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Ceratitis Capitata (Diptera: Tephritidae) Heredity (2002) 89, 480–487 2002 Nature Publishing Group All rights reserved 0018-067X/02 $25.00 www.nature.com/hdy Sexual selection on multivariate phenotype in wild and mass-reared Ceratitis capitata (Diptera: Tephritidae) MS Rodriguero1, MT Vera2, E Rial4, J-P Cayol3 and JC Vilardi1,5 1Laboratorio de Gene´tica de Poblaciones, Depto. Ecologi´a, Gene´tica y Evolucio´n, Fac. Cs. Exactas y Naturales, Univ. Buenos Aires, (1428) Buenos Aires, Argentina; 2Instituto de Gene´tica, “Ewald Favret”, CNIA, INTA Castelar, CC25. (1712) Castelar, Argentina; 3International Atomic Energy Agency. Wagramerstrasse 5, PO Box 100. A-1400 Vienna. Austria; 4Fundacio´n Barrera Zoofitosanitaria Patago´nica, Argentina Males with a larger thorax and narrower face were found to indicated that TL and FW are probable targets of directional be favoured by directional sexual selection in a previous field sexual selection independently of female strain. However, cage experiment where Ceratitis capitata males from the laboratory females were less selective than wild ones. Seib 6–96 genetic sexing strain competed with wild males Additionally, correlational sexual selection was detected act- from Alto Valle (Patagonia) for the possession of wild ing on the multivariate phenotype. The effects of corre- females. Targets of sexual selection, however, might differ lational selection overlap with those of directional selection between wild and laboratory females as a response to adap- on each single trait. The analysis of mating pair character- tation to mass-rearing conditions. To evaluate possible istics showed patterns that do not match the expectations divergences on the targets of sexual selection as a by-pro- for a random mating system. The current analysis indicates duct of adaptation to mass-rearing conditions, field cage that during mating pair formation two processes overlap. On tests were performed with both wild and laboratory females. the one hand, sexual selection favours males with larger size To avoid possible bias due to correlation among the meas- (TL) and narrower faces (FW). This effect occurs in both wild ured traits (eye length [EL], face width [FW], head width and laboratory females. In addition, assortative mating [HW], and thorax length [TL]), a multivariate analysis was based on both phenotype and origin was also observed. applied. Consistent with the previous experiment, the results Heredity (2002) 89, 480–487. doi:10.1038/sj.hdy.6800170 Keywords: female choice; morphometric traits; genetic sexing strains; correlational selection; assortative mating; Ceratitis capitata Introduction be indirect, ie the direct target of selection might be one or several size-related traits. The mechanisms responsible Darwin (1871) used the term sexual selection to explain for consistent relationships between any particular trait the evolution of elaborate male characters that are appar- and sexual selection are not well understood, and might ently detrimental under natural selection. Two different be related with characteristics of the mating behaviour. kinds of evolutionary processes can account for the evol- The Mediterranean fruit fly, Ceratitis capitata, is a useful ution of such traits (Partridge and Halliday, 1984; Møller, model to analyse the relationship between size-related 1994): intrasexual selection or competition for mates traits and sexual selection. This species displays a lek between members of the same sex, usually males, and mating system (Prokopy and Hendrichs, 1979; Whittier intersexual selection, that involves active choice of parti- et al, 1992). Male aggregations occur on the bottom sur- cular individuals of the opposite sex, usually female face of leaves where they produce a pheromone call to choice of mates. These categories form a continuum attract females (Whittier et al, 1994). After females’ arrival (Boake et al, 1996) and it is not always easy to determine the male begins to perform a complex courtship involv- which form of selection is acting on a particular trait ing also visual and acoustical stimuli (Fe´ron, 1962; Webb (Partridge and Halliday, 1984; Møller, 1994; Whittier et et al, 1983; Calcagno et al, 1999, 2002; Eberhard, 2000). The al, 1994). common finding that mating success is highly variable In many species body size was often found to be posi- among lekking males has led to the predominant view tively correlated with male mating success (see Partridge that mate choice is an important component of sexual and Halliday, 1984). However, such an association might selection in lek-mating systems (eg, Whittier et al, 1994; Jones et al, 1998). However, many traits can be correlated with mating success without being necessarily mate- ´ Correspondence: Dr JC Vilardi, Depto. Ecologia, Gene´tica y Evolucio´n, choice targets (Norry et al, 1999). Besides, aggressive Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires. interactions between males or male activity levels to (1428) Buenos Aires, Argentina. E-mail: [email protected] 5Member of Carrera del Investigador Cienti´fico, CONICET, attempt copulation are