مجلة ) - (Hebron University Research Journal-A (Natural Sciences جامعة الخليل للبحوث- أ (العلوم الطبيعيه

Volume 2 Issue 2 Article 1

2005

Effect of Different Prey Species on the Biology of the Predatory Bug, caliginosus Wagner [: ]

عبد الجليل حمدان [email protected] ,جامعة الخليل

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Recommended Citation ,Effect of Different Arthropod Prey Species on the Biology of the Predatory Bug" (حمدان, عبد الجليل (2005 Wagner [Hemiptera: Miridae]," Hebron University Research Journal-A (Natural .Vol. 2 : Iss. 2 , Article 1 :مجلة جامعة الخليل للبحوث- أ (العلوم الطبيعيه) - (Sciences Available at: https://digitalcommons.aaru.edu.jo/hujr_a/vol2/iss2/1

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Hebron University Research Journal. H.U.R.J. is available online at Vol.(2), No.(2), pp.(48 – 64 ) 2006 http://www.hebron.edu/journal

Effect of Different Arthropod Prey Species on the Biology of the Predatory Bug, Macrolophus caliginosus Wagner [Hemiptera: Miridae]

*Abdul-Jalil Salem Hamdan

Department of Plant Production and Protection, Faculty of Agriculture, Hebron University, P.o.box (40) , Hebron - Palestine

Abstract:

Laboratory experiment were done to examine the effect of different arthropod prey on the biology of predatory bug, Macrolophus caliginosus Wagner at 25 oC, 75% r.h. and 16:8 L:D photoperiod. The food eaten by M. caliginosus clearly had a major impact on most parameters of its life cycle. When either greenhouse , Trialeurodes vaporariorum (Westwood); eggs of Ephestia kuehniella Zeller, or potato peach , Myzus persicae Sulzer were offered as preys, during development M. caliginosus had high survival percentage (80%, 79% and 64% respectively) and developed more quickly with duration of development of (17.4, 15.6 and 15.7 days respectively) and were able to produce new generations. However, when greenhouse red spider mites, Metatetranycc chus urticae Koch or broad bean aphid, Aphis fabae Scopoli were offered as preys, the predator had high mortality (67% and 50% respectively), longer development, and shorter adult longevity and produced no offspring. The results also showed that M. caliginosus could obtain some nutrients from plants, as 10% of M. caliginosus nymphs survived to the fourth instar when reared on clean leaves of Pelargonium peltatium, although they did not survive to the adult stage.

Key words: Macrolophus caliginosus, Biological Control, & Predator Preys Relationship.

* Corresponding author: [email protected]

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تأثري نوع الفريسة من مفصليات األرجل على بيولوجيا حشرة البق املفرتس Macrolophus caliginosus [Hemiptera: Miridae]

امللخص:

لدرا�سة ت أ�ثري نوع الفري�سة )من مف�صليات أالرجل( على بيولوجيا البق املفرت�س Macrolophus caliginosus مت تنفيذ جتارب خمربية حتت ظروف حمكمة ) 25oC ، رطوبة ن�سبية %75، فرتة �إ�ضاءة L16 : D8(. أ�ظهرت نتائج الدرا�سة أ�ن لنوع الفري�سة ت أ�ثري وا�ضح على معظم اخلوا�ص البيولوجية للبق املفرت�س. فعند تغذية البق املفرت�س على ذبابة البيوت البال�ستيكية البي�ضاء، Trialeurodes vaporariorum Westwood أ�و من الدراق أالخ�رض Myzus persicae Sulzer أ�و بي�ض عثة الطحني Ephestia kuehniella Zeller لوحظ أ�ن �رسعة تطور البق كانت عالية حيث أن� فرتة التطور كانت 15.7، 17.4، ً يوماعلى التوايل و أ�ن ن�سبة عالية من البق و�صلت �إىل مرحلة الن�ضج و أ�جنبت ًجيال ً. جديدا يف املقابل، لوحظ أنه �عند تغذية البق املفرت�س على حلم البيوت البال�ستيكية أالحمر Metatetranychus urticae Koch أ�و على من الفول Aphis fabae Scopoli ف�إن ن�سبة عالية من ح�رشات البق نفقت قبل و�صولها ملرحلة الن�ضج، وان ن�سبة الوفيات اثناء التطور كانت %67، %50 على التوايل، يف حني كانت �رسعة تطور ح�رشات البق بطيئة، أ�ما ح�رشات البق التي متكنت من الو�صول ملرحلة الن�ضج فلم تنجب ًجيال ًجديدا. إبال�ضافة �إىل ذلك، أ�ظهرت نتائج الدرا�سة انه عند تربية ح�رشات البق على أ�وراق جريانيوم Pelargonium peltatiumغري م�صابة فقد متكن %10 فقط من ح�رشات البق من التغذية على النباتات وو�صلت يف تطورها ملرحلة احلورية الرابعة ومل يتمكن أ� ُي منها من الو�صول ملرحلة الن�ضج.

