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A Biological Study of Aspidiotus Nerii Bouché Conducted to Improve The 1 Assessment of life history parameters of Aspidiotus nerii (Hemiptera: Diaspididae) to 2 improve the mass rearing of Aphytis melinus (Hymenoptera: Aphelinidae) 3 4 5 Author names: J.E. González-Zamora, M.L. Castillo, C. Avilla 6 7 e-mail: [email protected] 8 9 Published version: 10 11 http://dx.doi.org/10.1080/09583157.2012.691157 12 -1- 13 Abstract 14 The biological control of Aonidiella aurantii (Maskell) on citrus can be achieved 15 with periodic releases of the parasitoid Aphytis melinus DeBach. A. melinus is 16 normally reared on parthenogenetic strains of the scale Aspidiotus nerii Bouché. 17 The development, crawler production and survival of A. nerii were studied at two 18 temperatures (25±1 and 30±1 ºC) and two relative humidities (normal 55±5 %, 19 and high 85±5 %) on squashes. The temperature had an important effect on 20 development, but not the relative humidity. At 30 ºC, no development was 21 observed, and therefore, no crawlers were produced. At 25 ºC, the development of 22 the L1, L2, and preoviposition stages required 24.4±0.7, 11.1±0.8, and 13.2±0.3 23 days, respectively. The total times until the appearance of gravid females, the 24 initiation of crawler production and the peak of crawler production were 48.7±0.1, 25 50.1±0.7, and 63.3±1.0 days, respectively. Crawler production lasted 42.6±1.9 26 days, and the 50 % production level was attained on the 13th day. Initial crawler 27 survival was higher at 30 ºC (65.9±2.8 %), whereas at the end of development, 28 survival was much higher at 25 ºC (88.0±2.1%). The relative humidity showed no 29 effect on initial and final survival. Progeny production was similar at the two 30 relative humidities, with an average of 28.0±2.8 crawlers per female at 25 ºC. The 31 relation between the number of crawlers and weight was Number of crawlers = 32 1,031.37 + 835,500.0*weight of crawlers (g), with R2=0.826. The importance of 33 selecting the correct strain of A. nerii is emphasised. 34 Keywords: Aspidiotus nerii; parthenogenetic strain; development; crawler 35 survival; biology. 36 -2- 37 1. Introduction 38 The importance of biological control has increased worldwide for many crops and pests due, 39 among different reasons, to the increasing restrictions on the use and registration of 40 pesticides, e.g., in the European Union with the Council Directive 91/414/EEC (European 41 Union 2011). A clear example of the expansion of this methodology is the extended use in 42 recent years of biological control techniques in commercial greenhouses in southeastern Spain 43 to control pests. The use of these techniques has increased from 1.3 % of the surface in 44 2006/2007 to 44.1 % in 2008/2009. In certain crops, such as sweet pepper or melon, 70 to 90 45 % of the surface is cultivated using integrated pest management criteria (Beltrán, Parra, 46 Roldán, Soler and Vila 2010; Blom, Robledo, Torres and Sánchez 2010). 47 Citrus is a good example for studying the importance of biological control. Since the 48 end of the 19th century, biological control has been implemented against several pests in 49 citrus (Kennet, McMurtry and Beardsley 1999), originally in the USA and later in many other 50 countries. Perhaps one of the most important pests in the citrus industry is Aonidiella aurantii 51 (Maskell) (Hemiptera: Diaspididae) or Californian Red Scale (CRS) (Jacas, Karamaouna, 52 Vercher and Zappalà 2010). Many parasitoids and predators have been considered for use 53 against this pest, but currently, its biological control relies mainly on hymenopteran 54 parasitoids, such as Aphytis africanus Quednau, Aphytis coheni DeBach, Aphytis lingnanensis 55 DeBach, Aphytis melinus DeBach, Comperiella bifasciata Howard, and Encarsia perniciosi 56 (Tower) (Kennet, McMurtry and Beardsley 1999), depending on the citrus region. With an 57 integrated pest management strategy and, of course, with organic management, biological 58 control is the basis of the control of CRS. Such control programmes may also involve the use 59 of several insecticides against this pest or other pests (García-Marí 2003; Grafton-Cardwell 60 2006; Jacas, Karamaouna, Vercher and Zappalà 2010), although mating disruption arises as a 61 promising new complementary strategy (Vacas, Alfaro, Navarro-Llopis and Primo 2009, -3- 62 2010). Conservation of natural enemies is the most rational approach (Jacas y Urbaneja 63 2010), but an augmentation strategy is used to help control CRS populations and is commonly 64 applied when the size of the parasitoid population is small at the beginning of the first CRS 65 generation (Moreno and Luck 1992; Sorribas and García-Marí 2010; University of California 66 2011). This strategy, based on the mass rearing of A. melinus (one of the most frequently used 67 species of parasitoid), is widely implemented in many citrus regions (Mazih, 2008; Zappalà, 68 Campolo, Saraceno, Grande, Raciti, Siscaro and Palmeri 2008; Zappalá 2010; Olivas, Lucas, 69 Calvo and Belda 2011; University of California, 2011), with subsequent releases of the 70 parasitoid in doses varying between approximately 25,000 and 250,000 individuals per ha and 71 year, depending on the climatic conditions, the densities of CRS and the parasitoids present. 