North-Western Journal of Zoology Vol. 4, No. 2, 2008, pp.189-202 [Online: Vol.4, 2008: 23]

Species richness of oak gall wasps (: Cynipidae) and identification of associated inquilines and parasitoids on two oak species in western Iran

Javad NAZEMI1, Ali Asghar TALEBI1,*, Seyed Ebrahim SADEGHI2, George MELIKA3 and Aurel LOZAN4

1. Dept. of Entomology, Faculty of Agriculture, Tarbiat Modares University, P.O.Box:14115-336, Tehran, Iran, 2. Division of Plant Pests and Diseases, Research Institute of Forests and Rangelands, Tehran, Iran, 3. Systematic Parasitoid Laboratory, Plant Protection and Soil Conservation Service of County Vas, Kelcz-Adelffy str. 6, Kőszeg, 9730 ; 4. Biological Centre, Institute of Entomology, Branišovská 31, 37005 Ceské Budìjovice, Czech Republic * Corresponding author: A.A. Talebi, e-mail: [email protected]

Abstract. The cynipid gall wasps (Hym., Cynipidae) are among the main associated with oaks (Quercus sp.) in Iran. The objective of this study was to infer the inquiline species and the parasitoids associated with the cynipid gall wasps while determining species richness of gall wasps in oak forests of Kurdistan, Ilam and Kermanshah provinces in the west of Iran. The cynipid gall wasps were collected from buds and leaves of Quercus infectoria Olivier and Q. brantii Lindley during 2004-2006. The collected galls were maintained under laboratory conditions at room temperature (20-25 ºC) in transparent plastic boxes until the cynipids, inquilines and their parasitoid adults were emerged. As a result of this study, 28, 22 and 4 species were collected in Kurdistan, Kermanshah and Ilam provinces, respectively. Five species of inquiline cynipids were reared from gall inducer wasps: Synergus umbraculus (Olivier), S. mikoi Melika & Pujade-Villar, S. acsi Melika & Pujade-Villar, S. bechtoldae Melika & Pujade-Villar and Saphonercus irani Melika & Pujade-Villar. Three parasitoid species include Glabobracon variator Nees, Ascogaster quadridentata Wesmael and Bassus tumidulus (Nees) (Hym., Braconidae) reared from amygdalana (Duponchel) (Lep., ) as inquiline species in oak galls related with Andricus cecconi Kieffer and A. multiplicatus Giraud. All three braconid species are reported here for the first time from Iran. The expected number of oak gall wasp species on Q. infectoria and Q. brantii was estimated to be 16 and 6, respectively. Species richness and geographical distribution of oak gall wasps, their inquilines and parasitoids are given and discussed.

Key words: Oak, Cynipidae, Braconidae, parasitoid, inquilines, species richness

Introduction are predominant; while in the eastern region, plant covering of juniper and Iran is located in the west of the deciduous or oak forests of semi-ever- Palaearctic region and its major plant green such as Q. infectoria, Q. brantii and covering is treeless steppe. Oak forms a Q. pubescens are predominant (Azizkhani very important component of the plant 2006) (Fig. 1). Western regions of Iran covering of forests in Iran. Oak forests are have a temperature range between -23.5 located as high as over 2000 meters above °C to +41.5 °C. The rainfall amount in sea level. In the North, South and West these regions is between 400 mm to 1000 of Iran, Q. pubescens Willd, Q. cerris L., Q, mm (Fattahi 1997). The oak forests extend infectoria Olivier and Q. macranthera Fisch northwards to the Caucasus Mountains

North-West J Zool, 4, 2008 Oradea, 190 Nazemi, J. et al. and westwards to the Talesh and Alborz 2006, Tavakoli et al. 2008). The Zagros Mountain range that surround the Mountains extend from the north of southern coast of the Caspian Sea. Oak Kurdistan province to the Shiraz area in forests also extend southwards along the South West of Iran. In this region oak Zagros mountain range. This area is forests are composed of drought-tolerant characterized by rich interesting fauna species including Q. libani and Q. brantii that is of most interest to many taxo- and Q. infectoria veneris (Azizkhani 2006). nomists (Melika et al. 2004, Sadeghi et al.

