Wing Dimorphism of European Mole Cricket Gryllotalpa Gryllotalpa (L.) (Orthoptera: Gryllotalpidae) in the North-West of Iran
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NORTH-WESTERN JOURNAL OF ZOOLOGY 9 (1): 45-50 ©NwjZ, Oradea, Romania, 2013 Article No.: 131102 http://biozoojournals.3x.ro/nwjz/index.html Wing dimorphism of European mole cricket Gryllotalpa gryllotalpa (L.) (Orthoptera: Gryllotalpidae) in the north-west of Iran Mohammad Hossein KAZEMI1*, Shabnam JAFARI1, Hosseinali LOTFALIZADEH2 and Mohammad JAFARLOO2 1. Department of Plant Protection, Tabriz branch, Islamic Azad University, Tabriz, Iran. 2. Department of Plant Protection, East-Azarbaijan Research Center for Agriculture & Natural Resources, Tabriz, Iran. *Corresponding author, M.H. Kazemi, E-mail: [email protected] Received: 24. December 2011 / Accepted: 25. September 2012 / Available online: 27. December 2012 / Printed: June 2013 Abstract. The seasonal study of wing dimorphism in the European mole cricket, Gryllotalpa gryllotalpa (Linnaeus, 1758) (Orthoptera: Gryllotalpidae), was carried out in northwest Iran. Based on present research, the long-winged (LW) morph appeared in early-mid spring when vegetation density is low and the crickets need to fly to search for food. Study of foretibiae showed that the dactyls are elongated and longer than short- winged (SW) morph’s dactyls, and that they are suitable for excavating hard soil in this period. SW adults were observed from May to July and the dactyls are shorter and broader, which are suitable for more tunneling in light soils in the middle of spring. Key words: European mole cricket, wing dimorphism, long-winged, short-winged, Iran, first report. Introduction pterous), or totally absent wings (apterous). In addition to differences in wing length, morphs are The European mole cricket, Gryllotalpa gryllotalpa often observed with differences in some other (Linnaeus, 1758), is one of the most important in- morphological, physiological and biological cha- sect pests in turf and field crops of Iran. They are racteristics such as degree of flight muscle deve- burrowing insects and feed on a variety of organ- lopment, duration of nymphal stage, time to first isms in the soil. These insects do not attack plants reproduction, fertility and diapause (Vepsäläinen directly, but by tunneling, extended surface tun- 1978, Harrison 1980). nels, cause significant damage to grass and crops Developing and maintaining the flight appara- of gardens, as they chop off any roots encountered tus carries a cost. For example, macropterous fe- when digging (Weiss & Dickerson 1918, Kazemi et males of the sand cricket, Gryllus firmus (Scudder), al. 2010). have an older reproductive age with shorter life- It was introduced from Europe into the United time fecundities when compared to their short- States about 1913. It has now been reported from winged counterparts (Elizabeth et al. 2011). Impor- Massachusetts, New Jersey, New York, and Penn- tantly, flight capability trades-off occur with re- sylvania (Nickle & Castner 1984). Weiss and productive effort: short-winged females start ovar- Dickerson (1918) determined that the original in- ian growth at an earlier age and often exhibit troduction of this species was in nursery stock greater overall fecundity than their long-winged from Holland and Belgium. It also is widespread counterparts. Long-winged females mainly allo- in the northern European part of Russia, southern cated energy from food to flight muscle develop- Ukraine and the northern Caucasus (Anonymous ment and general maintenance of the body rather 2010). This species in ecological conditions of than to egg production, whereas short-winged fe- north-west of Iran molts eight times during its de- males used it for egg production and longevity velopment. Small nymphs have no wings, but fifth (Zhao et al. 2010, Guerra 2011). The polymorphism instars have small wing pads that will grow in may be determined by genetic differences between later molting. There is only one generation annu- morphs (genetic polymorphism), environmental ally (Kazemi et al. 2010). conditions under which the morphs develop Wing polymorphism is commonly observed in (environmental polyphenism) or by a combination many orders of insects, especially species of of these two (Zera et al. 2007), however, wing Orthoptera, Coleoptera and Hemiptera (Harrison dimorphisms in insects are controlled by a single 1980). The polymorphism consists of discrete locus and polygenes, respectively (Roff & differences in wing length with morphs exhibiting Fairbairn 2007). fully developed (macropterous), reduced (brachy- The relationship of development and repro- 46 M.H. Kazemi et al. duction of flight capable (long-winged) and hind wings never reach to end of abdomen) individuals flightless (short-winged or wingless) morphs of could not fly in the air; thus, soil sampling is the main col- wing polymorphism with endocrine hormones has lection method. Mole crickets that surfaced