probably intrasexually selected Argentina traits in many lek-mating species. However, C. capitata Received 4 January 2002; accepted 28 July 2002 males only weakly defend their territories, and there is Sexual selection on multivariate phenotype in medfly MS Rodriguero et al 481 no evidence of a relationship between territory location emergence (“IMCO 20” Co60 irradiator, minimum and mating success (Whittier et al, 1992, 1994). absorbed dose = 100 Gy, maximum absorbed dose = 150 Ceratitis capitata is a major agricultural pest managed Gy) and packed and sent to testing area, in northern through control programmes that involve the Sterile Argentina. Insect Technique (Knipling, 1955). The optimisation of Wild flies were collected from infested figs and peaches this method requires an understanding of the type of from Alto Valle Region, Patagonia, Argentina. Fruits col- mating system and a good knowledge of sexual selection lected in the field and backyards were taken to the lab mechanisms (Liedo Ferna´ndez, 1997). Laboratory and placed in trays on sand litter. Sand was checked (hereafter lab) strains may possess behavioural and periodically to collect wild pupae, which were then car- physiological traits that diverge from those required for ried to the testing area. Once at the testing site pupae mating success by wild populations (Cayol, 2000). Parti- were placed in flasks until emergence. Virgin adults were cularly, mass-reared females become receptive at a much aspirated from the flask within 24 h after emergence in earlier age (eg, Ro¨ssler, 1975; Wong and Nakahara, 1978) order to separate the sexes. Once sexed, the flies were and are less selective than wild ones (Zapien et al, 1983; kept in separate rooms until they reached sexual maturity Kaneshiro et al, 1990; Calkins, 1991). Additionally, at the age of 6–8 days for lab flies and 8–10 days for crowded conditions preclude the formation of leks wild flies. (Cayol, 2000). The process of adaptation to mass-rearing conditions may also result in genetic differences between Field cage tests lab and wild flies. Therefore, behavioural differences The test was carried out at Estacio´n Experimental Agroin- between lab and wild flies are the consequences of both dustrial Obispo Colombres, Tucuma´n, Argentina. Out- environmental and genetic factors (Eberhard, 2000), and door cylindrical field cages (2.0 m high and 2.9 m diam- in field conditions they may lead to a different ability to eter, Saran screen 20 by 20 mesh) (Calkins and Webb, compete for mating (sexual selection) and/or 1983) with a young lemon tree were used to score male assortative mating. mating success under a mass selection experiment. Indi- Mating success in C. capitata is determined by factors viduals from different strains were identified with the such as morphology (Churchill Stanland et al, 1986; Hunt addition of a water-based paint dot on their notothorax. et al, 1998; Norry et al, 1999; Orozco and Lo´pez, 1993; Each test consisted of the release of 30 wild males and Rodriguero et al, 2002), nutritional level (Blay and Yuval, 30 lab sterile males at dawn, about 7.00 am local time. 1997), and behaviour (Calcagno, 1999, 2002). To date, Half an hour later, 30 wild females and 30 lab sterile there is very little information, if any, about the occur- females were released into the cage. During a 7-h obser- rence of assortative mating based on morphology, even vation period, mating pairs (successful) were scored and though there is evidence of departures from random mat- gently removed with the aid of a vial from the cage as ing respect to strain (eg, Zapien et al, 1983; McInnis et they formed. Strain origin was determined for both male al, 1996). and female and couples were placed in the shade until Analysing video recorded courtships of lab and wild the end of copulation. Couples were individually lab- medflies Norry et al (1999) were able to detect intersexual elled. Males which were not able to copulate along the selection on morphometric traits. In a previous work test were labelled as ‘unsuccessful’. This is a hetero- under field cage conditions (Cayol et al, 1999) male com- geneous group involving individuals that were not able petitiveness and sexual compatibility between wild to integrate to any lek, moved from lek to lek or stayed medflies from Patagonia (Argentina) and a mass-rearing inside the same lek throughout the whole observation strain (Seib 6–96) were evaluated. In cages where wild period. They were collected altogether in a single vial. and lab males competed for mating with wild females only (unisexual tests) Rodriguero et al (2002) observed the Morphometric analyses occurrence of sexual selection on some size-related traits. From a total of 2520 flies, 191 successful couples and 189 In the present work we analysed sexual selection and unsuccessful males, were selected at random and meas- assortative mating in bisexual tests involving the same ured for four body size related traits: eye length (EL), face strains as in those by Cayol et al (1999) and Rodriguero width (FW), head width (HW), thorax length (TL) (Figure et al (2002).
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