Figure 1. Distribution of oak species and sampling regions in Iran. b= Quercus brantii; c= Q. cerris; i= Q. infectoria; m= Q. macranteria; p= Q. pubescens. Grey colors indicate the provinces where the cynipid galls, their inquilines and parasitoids have been collected.

Some communities are simple enough and simplest measure of species richness. to permit a complete count of the number Complete counts can often be done on of species present, and this is the oldest bird communities in small habitat blocks

North-West J Zool, 4, 2008 Species richness of oak gall wasps in Iran 191 and on mammal communities in tem- Hymenenoptera (predominantly Chalci- perate and polar communities (Krebs doidea) that have become an important 1999). But it is often impossible to model system in the study of community enumerate every species in communities structure (Stone et al. 2002). of insects. One problem that frequently The tribe (Hym., Cyni- arises in comparing community samples pidae) is one of the common inquiline is that they are based on different sample groups (c.a. 175 species), whose members sizes. The larger sample exhibits the inhabit the galls of cynipid wasps (Stone greater expected number of species. One et al. 2002). The cynipid inquilines have way to overcome this problem is to lost the ability to induce galls. Cynipid standardize all samples from different inquilines larvae can develop inside the communities to a common sample size gall-inducer chamber and can usually such as the same number of individuals cause the death of the gall-inducer (Krebs 1999). Sanders (1968) proposed cynipid larvae. They can also locate and the rarefaction method for achieving this feed in the peripheral tissues of the gall goal. Rarefaction is a statistical method and can cause the death of the gall for estimating the number of species inducer larvae when the number of expected in a random sample of inquilines is high (Ronquist 1999, Csóka individuals taken from a collection. et al. 2004). Inquiline cynipids have a Rarefaction method answers this worldwide distribution, but the majority question: if the sample had consisted of are found in the Holarctic region and are n individuals (n

North-West J Zool, 4, 2008 192 Nazemi, J. et al. from the galls of B. pallida. Killing the 1 and 2). Only these two oak species were found larvae of the gall inducer has two in our study area. Weekly sampling of spring and autumn cynipid galls (Fig. 3) took place probable advantages for these moths: (1) from 22 April to 15 May and from 13 September these larvae probably represent an to 15 October in 2004-2006, respectively. Species important source of protein, supple- richness comparisons of oak gall wasps were menting the low nutritive value of the performed for 2005 and 2006 years. The optimal outer gall tissue, and (2) death of gall number of samples is determined according to inducer prevents the lignifications and Southwood & Henderson (2000): thus preserves the gall tissues as a N=(ts/Dm)2 suitable food source for the moth larvae (Ronquist 1999). Almost all the para- Where, t is students T-test of standard sitoids reared from cynipid galls are statistical tables, D is the predetermined half– specific to cynipid galls (Melika et al. width of the confidence limit for the estimation of the mean express as a decimal, m is sample 2004, Azizkhani et al. 2005). There are a mean and s is standard deviation. The optimum few exceptions indicated that some number of samples was determined to be 20 species of Chalcidoidea (such as the trees. All samples were collected by the first eupelmids, Eupelmus urozonus Dalman author (JN). The collected galls of cynipid wasps and E. vesicularis (Retzius) act as were held under laboratory conditions at room temperature (20-25 °C) in transparent plastic secondary parasitoids in the larvae of rearing boxes. The samples were observed daily (Eliason & Potter 2000, for emergence of inquiline cynipids and Schönrogge et al. 1995). parasitoid adults. All specimens were deposited The family Braconidae is the second in the collection of the Department of largest family (after the Ichneumonidae) Entomology at the Tarbiat Modares University, in the Hymenoptera, containing more Tehran, Iran. The species richness of oak gall wasps in than 15,000 valid species (Yu et al. 2005). Kurdistan, Kermanshah and Ilam provinces was Most braconid wasps are generalist estimated using the rarefaction equation that was parasitoids of Lepidoptera (Quicke & van first presented by Sanders (1968), and later Achterberg 1990). No other data is corrected independently by Hurlbert (1971) as available on the relationship between follows: ⎡ ⎛ − NN ⎞⎤ braconid parasitoids and inquilines ⎜ i ⎟ s ⎢ ⎜ ⎟⎥ ⎢ ⎝ n ⎠⎥ moth, P. amygdalana as well as on species SE n ∑ 1)( −= i=1 ⎢ ⎛ N ⎞ ⎥ richness of oak gall wasps in Iran. ⎢ ⎜ ⎟ ⎥ ⎢ ⎝ n ⎠ ⎥ Therefore, the objective of this study was ⎣ ⎦ to identify the inquilines of cynipid gall where E(Sn) is the expected species number inducing wasps and parasitoids of the in a random sample of n individuals; S is the total inquiline moth, P. amygdalana while number of species in the entire collection; Ni is determining species richness of oak gall the number of individuals in species i; N is the wasps in western Iran. total number of individuals in collection = ΣNi; n is the value of sample size (number of