after flushing the soil and with light traps were collected and counted been investigated (Zera 2003). There is a clear cor- on each sampling day. relation between activity of juvenile hormone (JH) during the last nymphal stadium and subsequent Morphological studies molt to either the long-winged (LW) or short- Morphological data were obtained from measurement of winged (SW) adult (Fairbairn 1994, Fairbairn & pronotum width, length of body, forewings, hind wings, Yadlowski 1997, Roff et al. 1997). Thus far, the pronotum and dactyls of foretibia using cullies (±0.01) on only direct test of the JH wing morph hypothesis both sexes. Two hundreds individuals were examined for these characters. According to Walker and Sivinski (1986), has been undertaken on the cricket, Gryllus rubens the ratios of hind wing (HW) to forewing (FW) length are Scudder (Zera & Denno 1997). Experimental eleva- used to discriminate among wing morphs. Data analyz- tion of JH titer during the last stadium in long- ing was done by t-test, and comparison of adult wing ra- winged-destined G. rubens redirected their devel- tio (HW/FW) in each sex was conducted. opment to the short-winged morph. Endo (2006) reported seasonal wing dimorphism of oriental mole cricket, Gryllotalpa orientalis (Brumeister, Results 1839), in Japan. It was the only report of wing di- morphism in mole crickets. In his study, the short- In ecological conditions Iran, the LW morph ap- winged morph appeared in early spring, and the peared in early-mid of spring when vegetation overwintering nymphs became long-winged density is low and the mole crickets need to fly to adults in summer. search for food and for mates. Habitat stability Considering economical importance of Euro- and quality also affect the fitness of each morph pean mole cricket in Iran, biological and ecological (Roff 1986, Denno et al. 1996). It is reasonable that study of this species is badly needed for making juveniles become SW adults after May (the habitat decision in integrated control of this pest. has good quality in this time of year), with more reproduction that has benefit to brachypterous in- dividuals. Materials and methods As the soil temperature rises in mid-March and April, mole crickets do more tunneling. The Study areas LW adult males in warm evening of late April or The study was done in turf grounds and field crops of May excavate an acoustic (funnel-shaped) cham- Tabriz (N 38º 6', E 46º 26'), Azarshahr (N 37º, E 45º) and ber with an opening to the soil surface for making Miandoab (N 36º, E 46º) in northwest Iran based on ob- servation of the mole crickets’ damage or presence of calling song. signs of tunneling. Mole crickets were sampled four times Females are attracted to the calling aggrega- monthly from mid-March to November in 2009 and 2010. tion, fly over the sound chamber created by the displaying males and then drop to the ground to Collecting access the selected male’s burrow (Hertl et al. Collection was made by two methods: 2001, Howard & Hill 2006, Kazemi et al. 2011). 1) The first method was to collect flying adults with Shortly after mating, oviposition starts. Small ultraviolet fluorescent light traps. The light traps were set for about four hours after sunset. Three light traps were nymphs continue to feed and grow through the set beside the selected areas and trapped mole crickets summer, and they are most destructive in this area were collected the next day. This method is suitable for during late July to August. Brandenburg & Wil- long-winged (LW with hind wings are longer than the liams (2002) believe that, for similar species, the abdomen) individuals. males die after mating and the females die shortly 2) The second method according to Nikouei et al. after oviposition. (2006) was to directly collect adults and nymphs from soil. Mole cricket damage was evaluated using a 0.6 m² frame divided into nine equal, square-shaped sections Morphological studies and damage was rated from zero to nine. The frame was Sex determination placed on the ground and the number of sections which contained mounds or tunnels was counted. Then, 0.6 m² Adults of both morphs have complete wings, and areas were flooded with 8 liters of soapy water. Mole the sexes can be distinguished by forewing vena- crickets that surfaced within the frame after flushing were tion. Forewings of the males have a pair of large collected and counted because short-winged (SW with triangular cells that have been described as harp- Wing dimorphism of Gryllotalpa gryllotalpa in Iran 47 Table 1. Measurment (in mm) of body parts in male European mole cricket (n=100). Long-winged Short-winged Body parts Mean ± SE Min Max Mean ± SE Min Max Length of body 42.7 ± 1.33 35 50 43.00 ± 0.95 38 48 Forewing length 15.7 ± 0.37 14 17 13.50 ± 0.43 12 16 Hind wing length 36.1 ± 0.77* 34 41 22.40 ± 0.73* 19 26 Pronotum length 14.5 ± 0.17 14 15 14.57 ± 0.18 14 15 Pronotum width 11.5 ± 0.17 11 12 11.57 ± 0.18 11 12 * significant difference at α = 5% Table 2.