Introduction: Almost all predators are carnn davets, 1995). This polyphagy allows nivorous with a varied diet them to feed on many arthropods such including , mites and some can as mites, , thrips or noctuid eggs, occasionally feed on vegetative matter. and thus survive when whitefly densitn Hemipteras are the main predators of ties are low. the (GHWF) and Macrolophus caliginosus Wagner is a have been studied mainly in Japan and polyphagous predator, which has been Europe, where the winters are too cold recorded feeding on whiteflies (T. vapocr for its free existence outdoors (Gerling, rariorum); aphids (M. persicae); thrips 1990). (different species) and mites (Metact Within the Hemiptera, several species tetranychus urticae) (Eremenko, 1984). feed on prey belonging to more than one In addition, Klapwijk (1995) reported arthropod family (Landis et al., 1958; M. caliginosus feeding on lepidopteran Ekbom, 1981; Kajita, 1982, 1984; Gerln eggs and caterpillars, leaf-miner larvae ling, 1990; Riudavets et al., 1993; Riudn and thrips and, when large numbers of

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the bug were present in the crop, the aphid (PPA) different ages, on tobacco bug helped to control those pests. leaf-discs. Moreover, Malausa et al. (1987) and 5. Aphis fabae Scopoli, Black bean Foglar et al. (1990) concluded that M. aphid (BBA) different ages, on leaves caliginosus appears to be a particularnl of broad bean. ly good polyphagous predator which 6. Control treatment with no prey but could be effectively used for biological clean leaves of Pelargonium peltatum control programmes. provided. A minimum of 20 replications of each Objective: treatment were initially used, each comprising one newly hatched nymph The objective of the present work was of M. caliginosus maintained on one of to examine the effect of different artn the 6 prey treatments. thropod prey species on the biology A record was kept of all life history parn and life-table parameters of the predatn rameters. Data for individuals which tory bug, M. caliginosus Wagner, under had been lost, accidentally killed or the laboratory conditions of 25oC, 75% which became stuck to the agar were r.h. and 16:8 L:D photoperiod. not included in the calculations. The M. caliginosus nymphs were reared individually in 5 cm diameter Materials and methods: Petri-dishes. Each Petri dish had a 2 cm diameter hole in the middle of the lid, Laboratory experiments were condn which was covered by 50 mesh cloth ducted in Wye College at University of to provide ventilation. The predant London to study the effect of different tors were provided with 5 cm diam. arthropod prey species on the rate of denv host plant leaf-discs heavily infested velopment, nymphal survival, fertility with assumed prey. Leaf-discs placed and life-table parameters of the predatn underside upwards on a 2 mm thick tory bug, M. caliginosus Wagner under agar media covered by a filter paper o laboratory conditions of 25 C, 75% r.h. to prevent the insects sticking to the and 16:8 L:D photoperiod. agar. The predators were transferred The experiment had 6 prey treatments to freshly prepared cages every day. as follows: To fix the relative humidity at 75% ± 1. Trialeurodes vaporariorum (Westnw 5%, 20 transparent plastic food boxes wood), Greenhouse whitefly (GHWF) (27 x 15 x 8.5 cm) were used. Each eggs supplied on tobacco leaf-discs. box included one cage from each prey 2. Metatetranychus urticae Koch, Green treatment in addition to two uncovered house red spider mites (GHRSM) of Petri-dishes which contained saturated different ages, on bean leaf-discs. solution of NaCl. 3. Ephestia kuehniella Zeller, eggs suppn Daily observations were made on the plied on Pelargonium peltatium leaves. frequency of moults and duration of survn 4. Myzus persicae Sulzer, Peach potato vival. Replicates in which the nymphs