individuals) chosen for standardization (n≤N); and ⎛ N ⎞ is the number of combinations of n Materials and Methods ⎜ ⎟ ⎝n ⎠

individuals that can be chosen from a set of N Oak cynipid galls were collected from individuals =N! /n! (N - n)! Quercus infectoria Olivier and Q. brantii Olivier in

Kurdistan, Kermanshah and Ilam provinces (Fig.

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Figure 2. Oak forests (Quercus infectoria) in the west of Iran.

Figure 3. Leaf galls induced by cynipid wasps on Querscus infectoria.

North-West J Zool, 4, 2008 194 Nazemi, J. et al.

Results Synergus bechtoldae Melika & Pujade-Villar, 2006 I. Inquilines Material examined: Kurdistan: Marivan, 1286-1320 m., 25.IX.2004, 2 ♂♂ As a result of this study, five in- & 2 ♀♀ ex galls of A. tomentosus on Q. quilines of cynipid gall wasps: Synergus infectoria. umbraculus (Olivier), S. mikoi Melika & Pujade-Villar, S. acsi Melika & Pujade- Saphonercus irani Melika & Pujade- Villar, S. bechtoldae Melika & Pujade- Villar, 2006 Villar and Saphonercus irani Melika & Material examined: Kurdistan: Pujade-Villar were reared from five gall Marivan, 1286-1320 m., 29.X.2005, 11 ♀♀ wasp species: Andricus tometosus, A. & 9 ♂♂ ex galls of Aphelinux persica on Q. caputmedusae, A. quercustozae, A. brantii. Ilam: 1320-1337m., 25.X.2005, 10 grossulariae and Aphelinux persica on ♀♀ & 9 ♂♂ ex galls of Aphelinux persica on Quercusn infectoria and Q. brantii. Q. brantii.

Synergus umbraculus (Olivier, 1971) Material examined: Kurdistan: II. Parasitoids of inquiline moth, Marivan, 1286-1320 m., 25.IX.2004, 4 ♀♀ Pammene amygdalana (Duponchel) ex galls of A. tomentosus on Q. infectoria; 13.X.2005; 27 ♀♀ and 21 ♂♂ ex galls of A. Three parasitoid wasp species include quercustozae on Q. infectoria; 2 ♀♀ ex galls Bracon variator Nees, Ascogaster of A. caputmedusae on Quercus infectoria. quadridentata Wesmael and Bassus Kemanshah: Javanrood, 1320-1340 m., tumidulus (Nees) (Hym., Braconidae) 25.IX.2004, 3 ♀♀ ex galls of A. were reared from the inquiline moth, P. caputmedusae on Q. infectoria; 21.IX.2004, 2 amygdalana (Lep., Tortricidae) associated ♀♀ ex galls of A. grossulariae (asexual) on with Andricus cecconi Kieffer and A. Q. infectoria; 21.IX.2004, 2 ♀♀ & 2 ♂♂ ex multiplicatus Giraud (Hym., Cynipidae). galls of A. quercustozae on Q. infectoria. All three braconid parasitoid species are newly recorded from Iran. The cynipid- Synergus mikoi Melika & Pujade- inquiline-parasitoid relationships are Villar, 2006 established for the first time in this paper. Material examined: Kurdistan: Marivan, 1286-1320 m., 3.X.2004, 9 ♀♀ & 1 Bracon variator Nees, 1812 ♂ ex galls of A. multiplicatus on Quercus Material examined: Kurdistan: brantii; 23.IX.2004, 1 ♂ ex galls of A. Marivan 1286-1320m., 27.V.2004; 8 ♂♂ ex grossulariae on Q. infectoria. galls of Andricus multiplicatus on Q. brantii. Synergus acsi Melika & Pujade- Villar, 2006 Bassus tumidulus (Nees, 1812) Material examined: Kermanshah: Material examined: Kurdistan: Javanrood, 1320-1340 m., 5.X.2005, 2 ♀♀ & Marivan, 27.V.2004, 12 ♀♀ ex galls of 2 ♂♂ ex galls of A. grossulariae (asexual) Andricus multiplicatus on Q. brantii. on Q. infectoria.

