J. Agric. Sci. Technol. (2002) Vol. 4: 1-10

The Relationship Between Oviposition Preference and Larval Performance in an Aphidophagous Hover , ribesii L. (Diptera: Syrphidae)

1 H. Sadeghi

ABSTRACT

The relationship between female oviposition preference and offspring performance is a key area of study on the evolution of host specificity. This study investigates host prefer- ence of the polyphagous hover fly (Diptera: Syrphidae) and compares it with some components of its offspring`s performance. Females and offspring were tested on six naturally used hosts (blackberry, dock, nettle, pea, rose, and sycamore ). In the laboratory, larvae were able to develop successfully on aphids which were rarely selected for oviposition by gravid females. Despite the relatively small differences in performance resulting from feeding on different aphids, there was evidence for a pref- erence-performance correlation in the generalist S. ribesii.

Keywords: Host choice, Host specificity, Fitness, Preference-performance.

INTRODUCTION specialization [3]. Numerous studies on a variety of phyto- In many where the newly hatched phagous insects have investigated the pref- offspring are relatively sessile and are not erence-performance correlation, but there is able to move any great distance to search for no consensus, with the results ranging from another appropriate host, host choice is strong [2, 29] to weak correlation [ 6, 7, re- made by the ovipositing female. In such view in 48]. Reasons put forward for the cases, offspring must often feed on the host variation in outcome of these experiments chosen by their mother. Whether or not the include variation among the measures of mother knows the best food for further de- performance used [24, 29, 47] and differ- ences in the ability of larvae to move to al-

Downloaded from jast.modares.ac.ir at 1:20 IRST on Thursday September 30th 2021 velopment of her offspring is a controversial point (7). As a result of natural selection, ternative food sources [30, 31]. Nothing is one might expect a positive correlation be- known about this matter in predatory insects. tween female oviposition preference and Larval performance depends mainly on offspring performance. A good correlation food quality and mortality due to natural between these suggests that host characteris- enemies. There is presumably little differ- tics including host chemistry and nutritional ence between the nutritional requirments of value, are key factors that influence off- predatory insects and the contents of their spring performance, determine host range, prey [4] and, as a result, the cost of capture and promote host specificity. Conversely, and the toxin content of prey are probably poor correlations are used as evidence that relatively more important to the overall costs other ecological factors, such as the selec- of feeding [23]. tion of enemy-free space are of greater im- In this research, the relationship between portance in the evolution of host choice and preference and performance in a predatory ______1 Department of Plant Protection, College of Agriculture, Ferdowsi University, Mashhad Islamic Republic of Iran.

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hover fly (Diptera: Syrphidae) was investi- Ruzicka [38]). A possible explanation for gated. In hover , like many other insects, these apparent contradictions lies in the fact ovipositing females select among food types that aphid toxins may be, at least in part, [41]. For a polyphagous syrphid, oviposi- sequestered from the host plants [17, 22, tional preference by females may have a 23], which are variable in toxin content in- profound effect on the performance of the ter- and intra-specifically and, in one indi- offspring, because syrphid larvae probably vidual plant, through time [see 44]. There have rather limited dispersal abilities, al- are a few explicit demonstrations of the way though some indications imply larvae can in which variation in host-plant chemistry move to new aphid colonies to a certain ex- affects the suitability of aphids as prey for tent. For example, Banks [1] suggested that syrphid larvae [9, 21, 22, possibly Paragus even quite small syrphid larvae must move longiventris in 43]. between plants and possibly 'considerable This study aims firstly, to compare the ef- distance'. Kan [19, 20] suggested that a sin- fect of different aphid species as larval food gle maple- or pea aphid-colony was insuffi- on the performance of Syrphus ribesii under cient to support larval development to ma- laboratory conditions, testing the null hy- turity, and noted that 'older larvae are ob- pothesis that all aphids are equally good as served actively migrating among maple food for the larvae. Secondly, we test branches' or pea plants. However, most au- whether there is an overall preference- thors agree that the larvae are more or less performance correlation in this species, us- sedentary (e.g. F. Gilbert; G. Rotheray, per- ing egg distribution among aphids in labora- sonal communication), and almost com- tory experiments as our preference measure. pletely dependent on their mothers to chose The hover fly chosen for use in this study a suitable host prey. is a widespread and very common species Rather few studies of syrphid larval devel- [14, 37]. The larvae are often the most opment compare quantitatively any aspect of common predators and have been reported larval performance on different aphids, with from colonies of more than 100 species of the notable exceptions of Schmutterer [43], aphids worldwide (F. Gilbert unpublished Ruzicka [38] and Malcolm [22, 23]. Al- data). It is oligovoltine, with 2-3 generations though involving disparate studies with dif- per year, overwintering as an exceptionally ferent aims and very different sets of meas- cold-tolerant larva [15]. Adult females are ured variables, some generalities can be normally ready to lay eggs 7-8 days after drawn. While development time and pupal emergence. The larvae of this syrphid are