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stuck to the agar were excluded. Upon the standard conditions of 25oC, 75% moulting to adult, each M. caliginosus relative humidity (r.h.), and 16:8 L:

was treated with CO2 so as they could D photoperiod. Daily observation for be weighed and measured and then hatching nymphs was done. All replicn each female was caged with one male cations in which the adult female died and reared together in a 5 cm diam before ovipositing were excluded from petri-dish cages which were changed the analysis. daily by freshly prepared cages through Data of age-specific survival and age- adult longevity of the adult female. specific fertility of M. caliginosus The adult M. caliginosus individuna which scored daily (Appendix 2) were als were transferred daily to freshly used to calculate life table parameters prepared cages and the assumed ovinp using a simple program of QBASIC position leaf-discs were kept for the version (Jervis and Copland, 1996). incubation period in incubator with

Results and Discussion: Growth and development of M. call tal nymphal developmental duration of liginosus nymphs offered different M. caliginosus was significantly shorter arthropod preys species: (15.7 days) on Ephestia eggs and (15.6 The developmental duration of M. calc days) on M. persicae, of medium length liginosus nymphs (particularly N3) was (17.4 days) on GHWF eggs and (18.2 significantly affected by the prey spenc days) on M. urticae and longest (22.3 cies offered (P < 0.05) (Table 1). The tont days) on A. fabae.

Table 1: Effect of prey species on the duration of each nymphal instar of M. caligicn nosus. Means (days)* ± s.e.

Treatment Duration of development (days) Prey (n)** N1 N2 N3 N4 N5 Total nymphs GHWF eggs 3.8±.42 3.4±.35 2.6 a ±.24 3.2±.31 4.3±.24 17.4 b ±.26 (16) GHRSM (6) 4.2±.40 2.7±.33 3.7 ab ±.62 3.5±.62 4.2±.31 18.2 b ±.48 Ephestia eggs 3.9±.19 3.0±.26 2.3 a ±.18 2.9±.43 3.7±.40 15.7 a ±.18 (15) PPA (14) 3.6±.20 2.4±.14 2.5 a ±.17 3.0±.30 4.0±.21 15.6 a ±.33 BBA (6) 5.3±.62 3.0±.36 4.7 b ±.88 4.2±.83 5.2±.87 22.3 c ±2.17 P. value .060 .118 .001 .462 .167 .001

*: Figures within columns with same letters do not differ significantly at p value 001 **: n= Number of replications which survived to become adult.

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Even though, whitefly predators are tabaci (Hoelmer et al., 1993) and then considered to be general predators, developed in 16 days on GHWF eggs previous studies have shown that diet at 26ºC (Hemachandra, 1994) and in 18 has an important effect on the duratn days on B. tabaci at 28ºC (Hoelmer et tion of development. M. caliginosus al., 1993). was recorded feeding on GHWF, Aphis The effect of diet on female longevity gosypii, Aulacorthum circumflexum, of M. caliginosus is shown in Table (2). Tetranychus turkestani and Ephestia The results of this study show that diet eggs (Fauvel et al., 1987). However, had a significant influence on the adult at 22ºC, M. caliginosus developed in longevity of M. caliginosus, with the 29.4 days on GHWF eggs, 28 days on greatest longevity on a diet of Ephesct either A. gosypii or T. turkestani, 26 tia eggs, medium longevity on GHWF days on either GHWF larvae or on A. eggs and M. persicae and the shortest circumflexum and in 22 days on Ephecs longevity on a diet of either M. urticae stia eggs. In addition, the coccinellid or A. fabae which was not enough to whitefly predator, Delphastus pusillus oviposition. With predatory Coccinelnl was also recorded feeding on GHWF lidae, Hemachandra (1994) found that (Hemachandra, 1994) and on Bemisia adult longevity could be affected by the

Table 2: Effect of prey species on female longevity of M. caliginosus. Means (days) ± s.e.*(n)**