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Ascogaster quadridentata Wesmael, species) in Kermanshah province. The 1835 highest number of species collected on Material examined: Kurdistan: this oak species was determined to be Marivan, 13.V.2004, 8 ♂♂ ex galls of seven. The expected number of species in Andricus cecconi on Quercus brantii. a sample with 2,000 individuals was six Kermanshah: Gahvareh 1320-1340m., species in both two years (Fig. 4, A). 18.V.2005, 4 ♂♂ ex galls of Andricus In Kurdistan province, the species cecconi on Q. brantii. richness of oak gall wasps on Q. infectoria decreased from 2005 to 2006, from 20 species to 16 species. The highest III. Species richness of oak gall expected number of cynipid species in a wasps sample with 2000 individuals was estimated as 20 and 16 in 2005 and 2006, The present study showed that respectively. Also this study showed that species richness of gall wasps on Q. the highest richness of oak gall wasps on infectoria in Kermanshah province was Q. brantii in this region were nine and similar in both 2005 and 2006. The eight species in 2005 and 2006, res- highest number of gall wasp species pectively (Fig. 4, B). collected on Q. infectoria in this region The species richness of gall wasps in was sixteen species. The expected Ilam province on Q. brantii was similar in number of oak gall wasp species in the 2005 and 2006 (four species). The highest sample with 1000 individuals was expected number of cynipid species in a estimated to be sixteen species in both sample with 200 individuals was also 2005 and 2006. Also in this study, species estimated to be four species in 2005 and richness of oak gall wasps on Q. brantii 2006 (Fig. 4, C). was similar in two sampling years (six

18 16 A 14 Q.infectoria 2005 12 Q.infectoria 2006 10 Q.brantii 2005 8 Q.brantii 2006

wasp species 6 4

Expected number of gal of number Expected 2 0 10 50 100 150 200 300 400 500 600 700 800 1000 Number of individuals

Figure 4./A. Comparison of species richness of oak gall wasps in Kermanshah province in 2005 and 2006.

North-West J Zool, 4, 2008 196 Nazemi, J. et al.

22 20 B 18 16 14 12 10 8 wasp specieswasp 6 Q.infectoria 2005 Q.infectoria 2006 4 Q.brantii 2005 Expected number of gal 2 Q. brantii 2006 0 10 100 200 400 600 800 1200 1600 2000

Number of individuals

4.5 p 4 C 3.5 3 2.5 2 species 1.5 Q.brantii 2005 1 Q.brantii 2006 0.5 Expected number of gall of was number Expected 0 10 50 100 150 200 300

Number of individuals

Figure 4./B.-C. Comparison of species richness of oak gall wasps in Kurdistan (B) and Ilam (C) provinces in 2005 and 2006.