Downloaded from jast.modares.ac.ir at 1:20 IRST on Thursday September 30th 2021 weights are often unaffected [8, 27], aphid- very generalized in their feeding habits, but prey species and/ or prey quality do seem to nonetheless show different oviposition pref- affect mortality. Several aphids are probably erences and field distributions on different toxic or partially toxic to the larvae of one or aphid species [40] and oviposition prefer- more species of hover fly [8, 9, 21, 22, 32, ences [41]. 38, 43]. Other variables have rarely been considered, but can be affected by food MATERIALS AND METHODS quality (e.g. ease of capture) [39, 52]. It is therefore unfortunate that several studies measured only development time or pupal Study Organism weight. While some aphids were consis- tently identified as unsuitable (e.g. Megoura The individuals of the syrphus ribesii L. viciae aphids studied by Milne [25], Ruzicka (Diptera: Syrphidae) used in the experiment [38] and Xiong and Dong [53]), several came from gravid females collected from aphids were deemed highly toxic in one Nottingham University Campus. The aphids study, but suitable in another (e.g. Bre- used in this experiment were chosen for their vicoryne brassicae by Schmutterer [42] and availability, and because in both field and by

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experiment [40] they form a continuum from ony; presentations continued throughout the good to poor prey. Pea aphids Acyrthosiphun oviposition preiod of 14–38 days. 13 syrphid pisum (Harris) were obtained from stock females were used in the oviposition prefer- cultures on broad bean (Vicia faba L.). All ence experiment. other aphids were collected from the Uni- versity Campus including Aphis fabae L. Larval Performance from dock (Rumex crispus L.), Microlo- phium carnosum (Buckton) from nettle (Ur- tica dioica L.), Macrosiphum rosae (L.) The culturing of Syrphus ribesii follows from rose (Rosa canina L.), Aphis ruborum the methods of Frazer [11] and Hart and Börner from blackberry (Rubus fruticosus Bale [15, 16] with slight modifications. L.), and Drepanosiphum plantanoidis Adults were kept in a constant environment Schrank from sycamore (Acer pseudoplata- of 22-23°C with a 16 hour photoperiod, and nus L.). All are known to be the natural prey were provided with bee-collected pollen of larvae from this hover fly; the larvae have (Sigma Ltd, Cat. No.P-8753), crystalline been found in colonies of all these aphid sugar placed on petri-dish lids on the floor species [40]. of the cage, and water on a soaked pad of cotton wool in a conical flask. Both pollen and water were changed every 2-3 days. Oviposition Preference To obtain a group of larvae of the same age, females were induced to lay eggs on cut For experimenting on oviposition prefer- sections of broad bean plant (Vicia faba var. ence, females of known age were used. Fe- Aquadulce claudia) infested with pea males were initially naive, having had no aphids. For experimental purposes, a batch previous exposure to aphids. On the day of of eggs laid over a period of three hours was eclosion, an equal number of females and selected and placed in a large Petri dish to males were transferred to the adult rearing hatch. The percentage mortality of the early cages. Mating was completed after 3-4 days, larval instar is normally higher than the and the ovaries began to enlarge about a other instars, because larvae at this time are week after emergence. Once the majority of delicate and difficult to handle. In addition, females contained some mature eggs (easily in our experiments there was a low percent- seen through the transparent abdominal age of egg hatch, and together, these factors pleurites), individual gravid females were decreased the number of larvae available for transferred to separate cages. Aphids were study. To counter this, for the first three Downloaded from jast.modares.ac.ir at 1:20 IRST on Thursday September 30th 2021 then offered to each female on a newly-cut days, larvae were left in groups and allowed section of their host-plant standing in water; to feed on pea aphids (during this period, great care was taken to ensure that all cut only one or two aphids are normally con- sections were about the same size and with sumed). At the beginning of the fourth day, the same number of aphids of various in- larvae were weighed individually, trans- stars. Each day, aphids were presented in a ferred to experimental Petri dishes, and randomized sequence to each syrphid female given aphids of the selected species. (i.e, a no-choice situation, with only one Each day, the larvae were weighed, and aphid species available at any time). Each enough similarly-sized aphids to supply food presentation of an aphid species lasted for 30 for 24 hours were weighed and added to the minutes. The number of eggs laid in re- Petri dishes, the remains being weighed sponse to the presented aphid were counted, again the following day. Each larva received and the aphid replaced by another aphid spe- only one aphid species. Depending on avail- cies, continuing until all aphids had been ability, 10 to 16 replicates were used, except presented. Each day females had the choice in the case of blackberry aphid, for which of ovipositing in any of the test aphid col- only 6 larvae were reared because of the