Treatment Pren Ovip.n period Postn Average female ovip. ovip. longevity GHWF eggs 3.5±.35 (2) 4.0±2.12 b (2) 1.0±0.99 (2) 8.8±2.2ab (8) GHRSM *** nn nn nn 4.2±.85b (4) Ephestia eggs 4.0±.3 (7) 7.6±1.42a (7) 2.4±1.8 (7) 12.6±1.87a (8) PPA 5.2±0.79 (6) 3.2±1.42 b (6) 1.5±0.67 (6) 8.9±1.32ab (7) BBA*** nn nn nn 5.7±1.76b (3) P. value .24 .149 .838 .066

*: Figures within columns with different letters differ significantly according to Fisher’s pairwise comparisons. **: n= Number of females survived to become adult. ***: No oviposition recorded when fed on these treatments.

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adult diet, so that longevity of D. pusillc no oviposition when fed on M. urticae lus was longer on GHWF eggs than on and A. fabae. Furthermore, M. cacl GHWF larvae. liginosus females stopped ovipositing The pre-oviposition period varied bent half a day before death when fed on tween 3.5 days on GHWF eggs, 4 days Ephestia eggs, one day before death on on Ephestia eggs and 4.9 days on M. GHWF eggs and 1.5 day before death persicae , but these periods did not difnf on M. persicae. fer significantly (Table 2). In addition, The prey had a highly significant effn diets did have an effect on the ovipons fect on the growth rate of the predator, sition period, the longest being with with greatest mean relative growth rate Ephestia eggs (7.6 days) and the shorten of M. caliginosus when fed on Ephecs est when fed on GHWF eggs (4 days) stia eggs and least on A. fabae (Table and M. persicae (3.2 days). There was 3). Final adult weight was also greatest

Table 3: Effect of prey species on MRGR* of M. caliginosus. MRGR (mg/mg/ day) ** ± s.e.(n)***

Treatment M. caliginosus sex (Prey species) Females Males P value** Mean GHWF eggs 0.172 b±0.005(8) 0.157 b ± 0.004(8) 0.020 0.164 b ±0.004(16) GHRSM 0.164 bc ±0.006(4) 0.143 b ±0.009(2) 0.30 0.157 b ±0.006(6) Ephestia eggs 0.209 a±0.003(8) 0.190 a ±0.005(7) 0.009 0.200 a±0.004(15) PPA 0.191 ab 0.008(7) 0.182 a ±0.006(7) 0.39 0.187 a±0.005(14) BBA 0.120 c±0.019(3) 0.131 b0.017(3) 0.70 0.125 c±0.012(6) P value .001 .001 .001

*: Mean relative growth rate (mg/mg/day) = Loge adult weight - Loge initial weight total period of nymphal development where initial weight = mean weight of random sample of newly hatched nymphs. **: Figures within columns with different letters differ significantly at p value 0.001. ***:n= Number of replications survived to become adult

Table 4: Effect of prey species on adult weight of M. caliginosus. Means (mg)* ± s.e.(n)**

Treatment M. caliginosus sex (Prey species) Females Males P value Mean GHWF eggs .691 b±.023(8) .506 b±.016(8) 0.000 .598 b±.027(16) GHRSM .652 b±.021(4) .476 b±.022(2) 0.029 .593 b±.040(6) Ephestia eggs .956 a±.026(8) .643 a±.027(7) 0.000 .803 a±.045(15) PPA .702 b±.044(7) .546 b±.024(7) 0.012 .624 b±.032(14) BBA .573 b±.047(3) .438 b±.047(3) 0.130 .505 b±.042(6) P value .001 .001 .001

*: Figures within columns with different letters differ significantly at 0.001 level. **: n= Number of replications survived to become adult