Lists of oak gall wasps and their host wasps was related with Q. infectoria. plant associations as well as location of Sixteen and four gall wasp species were galls on oak trees, number of chambers in collected on shoots and leaves of Q. the galls, sexual or asexual generation infectoria, respectively (Table 1). There and time of gall formation are shown in were both oak species (Q. infectoria and Table 1 and Table 2. Our finding showed Q. brantii) in Kurdistan and Kermanshah that only asexual generation of cynipid provinces, whereas only Q. brantii was

North-West J Zool, 4, 2008 Species richness of oak gall wasps in Iran 197 found in Ilam province (Table 2). We grossulariae respectively on Q. infectoria collected four and three gall wasp species and Q. brantii. Except for the sexual on catkins and leaves of Q. brantii, generation of N. saliens reared on Q. respectively. One species of gall wasp brantii in Ilam province, no sexual was also collected on shoots, acorns and generation on this oak was recovered in branches of this oak species (Table 2). Ilam province. All species formed galls Here all sexual generations of oak gall on catkins collected on Q. brantii in the wasps are formed on Q. brantii (Table 2). provinces of Kurdistan and Kermanshah There was host alternation concerning (Table 2). the asexual and sexual generations of A.

Table 1. Gall wasp species associated with Quercus infectoria in Kurdistan and Kermanshah provinces in 2004-2006.

Gall wasp species Location of Number of Sexual or Season of Location of gall larval chamber asexual gall collection formation generation formation Andricus quercus leaf unichamber asexual summer Kurdistan & (Fourcroy, 1785) Kermanshah Cynips quercusfolii leaf unichamber asexual summer Kurdistan & (Linnaeus, 1758) Kermanshah A. caputmedusae (Giraud, shoot unichamber asexual summer Kurdistan & 1859) and Kermanshah autumn A. coriarious (Hartig. 1838) shoot unichamber asexual summer Kurdistan & Kermanshah A. moreae Melika, shoot unichamber asexual summer Kurdistan & Atkinson, Sadeghi, Kermanshah Tavakoli & Stone, 2003 A. sternlichti Bellido, shoot unichamber asexual summer Kurdistan & Pujade-Villar & Melika, Kermanshah 2002 A. lucidus (Harting, 1843) shoot multichamber asexual summer Kurdistan & and Kermanshah autumn A. curtisii (Muller, 1970) shoot unichamber asexual summer Kurdistan & Kermanshah A. grossulariae Giraud, shoot multichamber asexual summer Kurdistan & 1859 Kermanshah A. pseudoaries Melika, shoot unichamber asexual summer Kurdistan Stone & Sadeghi, 2004 A. megalocidus Melika, shoot ultichamber asexual summer Kurdistan & Stone, Sadeghi & Pujade- and Kermanshah Villar, 2003 autumn A. polycerus (Giraud, 1859) shoot unichamber asexual summer kurdistan A.tomentosus (Trotter, shoot unichamber asexual summer Kurdistan & 1901) Kermanshah

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Table 1. (Continued)

Gall wasp species Location Number of Sexual or Season of Location of of gall larval chamber asexual gall collection formation generation formation A. insana (Bosc, 1792) shoot unichamber asexual summer Kurdistan & and Kermanshah autumn A. stonei Melika, Tavakoli & shoot unichamber asexual summer Kurdistan Sadeghi, 2005 A. trunicicolous (Giraud, shoot unichamber asexual summer Kurdistan & 1859) Kermanshah A. askewi Melika & Stone, shoot unichamber asexual summer Kurdistan & 2001 Kermanshah Synophrus politus Hartig, shoot unichamber asexual summer Kurdistan 1843 Neuroterus quercusbaccarus leaf unichamber asexual summer Kurdistan & (Linnaeus, 1758) Kermanshah N. numismalis (Fourcroy, leaf unichamber asexual summer Kurdistan & 1785) Kermanshah

Table 2. Gall wasp species associated with Quercus brantii in Kurdistan, Kermanshah and Ilam provinces in 2004-2006.