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scarcity of this aphid in the field. The result- RESULTS ing pupae were weighed, and the potential fecundity of the resulting adult females Oviposition Preference measured by dissecting them and counting the ovariole numbers of both ovaries. Individual fitness was calculated (r) as a The distribution of eggs among aphids performance measure [24] by integrating showed significant differences in the relative development time (D: egg - pupa), survival preference of aphids by females (KW = (m=1 or 0) and potential fecundity (V) via 76.5, P< 0.001; Fig. 1). With a few excep- the equation: r = [Ln (m.V)] / D, where Ln = tions, sycamore aphid was ranked the high- natural logarithms. Fitness was calculated est by all females, with on average about twice, once for surviving females only, and 26.6% of eggs, followed by rose aphid then for all females including non-survivors. (24.5%) and pea aphid (20.8%). The lowest In the latter, because female larvae cannot preference was for nettle aphid, with about be differentiated from males, I assumed half 5.9% of the eggs. There were some differ- the mortality to be female, rounding up ences among different aphids in terms of when an odd number of larvae had died. acceptability by individual females. For ex- Differences among means were assessed ample, only two out of 13 females laid eggs using a one-way Anova, except for individ- on nettle aphid colonies on the first day of ual fitness including that of non-survivors, oviposition, but the rest oviposited on the where a non-parametric Anova test was used colonies of this aphid after days 3 or 4 of the (H, distributed as χ²); survival differences experiment. All the other aphids were used were tested using χ2. for oviposition by females from the first day of oviposition. Downloaded from jast.modares.ac.ir at 1:20 IRST on Thursday September 30th 2021

Figure. Percentage ( ± se) of lifetime fertility of female Syrphus ribsii laid in response to particular aphid species averaged across females (n=13). The differences are highly significant (KW= 76.5, P< 0.001).