72 Biological control by augmentation requires the regular production of natural enemies 73 by insectaries that seek to maximise production at the lowest cost and with a standard quality 74 (Hoy 2000; Van Lenteren 2003; Warner and Getz 2008). As demand increases under new 75 pest-control scenarios, commercial insectaries must meet these requirements. The production 76 of A. melinus is an appropriate example. The species has been produced in insectaries since 77 the 1950s using a well-known and widespread technology (DeBach and White 1960; Rose 78 1990) based on the use of a parthenogenetic strain of Aspidiotus nerii Bouché (Hemiptera: 79 Diaspididae) as the host and different species of squash as the feeding substrate for the host. 80 The parthenogenetic strain of A. nerii was described first in 1901 and subsequently by 81 several authors for different plants and regions of the world (a brief review of the topic 82 appears in Gerson and Hazan 1979; Provencher, Morse, Weeks and Normark 2005). 83 Parthenogenesis is caused by bacteria of the genus Cardinium (Provencher, Morse, Weeks 84 and Normark 2005; Gruwell, Wu and Normark 2009). The parthenogenetic strain is widely 85 used in laboratories and insectaries for the mass rearing of different parasitoids and predators 86 (Uygun and Elekçioglu 1998; Raciti, Saraceno and Siscaro 2003; Silva, Guerreiro, Michelotto -4- 87 and Busoli 2003). However, although the demand for natural enemies reared on this host is 88 increasing (as in the case of A. melinus), few recent studies have focused on the influence of 89 various factors on its biology and progeny production (DeBach and Fisher 1956; Gerson and 90 Hazan 1979; Rocha, Silva, Michelotto and Busoli 2006). This information is important for 91 improving the efficiency of the production of parasitoids and predators. 92 For these reasons, the biology of a parthenogenetic strain of A. nerii under different 93 temperatures and relative humidities and the effect of these factors on crawler survival and 94 development on squashes are the primary emphases of this work. Similarly, crawler 95 production and its pattern over time were studied to support the scheduling of production. 96 Finally, a relationship between the weight and the number of crawlers is presented. This 97 relationship is of great interest in studies of mass production. 98 99 2. Material and Methods 100 2.1 Insect origin and mass rearing 101 The uniparental (parthenogenetic) strain of A. nerii was originally supplied by the INRA 102 laboratory of Vallbone (France) in January 2009. The original insects were received on 103 potatoes, and the crawlers produced were transferred to squashes (Cucurbita moschata Duch 104 ex Lam.). Thereafter, the colony has been continuously reared on squashes in our facilities at 105 25±1 ºC and 55-65 % RH, following the original methodology of DeBach and White (1960) 106 and Rose (1990) with minor adjustments to our local conditions, as is also the case in other 107 insectaries (Raciti, Saraceno and Siscaro 2003). 108 The methodology involves a procedure in which scale-infested squashes are placed on 109 steel shelves. Below, on other shelves, are new, clean squashes, ready to be colonised. 110 Attracted by light, the mobile first-stage nymphs (crawlers) walk towards the tip of the 111 mother squash to search for a place to attach and feed. They fall to the squashes below and -5- 112 proceed to colonise. We used a variation of the previous methodology (as in Raciti, Saraceno 113 and Siscaro 2003). No squashes were placed on the shelf below. Instead, a sheet of paper was 114 placed on the shelf. The crawlers were collected every day and then distributed onto new 115 squashes using a dispenser resembling a salt shaker. This technique produced a regular 116 distribution of the scale insects on the surface of the newly infested squashes. 117 118 2.2. Survival and development studies 119 Squashes of approximately 15-20 cm in length and 8-12 cm in diameter were selected for the 120 development and fertility studies. The squashes were washed with a sponge soaked in soapy 121 water. They were then rinsed with tap water, thoroughly dried with a paper towel and placed 122 at 25 ºC to reach room temperature prior to infestation with the crawlers. On the day of 123 infestation, the crawlers were collected from the rearing facility and distributed over half of 124 the squash surface.
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