Gall wasp species Location of Number of Sexual or Season of Location of gall larval chamber asexual gall collection formation generation formation Andricus grossulariae catkin unichamber sexual spring Kurdistan & (Giraud, 1859) Kermanshah A. multiplicatus Giraud, catkin unichamber sexual spring Kurdistan & 1859 Kermanshah A. cecconii Kieffer, 1901 catkin unichamber sexual spring Kurdistan & Kermanshah Aphelonyx persica Melika, shoot unichamber asexual summer Kurdistan, Stone, Sadeghi et Pujade- Kermanshah Villar, 2004 & Ilam Neuroterus lanuginosus leaf unichamber asexual summer Kurdistan, (Giraud, 1859) Kermanshah & Ilam N. saliens (Kollar, 1857) acorn multichamber sexual spring Kurdistan, Kermanshah & Ilam N. saliens (Kollar, 1857) leaf unichamber asexual summer Ilam & Kermanshah N. macropterus (Hartig, branch unichamber asexual summer Kurdistan & 1843) Ilam Chilaspis israeli (Sternlicht, catkin unichamber sexual spring Kurdistan 1968) mikoi leaf unichamber sexual spring Kurdistan Tavakoli, Stone & Azizkhani, 2005

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Discussion Tortricidae). Meanwhile, this parasitoid has also been reared from Cydia splendana We reared five species of the (Hübner) and C. fagiglandana (Zeller) Synergus from the cynipid galls in the (Lep., Tortricidae) in (Rotundo & west of Iran. These five inquiline species Giacometti 1986). In New Zealand, larvae have been recently described from of Cydia pomonella were parasitized by A. cynipid gall inducers in Iran (Sadeghi et quadridentata (Tomkins et al. 1987) as well al. 2006). Populations of Callirhytis as in Moldavia (Babidorich & Sharov cornigera (Osten Sacken) (Hym., 1986) and Sweden (Subinprasert 1987). In Cynipidae) and its associated parasitoids Russia, A. quadridentata has been and inquilines were monitored on pin considered as the most effective bio- oak, Quercus palustris in USA from 1997 control agent of Cydia splendana (Hübner) to 1999. The most abundant inquilines in (Terent'ev 1978). Bogenschutz & Franz cynipid gall inducers were the species of (1975) reported A. quadridentata as a the genus Synergus (Eliason & Potter larval parasitoid of Cydia pactolana 2000). Nieves-Aldrey & Pujade-Villar (Zeller) in . Kisakurek (1972) (1986) recorded eleven species of the found that A. quadridentata had a high genus Synergus from oak gall wasps in level parasitism on larvae of Lobesia Spain. Baldassari and Baronio (1996) botrana (Denis & Schiffermüller) (Lep., reared Synergus rotundiventris as a new Tortricidae) on grape vines in Turkey. Yu record from Callirhytis glandulosa in Italy. et al. (2005) reported that this parasitoid Our findings are in agreement with is quite polyphagous but mostly previous data demonstrating that the restricted to Tortricidae. species of Synergus are the most Bassus tumidulus (Nees), a parasitoid important inquilines of oak gall wasps. of Gypsonoma aceriana (Duponchel), was They cannot induce galls and their larvae collected during the period of 1989-2000 live inside the galls formed by other in (Georgiev & Delkov 2003). cynipids (Ronquist 1999, Csóka et al. Bassus linguarius (Nees) and B. tumidulus 2004). were recorded as parasitoids of Cydia There is no other data on parasitism pomonella (Cetin & Beyarslan 2001). of the inquiline moth, Pammene amygda- Nuzzaci and Triggiani (1982) found lana by B. variator, Bassus tumidulus and Lobesia botrana (Denis & Schiffermüller) A. quadridentata. However, all three bra- (Lep., Tortricidae) to be parasitized by B. conids have previously been reported linguarius in Italy. Yu et al. (2005) listed from many species of Lepidoptera and about 20 hosts for this parasitoid all over Coleoptera. All known parasitoids that the world. attack lepidopterous inquilines are Glabrobracon variator was reported as wasps, including members of Ichneu- an effective agent in control of Pissodes monidae and Braconidae (Quicke & van validirostris (Sahlberg) (Col., Curcu- Achterberg 1990). lionidae) in the Bryansk region of the Rotundo & Tremblay (1993) already Soviet Union in 1966-68 (Smetanin 1970). quoted A. quadridentata as a parasitoid of Yu et al. (2005) listed 69 hosts for this Pammene fasciana (Linnaeus) (Lep., parasitoid, which include Chrysome-