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Table 2. Hierarchy of aphid species in relation to oviposition preference and suitability for lar- val development (from 41). Oviposition Suitability for lavral Aphid species Plant species preferencea development allb survivorsc Acyrthosiphon pisum Vicia faba 18.9 11.8 14.8 Apbis fabae Rumex obtusifolius 11.1 8.3 13.8 Aphis grossulariae Epilobium hirsutum - - - Aphis pomi Malus domestica - - - Aphis ruborum Rubus fruticasus 7.0 9.5 14.2 Aphis sambuci Sambucus nigra 6.9 - - Cavariella sp Haracleum sphondylium 9.8 - - Drepanosiphum plantanoidis Acer pseudoplantanus 21.4 15.9 15.9 Marasipbum rosae Rasa sp 20.3 15.7 15.7 Microlophium carnosum Urtica dioica 4.6 12.0 15.0 Phyllaphis fagi Fagus sylvatica - - - Schizoneura ulmi Ulmus sp - - - a Mean percentage of total egg load laid on an aphid during no-choice sequential oviposition tests b Mean individual fitness of all individuals tested, calculated according to McGraw and Caswell (25). c Mean individual fitness of those surviving to adulthood, calculated according to McGraw and Casweel (25).

Larval Performance The number of ovarioles of adult females ranged from 61 to 120, but the mean values The performance measures of S. ribesii were not significantly different. reared on the different aphid species are Individual fitness of survivors (Table 1) given in Table 1. The results showed only differed significantly among aphid-prey relatively small but nevertheless significant treatments, with larvae fed on sycamore and differences in the suitability of most aphid rose aphids having greater fitness. The in- species as larval food. creased variance associated with the inclu- There were no significant differences in sion of non-survivors (i.e. individual females survival among treatment groups. The sur- with zero fitness) led to no significant dif- vival rate until pupation ranged from 75 % ferences among the mean fitnesses of all for dock aphid to 100% for sycamore and females, although a non-parametric ANOVA Downloaded from jast.modares.ac.ir at 1:20 IRST on Thursday September 30th 2021 rose aphids. Substantial mortality occurred remained significant (Kruskal-Wallis H = during the pupal period, particularly on rose 13.1, p < 0.05). aphids. The mean larval development period ranged from 9.6 to 11.6 days. The mean val- Oviposition Preference Versus Larval ues were significantly different among Performance groups (F5,59= 8.76; p<0 .001), being slightly longer on blackberry and dock aphids than Testing for a positive (rank) correlation the very similar but more rapid times for the between oviposition preference and the per- other four aphid species. formance of offspring measured by individ- Pupal weights ranged from 32.4 mg (on ual fitness (data in Table 2), there were indi- dock aphid) to 70.8 mg (on sycamore aphid), cations of a positive relationship between and were significantly different among oviposition preference versus larval per- aphid-prey groups. Larvae fed on rose and formance (r =0.60, n=6, p=0.09). sycamore aphids on average resulted in s heavier pupae, whereas the pupae of larvae reared on blackberry and dock were lighter.

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DISCUSSION larvae in this study, one might expect to see a better performance in the larvae fed on pea A number of factors, including those men- aphids. tioned by Thompson and Pellmyr [48], may The results showed that even in this very be the cause of a weak correlation between generalized predator, there are significant oviposition preference and larval perform- differences in the distribution of eggs among various aphids. This supports suggestions in ance in the syrphid studied here. In the field, it is possible that the relation- the literature of selectivity of oviposition [5, ship between oviposition preference and per- 25, 26, 29, 35]. formance is influenced by differential levels As in other studies[33, 34, 36, 45, 50, 51], of attack by natural enemies on preferred or the evidence suggests that the larvae of this non-preferred aphid species: the proportion syrphid species are able to develop success- of syrphid larvae killed by specialist parasi- fully on nearly all aphid species offered to toids can be very high [14, 37]. However, them. In other words, the host range of the the effect of natural enemies on host choice larvae, as measured by different components by ovipositing females was not investigated of larval performance, is wider than that of in the present study. Studies have also the ovipositing females. shown that factors such as egg load and age Larval conservatism (i.e. being generalist may affect a female’s oviposition preference in feeding behaviour) along with the results [e.g. 10]. Earlier works [41, 42] showed that, of studies on host shifts [18, 46] are used as as a female ages or when she has accumu- evidence that host shifts only involve a be- lated eggs, the discrepancy among preferred havioural change in host preference. It has and less preferred aphid species decreases. been argued that, in general, behavioural One important point to be mentioned is the (i.e. preference) flexibility should be more possible effect of host plant on the oviposi- evolutionarily labile than morphological or tion preference of females. The treatments in physiological (i.e. performance) traits, and oviposition preferences were aphids with a for this reason, the more labile characters cut section of their host plant. Consequently, should be the initiators of new directions in there is the possiblity that the significant the evolution of host utilization [49]. Other differences among treatments were due to authors [12,13, 36, 44, 45] have also stressed different aphids, plants, or both. For exam- the role of behavioural adaptation in the spe- ple, plant traits such as the presence of cialization process, and the leading role of trichomes and many others [see 44] are often the behaviour of ovipositing females in evo-