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on Ephestia eggs and least on A. fabae tor to have a high rate of development. (Table 4). Males and females were affn Other species, such as M. urticae or fected similarly by each treatment, altn Aphis gossypii, did not. though females always weighed about 30% more than males. Size of adult M. caliginosus fed on The variation in nymphal growth rate different arthropod prey species: and developmental period of whitefly Prey species had a significant effect on predators have been shown to be reln the size of the predator when measured lated to the diet offered. Examples of at eclosion (Table 5). Body length, studies demonstrating this effect on head width, eye width and pronotum the hemipteran whitefly predators are width of both sexes were all significn those done using Anthocoris nemorc cantly different depending on the food rum (Russel, 1970; Campbell, 1977; offered during development. However, Ekbom, 1981), Macrolophus costacl vertex width and hind tibia length did lis (Brzezinski, 1982), M. caliginosus not differ. (Fauvel et al., 1987), Deraeocoris sp., When M. caliginosus was offered Ephecs Campylomma nicolasi and Geocoris stia eggs, the adults were significantly ochropterous (Kapadia and Puri, 1991) larger than when fed on M. persicae, and on the coccinellid whitefly predator GHWF eggs, spider mite or A. fabae. using D. pusillus (Hoelmer et al., 1993; In addition, head width and pronotum Hemachandra, 1994). width were greater when fed on either In this study, the diet offered to M. calc Ephestia eggs or M. persicae, rather liginosus significantly influenced many than on GHWF eggs, spider mites or A. of the biological parameters of the fabae. predator. Thus, when fed on Ephestia The size of males and females were eggs, M. persicae or GHWF eggs, M. found to differ in relation to the prey caliginosus had a shorter developmennt species. The pronotum widths of femn tal time and greater rate of growth than males fed on Ephestia eggs, GHWF when fed on M. urticae or A. fabae. eggs or M. persicae were larger than The present study is in general agreenm those of males. However, eye width, ment with the observations made on hind tibia and 2nd-antennal segment of M. caliginosus by Fauvel et al. (1987) males were larger than those of females when fed on 6 alternative diets. Fauvel when fed on Ephestia eggs, GHWF et al. (1987) also reported that different eggs or M. persicae. biological characteristics of M. caligicn nosus were significantly influenced by the prey species offered. They found that prey species such as T. vaporarioc orum or M. persicae enabled the predant

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Table 5: Size of adult M. caliginosus fed on five prey species. Means* ± s.e.**

*: Figures within rows with same letters do not differ significantly at 5% level according to Fisher’s pairwise comparisons. **: n= Number of replications survived to become adult Mortality of M. caliginosus nymphs ing from 20% on the GHWF eggs treatmn offered different arthropod prey specc ment to 83% on A. fabae and to 100% cies: in the control treatment. The percentan In all treatments, greatest mortality occn age of total mortality was significantly curred in the 1st instar whilst no mortalin lower when fed on GHWF eggs, Ephecs ity was observed during the 3rd instar stia eggs or M. persicae than on M. (Table 6 and Figure 1). In the control urticae or A. fabae with total mortalni treatment, where M. caliginosus nymphs ity of 20%, 21.05%, 36.36%, 70% and were reared on clean leaves of Pelargonn 82.86% respectively (P value = 0.000). nium, 80% died as 1st-instar nymphs, Therefore, M. caliginosus had the highne 10% survived to moult to a 4th-instar est survival rate when fed on GHWF nymph but none became adult. eggs and Ephestia eggs, medium surnv In addition, the results in Table 6 and vival on M. persicae but poor survival Figure 2 show that there was a wide on M. urticae and A. fabae. variation in the total nymphal mortality The percentage of female nymphs between the different treatments, rangni which survived to become adult was

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about 50% on all diets except on M. urtc through all age classes, although there ticae where it was 67%. Survivorship was accelerated mortality in the late curves were drawn using data of propn adult stage. portion of cohorts surviving on the Y- The prey-related differences in predator axis against age in days on the X-axis. survival were assumed to be due to factn The percentage surviving egg incubant tors such as differences in the qualitant tion was assumed to be 100 according tive attributes of the prey species such to Koppert observations (Klapwijik, as prey size, mobility and hardness 1995), while the data for survival durin (Jervis and Copland, 1996). Results of ing nymphal development and during this study show that diet had a remarkna adult stage were calculated from the able influence on the juvenile mortality data of the present experiment. of the predator. When M. caliginosus The results in Fig. 3 show that M. caligci was fed on GHWF eggs, Ephestia eggs inosus survivorship was greatest when or M. persicae, it had a significantly fed on Ephestia eggs > GHWF eggs > lower mortality during development M. persicae (PPA) > A. fabae (BBA) > than when fed on either M. urtica or A. spider mite (GHRSM). The survivorsn fabae. Previous studies on M. caligicn ship curves of M. caliginosus fed on eint nosus (Fauvel et al., 1987) and on M. ther Ephestia eggs, GHWF eggs or M. costalis (Brzezinski, 1982) have shown persicae fit a type I survivorship curve similar effects of diet on the mortality described by Jervis and Copland (1996), of the predators. Fauvel et al. (1987) where mortality is concentrated in the found that juvenile mortality for M. oldest age classes. However, when fed caliginosus was 11.8 % when fed on on either M. urticae or A. fabae. The GHWF at 25ºC and 20 % when fed on survivorship curves nearer a type II, Ephestia eggs at 20ºC, while Brzezinns where mortality is distributed equally ski (1982) found that the mortality of