North-West J Zool, 4, 2008 200 Nazemi, J. et al. lidae, Curculionidae (Coleoptera), Ten- Host plant traits (e.g. morphology, thredinidae (Hymenoptera), Tephritidae phenology, plant vigor) are known to (Diptera) and several families of influence host choice by gall-inducing Lepidoptera including Tortricidae. P. insects (Anderson, et al. 1989, Eliason & amygdalana is reported here as new host Potter 2000). Our findings have shown record for this parasitoid. that gall wasp inducers also discriminate Little data are available on species among closely related species of Q. brantii richness of oak gall wasps on Q. infectoria and Q. infectoria. and Q. brantii in Iran and other countries. In our research all sexual generations of gall wasps were collected from Q. brantii Acknowledgments. This research was financially in Kurdistan and Kermanshah as these supported by the Department of Entomology, Tarbiat were the only oak species in these Modares University, Tehran, Iran. The authors are most grateful to the two reviewers, Prof. G. Delvare and Dr. provinces. Sexual generation of N. saliens H. Lotfalizadeh for their valuable comments and was only collected on Q. brantii in Ilam suggestions on the manuscript. province. Azizkhani (2006) reported 24 gall wasp species on Q. infectoria (13 References species) and Q. brantii (11 species) in Lorestan province. All sexual generations Abrahamson, W. G., Hunter, M. D., Melika, G., Price, P. W. (2003): Cynipid gall-wasp communities correlate on catkins was collected on Q. brantii. He with oak chemistry. Journal of Chemical Ecology, also found asexual forms of oak gall 29(1): 209-23. wasps only on Q. infectoria. Shojai (1980) Aguilar, J. M., Boecklen, W. J. (1992): Patterns of herbivory in the Quercus grisea X Quercus gambelii reported 36 gall wasps species related species complex. Oikos 64: 498–504. with Q. infectoria from Iran. He proved Anderson, S. S., McCrea, K. D., Abrahamson, W. G., that the most of the asexual generations Hartzel, L. M. (1989): Host genotype choice by the ball gallmaker Eurosta solidaginis (Diptera: were formed on Q. infectoria. Oak gall Tephritidae). Ecology 70: 1048–1054. wasps exhibit a high degree of host plant Azizkhani, E., (2006): Parasitoid fauna of oak gall wasps specificity and physiology and chemistry in Lorestan province. PhD thesis, Faculty of Agriculture, Tehran University, 277pp. may be crucial for the success of these Azizkhani, E., Rasulian, G. R., Kharazi-Pakdel, A., insects (Shorthouse 1982, How et al. Sadeghi, S. E., Tavakoli, M., Melika, G. (2005): Report of eight species of parasitoid wasps 1993). A survey on oak hybrids has belonging to Chalcidoidea from cynipid galls on demonstrated that gall inducers can oak trees. Journal of Entomological Society of Iran distinguish closely related host plant 25(1): 79-80. Babidorich, M. M., Sharov, A. A. (1986): Coherence of species (Aguilar & Boecklen 1992). seasonal cycles of the codling moth Laspeyresia Among gall-inducing insects, cynipid pomonella L. (Lepidoptera, Tortricidae) and its wasps are noted for their fidelity to their parasites in Moldavia. Entomologicheskoe Obozrenie 65(3): 470-478. host plants. Approximately 87% of all Baldassari, N., Baronio, P. (1996): Infestation of cynipids attack species of Quercus sp. and Callirhytis rufescens (Mayr) (Hymenoptera the host range of each wasp species is Cynipidae) on Quercus pubescens Willd. and Q. robur L. growing on reclaimed land at Maccarese (Roma). generally restricted to one or a few oak Informatore Fitopatologico 46(1): 19-22. species (Burks 1997). Plant chemistry Becerra, J. X. (1997): Insects on plants: may play an appreciable role in the macroevolutionary chemical trends in host use. Science 276: 253–256. selection of plant species by gall wasps Bogenschutz, H., Franz, J. M. (1975): Parasitenstudien als (Becerra 1997, Abrahamson et al. 2003). Entscheidungshilfen bei der integrierten

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