Downloaded from jast.modares.ac.ir at 1:20 IRST on Thursday September 30th 2021 assumed to influence the host plant selection lution of host utilization [18]. The behav- behaviour. As a result, it is possible that the ioural sequence followed by gravid females oviposition preferences observed could be during oviposition may shed more light on due either to the aphid or plant effect. syrphid specialization, especially in the light There is plenty of evidence on the effect of of the hypothesis of selective attention (i.e. learning on host selection by adult females the costs associated with obtaining and as- [see 7], but very little is known of the effect sessing information for generalists are of prior experience of larvae on their per- avoided by specialists, so the specialist spe- formance. Without providing data, Schmut- cies will make faster and more accurate de- terer [43] notes that small larvae of Afrosyr- cisions than generalists). This could be the phus varipens collected from dactynotus next study to be undertaken in unravelling compositae on Vernomia lasiopus developed the evolution of prey specialization in the normally when transferred to Toxoptera ci- aphidophagous Syrphidae. tricidus, but were killed by Aphis fabae so- lanella. However, if there was any effect of early feeding on the performance of the test

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ACKNOWLEDGEMENTS 10. Fitt, G. P. 1986. The Influence of a Shortage of Hosts on the Specificity of Oviposition Behaviour in Species of Dacus (Diptera, Francis Gilbert kindly supervised the pro- Tephritidae). Physiol. Entomol., 11: 133- ject. Many other people, specially Andy 143. hart, Jeff Bale, Graham Holloway and Rich- 11. Frazer, B. D. 1972. A Simple and Efficient ard Harington helped with rearing aphids Method of Rearing Aphidophagous Hover- and syrphids. The greenhouse staff of Not- flies (Diptera: Syrphidae). J. Entomol. Soc. tingham University helped me with culturing Brit. Columb., 69: 23-24 broad bean plants during experimental peri- 12. Futuyma, D. J. 1983. Selective Factors in the ods and for this I am deeply grateful to all of Evolution of Host Choice by Phytopha In- sects. In: “Herbivorous Insects: host Selec- them. tion Behaviour and Mechanisms”. (Ed.), Ahmad S. Academic Press, New York. pp. REFERENCES 227-244. 13. Futuyma, D. J., Cort, R. P. and van Noord- wijk, I. 1984. Adaptation to Host Plants in 1. Banks, C. J. 1968. Effects of Preda- the Fall Cankerworm (Alsophila pometaria) tors on Small Populations of Aphis fabae in and its Bearing on the Evolution of Host Af- the Field. Entomol. Exp. Appl., 11: 169-176. filiation in Phytophagous Insects. Amer. 2. Barker, A. M. and Maczka, J. M. 1996. The Naturalist., 123(3): 287-196. Relationships Between Host Selection and 14. Gilbert, F. S. 1993. “”. Natural- Subsequent Larval Performance in Three ists’ Handbooks 5, 2nd edition. Richmond Free-living Graminivorous Sawflies. Ecol. Press. 