Table 6: The effect of six prey species on the mortality of M. caliginosus during the nymphal stages. % mortality to the next instar ± s.e.*

*: s.e. calculated from the equation: s.e. =100√p q/n, where p = proportion died, q = 1np. **: n = number of replication. ***: P value calculated by χ2 analysis

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100 90 80 70 60 50 40 % mortality 30 20 10 0 GHWF GHRSM EPH PPA BBA Control eggs eggs

Prey species

1st instar 2nd instar 3rd instar 4th instar 5th instar

Figure 1: Effect of prey species on the distribution of mortality over the nymphal stages of M. caliginosus.

100 90 80 70 Mc 60 50 40 30 % mortality of 20 10 0 GHWF GHRSM EPH PPA BBA Control eggs eggs

Prey species

Figure 2: Effect of prey species on total mortality of M. caliginosus during nymnp phal stages.

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M. costalis was nil when fed on either on S. cerealella but 71.5 % when fed on GHWF or M. persicae, 25 % when fed M. urticae.

Figure 3: Survivorship curves of M. caliginosus when fed on GHWF eggs( ), GHRSM ( ), Ephestia eggs ( ), PPA ( ) and BBA ( ).

Fertility and life-table parameters the number of viable progeny that enns of M. caliginosus fed on different artc sue (i.e. the number of nymphs emergin thropod prey species: ing from the eggs laid) and females that In the case of M. caliginosus, it was produced nymphs are referred to as not easy to detect all eggs laid, and “fertile females”. therefore, the number of nymphs that M. caliginosus failed to produce nymphs emerged from the leaf tissue was consn when fed on M. urticae or on A. fabae. sidered to be a measure of the fertility. This was because they died during the Therefore, “fertility” here refers only to preoviposition period (Tables 7 and 8).

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Nymphs emerged only when M. caligicn nymphs produced per surviving female nosus had a diet of Ephestia eggs, M. of M. caliginosus was significantly persicae and GHWF eggs, with signifinc affected by its diet. The cumulative cantly more nymphs being produced number of nymphs produced when fed when M. caliginosus fed on Ephestia on Ephestia eggs was much higher than eggs than on GHWF eggs and M. persc that when fed on either M. persicae or sicae (P < 0.05) and with a significantly GHWF eggs. In addition, the oviposn higher percentage of “fertile” females sition period of M. caliginosus fed on when fed on both Ephestia eggs and Ephestia eggs were significantly longer GHWF eggs offered on tobacco + Pelca than that fed on GHWF eggs or on M. argonium than when fed on GHWF persicae . However, diet had no significn eggs on tobacco alone. In addition, Fig. cant effect on the incubation period of 4 shows that the cumulative number of M. caliginosus eggs (Table 8).

Table 7: : Percentage of fertile females and number of offspring produced when fed on each of different five prey species.

Table8 : Incubation period of M. caliginosus eggs laid by females reared on differen ent prey species. Means ± s.e.

Diet (n)* Oviposition host plant Incubation period (days) GHWF eggs (7) tobacco 10.6 ± .20 Ephestia eggs (58) Pelargonium 10.5 ± .14 M. persicae(15) tobacco 10.7 ± .25 P. value 0.899

*: n= Number of nymphs emerged

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Feeding on GHWF eggs, M. persicae lar incubation period, averaging 10.6 or on Ephestia eggs resulted in a siminl days.

