems to be very specific of this Alloxysta species. While some host records for this species seem doubtful (Dysaphis mali; Belizin, 1966), the other host aphids (Eucallipterus tiliae, Tuberculoides annulatus, Pterocallis alni, and Myzocallis coryli) have a strict affinity to Chromaphis juglandicola, so that the primary parasitoid was Trioxys pallidus.

On the basis of the studied material from all over the world, and also taking into account data from the Charipinae worldwide catalogue (Ferrer-Suay et al., 2012a), there are some complexes that repeat in different Alloxysta species, such as, for example: Ephedrus persicae – Dysaphis plantaginea in A. arcuata and A. brevis; Lysiphlebus fabarum – Aphis craccivora in A. arcuata, A. brevis, A. circumscripta, and A. victrix; Lysiphlebus fabarum – Aphis urticata, Aphis fabae in A. consobrina, A. melanogaster, and A. pleuralis; and Praon volucre – Aphis fabae in A. arcuata, A. brevis, and A. castanea. Alloxysta circumscripta was found frequently in association with Urolecucon aphids, too. Therefore, it seems that there is slight host selection, but it is more accurate for the primary parasitoids than for the aphids. It is necessary to continue collecting data about the trophic associations related with the Charipinae in order to get a better idea of the possible host selection in this subfamily.

For this reason, it is important to continue with the collections and revisions of the Charipinae material from Iran and other poorly known regions in order to improve our knowledge about their trophic relationships. In this way, we get to know the trophic relationships in which they are involved, and, with this, be able to predict the impact that their presence can have on important crops in the region.

AcknowledgmentsThis research was supported by the projects CGL2008-00180 of the Science and Innovation Ministry of Spain and the grant AP2009-4833 of the Education Ministry of Spain. The participation by E Rakhshani was supported by grant No. 89-9198, University of Zabol.

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