72 pp. Entomol., 21: 317-327. 15. Hart, A. J. and Bale, J. S. 1997a. Evidence 3. Bernays, E. A., and Graham, M. 1988. On for the First Strongly Freeze Tolerant Insect the Evolution of Host Specificity in Phyto- Found in the UK. Ecol. Entomol., 22: 242- phagous . Ecology., 69: 886- 892 245. . 16. Hart, A. and Bale, J. S. 1997b. A Method for 4. Bristowe, C. M. 1988. What Makes a Preda- Mass-rearing the Episyrphus bal- tor Specialize ? Trend. Ecol. Evolut., 3: 1-2. teatus (Diptera, Syrphidae). Dipterists Di- 5. Budenberg, W. J., and Powell, W. 1992. The gest., 4(1): 1-3. Role of Honeydew as an Oviposition Stimu- 17. Hodek, I. 1993. Habitat and Food Specificity lant for Two Species of Syrphids. Entomol. in Aphidophagous Predators. Biocontrol Exp. Appl., 64: 57- 61. Sci.Technol., 3: 91-100. 6. Burstein, M., and Wool, D. 1993. Gall 18. Janz, N., Nylin, S. and Wedell, N. 1994. Aphids do not Select Optimal Galling Sites. Host Plant Utilization in the Comma Butter- Downloaded from jast.modares.ac.ir at 1:20 IRST on Thursday September 30th 2021 Ecol. Entomol., 18: 155- 164. fly: Source of Variation and Evolutionary 7. Courtney, S. P., and Kibota, T.T. 1990. Implications. Oecologia., 99: 132-140. Mother Does not Know Best: Selection of 19. Kan, E. 1988a. Assessment of Aphid Colo- Hosts by Ovipositing Insects. In: “Insect- nies by Hoverflies. 1. Maple Aphids and plant interactions II” E. A. Bernays. CRC Episyrphus balteatus (de Geer) (Diptera: Press. (Ed.) pp. 161-188. Syrphidae). J. Ethol., 6: 39-48. 8. Du, Y-Z., and Chen, X-Z. 1993. Influence of 20. Kan, E. 1988b. Assessment of Aphid Colo- Different Aphid Prey on the Development of nies by Hoverflies. II. Pea Aphids and 3 Metasyrphus corollae (Dip., Syrphidae) (in Syrphid Species: Betasyrphus Serarius Chinese with english summary). Chinese J. (Wiedemann), Metasyrphus Frequens Ma- Biol. Control., 9: 111-113. tsumura and (Meigen) 9. Emerich, B. H. 1991. Acquired Toxicity of Diptera: Syrphidae). J. Ethol., 6: 135-142. the Lupin Aphid, Macrosiphum albifrons, 21. Kaufmann, T. 1973. Biology of Paragus and its Influence on the Aphidophagous borbonius (Diptera: Syrphidae) as a predator Predators Coccinella septumpunctata, Epi- of Toxoptera aurantii (Homoptera: Aphidi- syrphus balteatus and Chrysoperla carnea dae) Attacking Cocoa in Ghana. Amer. Mid- [in german with summary in English]. Z. land Naturalist., 90: 252-256. pflanzenkr. Pflanzensch., 98: 398-404.

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ﺭﺍﺑﻄﻪ ﺑﻴﻦ ﺭﺟﺤﺎﻥ ﺗﺨﻢ ﺭﻳﺰﻱ ﻭ ﺭﺷﺪ ﻭ ﻧﻤﻮ ﻻﺭﻭﻫﺎ ﺩﺭ ﻳﻚ ﮔﻮﻧﻪ ﻣﮕﺲ ﮔﻞ Syrphus ribesii L. (Diptera: Syrphidae)