70

60 GHWF eggs Ephestia eggs 50 M. persicae

40

30

20

Cumulative nymphs/female/day 10

0 25 30 35 40 45

Age (days)

Figure 4: Cumulative number of nymphs emerging from eggs laid per day age of M. caliginosus females fed on GHWF eggs, Ephestia eggs and M. persicae.

The present results are similar to those depending on the prey species and on of Fauvel et al. (1987) for M. caligicn the host plant species. nosus and Albajes et al. (1996) for Dicc The effect of the diet offered to M. cyphus tamanini. Fauvel et al. (1987) caliginosus on life-table parameters found that M. caliginosus had a high showed that the gross reproductive rate cumulative fecundity on Ephestia eggs, (GRR), net reproductive rate (Ro), innt medium on both GHWF eggs and M. trinsic rate of increase (rm) and generant persicae, and a low cumulative fecundn tion time (T) of the predator were highen dity on both A. gossypii and Tetranycc est with a diet of Ephestia eggs (Table chus turkestani. Albajes et al. (1996) 9). The doubling time (DT) was also had also found that the fecundity of shortest with Ephestia eggs. these mirid predators was quite variable

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Feeding on either GHWF eggs or on On the other hand, the intrinsic rate of M. persicae offered on tobacco leaf- increase of M. caliginosus when fed discs resulted in low reproductive rates, on Ephestia eggs offered on Pelargocn with even the intrinsic rate of increase nium leaves alone was 0.0825 insects being negative. However, when Pelarcg per day, resulting in a doubling time of gonium leaves were offered as well as 8.4 days, while on GHWF eggs offered the tobacco leaf-discs at the adult stage, on tobacco leaf-discs + Pelargonium the reproductive rates of the predator leaves, the intrinsic rate of increase was fed on GHWF eggs became higher and only 0.0423 per day resulting in a the doubling time was shorter (Hamdn doubling time of about 16.4 days. dan, 1997).

Table 9: Life-table parameters* of M. caliginosus when fed on different prey spenc cies.

Diet treatments/oviposition site (n) Lifectable GHWF eggs/ GHRSM/ Ephestia eggs/ PPA / BBA/ parameters tobacco beans Pelargonium tobacco broad bean (8) (4)** (8) (7) (3)** % fertile 25% 0.0 87.5% 85.5% 0.0 females GRR 1.7 nn 34.8 3.2 nn Ro 0.9 nn 18.2 1.0 nn rc n0.00175711 nn 0.08131106 0.001326906 nn rm n0.00175711 nn 0.08254354 0.001327247 nn Tc 32.8 nn 35.7 33.5 nn T 32.8 nn 35.1 33.5 nn λ 0.9982444 nn 1.086046 1.0013228 nn DT n394.5 nn 8.4 522.2 nn

*: Lifentable parameters: gross reproductive rate (GRR) cohort generation time (days) (Tc) net reproductive rate (Ro) generation time (days) (T) capacity for increase (rc) finite capacity for increase (λ) intrinsic rate of increase (rm) doubling time (days) (DT) **: The lifentable parameters could not be calculated because no nymphs emerged at these diets.

Heteroptera and auchenorrhynchan bipunctata) (Hurst et al., 1993 and Ottn bugs always reproduce bisexually. tenheim et al., 1992). However, there Mating is usually essential to the prond were no reports of a female biased sex duction of viable offspring, and the sex ratio in mirid bugs. On the other hand, ratio of offspring is close to unity. The the results of this research showed that resultant eggs are laid (ovipary) on folian the percentage of female M. caliginosus age, bark or litter, or inserted into plant under the laboratory conditions of 25C, tissues (McGavin, 1993). 16:8 L:D and 75% r.h. were: < 50% Female biased sex ratio has been recorden when nymphs have been feeding on ed in two species of predatory ladybirds either GHWF eggs or larvae; 50% on (Coccinella septempunctata and Adalia either of M. persicae or A. fabae; 53%