ﺡ. ﺻﺎﺩﻗﻲ

ﭼﻜﻴﺪﻩ

ﺭﺍﺑﻄﻪ ﺑﻴﻦ ﺭﺟﺤﺎﻥ ﺗﺨﻢﺭﻳﺰﻱ ﻣﮕﺲ ﮔﻞ ﻣﺎﺩﻩ ﻭ ﺭﺷﺪ ﻭ ﻧﻤﻮ ﻧﺘﺎﺝ ﻳﻜﻲ ﺍﺯ ﻋﺮﺻﻪ ﻫﺎﻱ ﺍﺻﻠﻲ ﻣﻄﺎﻟﻌﺎﺕ ﻣﺮﺗﺒﻂ ﺑﺎ ﻓﺮﺁﻳﻨﺪ ﺗﻜﺎﻣﻠﻲ ﺍﺳﺘﻔﺎﺩﻩ ﺍﺯ ﺗﺨﺼﺺ ﺩﺭ ﻣﻴﺰﺑﺎﻥﻳﺎﺑﻲ ﺍﺳﺖ. ﺍﻳﻦ ﻣﻄﺎﻟﻌﻪ ﺭﺟﺤﺎﻥ ﻣﻴﺰﺑﺎﻧﻲ ﺭﺍ ﺩﺭ ﻣﮕﺲ ﭘﻠﻲ ﻓﺎﮊ Syrphus ribesii ﺑﺮﺭﺳﻲ ﻭ ﺁﻧﺮﺍ ﺑﺎ ﺑﺮﺧﻲ ﺍﺟﺰﺍ ﺭﺷﺪ ﻭ ﻧﻤﻮ ﻧﺘﺎﺝ ﻣﻘﺎﻳﺴﻪ ﻣﻲﻛﻨﺪ. ﺣﺸﺮﻩ ﻣﺎﺩﻩ ﻭ ﻻﺭﻭﻫﺎ ﺭﻭﻱ ﺷﺶ ﮔﻮﻧﻪ ﺍﺯ ﺷﺘﻪﻫﺎﻱ ﻣﻴﺰﺑﺎﻥ ﻃﺒﻴﻌﻲ ﺁﻧﻬﺎ: ﺷﺘﻪ ﺗﻤﺸﻚ، ﺭﺯ، ﻧﺨﻮﺩ، ﺍﻓﺮﺍ، ﺗﺮﺷﻚ ﻭ ﮔﺰﻧﻪ ﻣﻮﺭﺩ ﺁﺯﻣﺎﻳﺶ ﻗﺮﺍﺭ Downloaded from jast.modares.ac.ir at 1:20 IRST on Thursday September 30th 2021 ﮔﺮﻓﺘﻨﺪ. ﺩﺭ ﺷﺮﺍﻳﻂ ﺁﺯﻣﺎﻳﺸﮕﺎﻫﻲ ﻻﺭﻭﻫﺎ ﺑﺎ ﻣﻮﻓﻘﻴﺖ ﺑﻴﺸﺘﺮﻱ ﺭﻭﻱ ﺷﺘﻪ ﻫﺎﻳﻲ ﻛﻪ ﺑﻨﺪﺭﺕ ﺑﺮﺍﻱ ﺗﺨﻢ ﺭﻳﺰﻱ ﺍﻧﺘﺨﺎﺏ ﻣﻲﺷﻮﻧﺪ ﺭﺷﺪ ﻳﺎﻓﺘﻨﺪ. ﻋﻠﻴﺮﻏﻢ ﺍﺧﺘﻼﻑ ﺟﺰﻳﻲ ﺩﺭ ﺭﺷﺪ ﻭ ﻧﻤﻮ ﺩﺭ ﻧﺘﻴﺠﻪ ﺗﻐﺬﻳﻪ ﺍﺯ ﺷﺘﻪﻫﺎﻱ ﻣﺨﺘﻠﻒ، ﺷﻮﺍﻫﺪ ﺣﺎﻛﻲ ﺍﺯ ﻭﺟﻮﺩ ﻳﻚ ﻫﻤﺒﺴﺘﮕﻲ ﺑﻴﻦ ﺭﺟﺤﺎﻥ ﺗﺨﻢﺭﻳﺰﻱ ﻭ ﺭﺷﺪ ﻭ ﻧﻤﻮ ﻻﺭﻭﻫﺎ ﺩﺭ ﻣﮕﺲ ﺳﻴﺮﻓﻴﺪﻩ Syrphus ribesii ﺑﻮﺩ.

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