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on Ephestia eggs and 67% on M. urticc when fed on M. persicae on tobacco cae. Therefore, it is concluded that the leaf-discs but rm was high (0.08254) sex ratio of M. caliginosus could be afnf and the doubling time was 8.4 days fected by the prey quality which might when M. caliginosus fed on Ephestia affect the survivorship of each sex. eggs offered on Pelargonium leaves. The results of this experiment showed In addition, the results of the previous that the host plant species offered for experiment (Hamdan, 1997) show that oviposition had affected the fertility rm was 0.04229 and the doubling time of M. caliginosus. The intrinsic rate was 16.4 days when M. caliginosus of increase (rm) of M. caliginosus was fed on GHWF eggs offered on tobacco negative (- 0.00175 female/female/day) leaf-discs in the presence of Pelargocn when fed on GHWF eggs offered on nium leaves as an alternative ovipositn tobacco leaf-discs, very low (0.00133) tion sites.

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7. Foglar, G., Malausa, J. C. and Wajnberg, References: E. (1990). The functional response and prefen 1. Albajes, R., O. Alomar, J. Riudavets, C. erence of Macrolophus caliginosus [Heteroptn Castane, J. Arno and R. Gabarra (1996). tera: Miridae] for two of its prey: Myzus percs The mirid bug Dicyphus tamininii: an efnf sicae and Tetranychus urticae. Entomophaga, fective predator for vegetable crops. IOBC/ 35(5), 465-474. WPRS Bulletin, The International Organizatc 8. Gerling, D. (1990). Natural enemies of tion for Biological and Integrated Control of whitefly: Predators and parasitoids. In: Gerln Noxious and Plants, West Palaearctc ling, D. (ed.), Whiteflies; Their Bionomics, tic Regional Section, 19(1), 1-4. Pest Status And Management, Intercept Ltd. 2. Brzezinski, K. (1982). Report from investn UK, pp. 147-185. tigations on morphology, biology and ecology 9. Hamdan, A. J. S. (1997). Biological and of Heteroptera Macrolophus costalis (Fieb.) Ecological Studies on the Predatory Bug (Heteroptera, Miridae) and its predacity in Macrolophus caliginosus Wagner [Heteroptn relation to greenhouse whitefly Trialeurodes tera: Miridae] as a Biocontrol Agent Against vaporariorum (Westw.). Materialy, XXII I the Greenhouse Whitefly Trialeurodes vapocr XXIII Sesji Naukowej Instytutu Ochrony Roslc rariorum (Westwood) [Homoptera: Aleyrodidn lin, Poznan, 285-292. dae]. Ph.D Thesis, Wye College, University 3. Campbell, C. A. M. (1977). A laborant of London, 254 pp. tory evaluation of Anthocoris nemorum and 10. Hemachandra, K. S. (1994). Developmn nemoralis (Hemiptera: Anthocoridae) as ment, Reproduction And Behaviour Of Delcp predators of Phorodon humuli (Homoptera: phastus pusillus, A Coccinellid Predator Of Aphididae). Entomophaga, 22, 309-414. Glasshouse Whitefly, Trialeurodes vaporarioc 4. Ekbom, B. S. (1981). Efficiency of the orum . M.Phil. Thesis, Wye College, Universn predator Anthocoris nemorum (Hemiptera: sity of London, 169pp. Anthocoridae) against the greenhouse whitefn 11. Hoelmer, K.A., Osborne, L. S. and Yokomc fly, Trialeurodes vaporariorum (Homoptera: mi, R. K. (1993). Reproduction and feeding Aleyrodidae). Zeitschrift fur Angewandte Enct behaviour of Delphastus pusillus (Coleoptera: tomologie, 92, 26-34. Coccinellidae), a predator of Bemisia tabaci 5. Eremenko, A. P. (1984). The biological (Homoptera: Aleyrodidae). Journal of Ecocn method in glasshouses. Zashchita Rastenii, nomic Entomology, 86(2), 322-329 11, 18-19. 12. Hurst, G.D.D., Majerus, M. E. N. and 6. Fauvel, G., Malausa, J. C. and Kaspar, Walker, L. E. (1993). The importance of B. (1987). Etude en laboratoire de principals cytoplasmic male killing elements in natural characteristics biologiques de Macrolophus populations of the two spot ladybird, Adalia caliginosus (Heteroptera: Miridae). Entomn bipunctata mophaga, 32, 5, 529-543.

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