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Ceratitis Capitata (Diptera: Tephritidae)! Pacific Science (1989), vol.43, no. 2 © 1989 by University of HawaiiPress. Allrights reserved Sexual Selection and Lek Behavior in the Mediterranean Fruit Fly, Ceratitis capitata (Diptera: Tephritidae)! LORNA H. ARITA 2 AND KENNETH Y. KANESHIR0 3 ABSTRACT: Field and laboratory studies were conducted to describe the complex lek mating system of the Mediterranean fruit fly, Ceratitis capitata (Wiedemann). Results of these studies indicate that there are two phases in the mating system: (I) lek behavior and (2) courtship. In the initial lek behavior phase, males acquire and defend territories that are used as courting sites for receptive females. An aggregation of these territories that form a communal display area is known as a "lek" and once a female enters the lek and arrives at the territory of a male, a series ofcourtship actions are reciprocated between the pair that may result in copulation. Descriptions of other environmental parameters that appear to be important features of the lek system are also presented. SEXUAL REPRODUCTION is one of the most territories frequented by receptive females, importantfeaturesin thelife history of most resulting in the evolution .of sexually die higher organisms. The processes that result in morphic characteristics that would give better the bringing together of individuals of the endowed individuals a higher success rate opposite sex and the subsequent mating of in defending such preferred territories. In these individuals are essential to the perpetua­ epigamic selection, the performance ofcourt­ tion of the species. In polygamous popula­ ship displays by the male (which may involve tions, there is often differential mating success visual, chemical, or acoustical signals) must among the males. This differential mating be able to satisfy the mating requirement of success is a result of the sexual selection pro­ the female. Those males whose courtship cess, which ensures that individuals whose ge­ actions are performed in the "right" sequence netic combination results in high Darwinian and temporal pattern will satisfy the mating fitness are those that will participate in the requirements of a higher number of females mating community. in the population, resulting in differential There are two aspects to sexual selection: mating success among the males. Also, any (1) intrasexual selection, which involves morphological embellishment that provides competition among individuals of the same males with an advantage over othermales that sex (usually males), and (2) epigamic selec­ lack such features will evolve as part of the tion, which involves interaction between intraspecific mate recognition system of the individuals of the opposite sex (see Thornhill population. and Alcock [1983] for a general review of this The lek mating system, which has been topic). In the former , males may form mating observed in a .wide variety of organisms (Bradbury and Gibson 1983), involves the for­ mation of an aggregation ofterritories called a I Financial support for this study was partially lek. In most lek systems, each territory within provided by the USDA Cooperative Agreement No. 58-9AHZ-1-627, USDA Competitive Grant No. GAM­ a lek is occupied by a single male to which 8600216, and a grant from the Threshold Foundation. receptive females are attracted for the sole Manuscript accepted August 1988. purpose of mating. Males compete for posses­ 2 University of Hawaii at Hi10,College of Agriculture, sion of these territories and in some cases, 523 West Lanikaula Street, Hilo, Hawaii 96720. 3 University of Hawaii at Manoa, Hawaiian Evolu­ "preferred" territories have been observed , as tionary Biology Program, 3050 Maile Way, Honolulu, evidenced by the number of males competing Hawaii 96822. for the territory as well as the number of fe- 135 136 PACIFIC SCIENCE, Volume 43, April 1989 males frequenting it. However, in addition to of C. capitata focused mainly on courtship defending and occupying a territory that pro­ (Martelli 1910, Back and Pemberton 1918). A vides a male with the opportunity to encounter more complete description by Feron (1962) receptive females, males must also be able to separated the mating behavior of C. capitata perform precise courtship displays to satisfy into three stages, which he termed: (1) the the receptivity threshold of those females. awaiting period, (2) the arrival of the female, Thus, the dynamics of lek systems represents and (3) assault and coupling. However, it was a highly intense form of sexual selection in not until the work ofProkopy and Hendrichs natural populations where both intrasexual (1979) and Arita and Kaneshiro (1983, 1985) and epigamic selection contribute to mating that the lek system was proposed as an in­ success. tegral part of the mating behavior of this The Mediterranean fruit fly, Ceratitis species. Other authors have researched the capitata (Wiedemann), is considered one of role of pheromones (Ohinata et al. 1972, the most damaging insect pests because ofthe Jacobsen et al. 1972, McDonald 1987), audio wide variety offruits and vegetables that serve stimulation (Rolli 1976, Webb et al. 1983), as hosts for the developing larvae. During the and environmental parameters (Myburgh past three decades, efforts to control or eradi­ 1962,Causse and Feron 1967)as contributing cate C. capitata have been initiated by many factors in courtship. However, though these countries hoping to increase their fruit exports references present a general scheme of the as well as curb their current fruit losses. One mating behavior of C. capitata, Burk and of the major control methods that has been Calkins (1983) concluded that much more in­ applied to C. capitata is the Sterile Insect formation about its mating behavior was Release Method (SIRM). This technique, essential. which was first proposed by Knipling (1955), In this paper, we describe the mating be­ utilizes sterilized laboratory-reared insects havior of C. capitata based on field and labo­ that are released into the field to control wild ratory observations. Such a description pro­ population levels, leading to the eradication vides a basis for subsequent research in this of the pest from an area. The application of area, especially in our understanding of the this technique was initially successful when dynamics of the sexual selection process and used for the control of the screwworm, its influence on regulating effective popula­ Cochliomyia hominivorax (Coquerel). How­ tion size. We hope that our results will ulti­ ever, problems with the mating competitive­ mately contribute toward the development ness ofthe laboratory stock of C. hominivorax of more effective control programs for this greatly diminished its effectiveness against species. wild populations in subsequent control at­ tempts (Richardson et al. 1982). The relatively frequent infestations of C. capitata in parts of southern California em­ MATERIALS AND METHODS phasize the need for a more comprehensive understanding of the behavioral ecology of Field Observations this pest so that more efficient control pro­ Preliminary field studies were conducted in grams -can be developed. To avoid setbacks various localities within the Hawaiian Islands. similar to those experienced in the screw­ From these studies, two sites were chosen for worm programs, the genetic quality of the more in-depth observations of the mating laboratory stock of C. capitata must be re­ behavior of C. capitata. The areas chosen established and maintained. Yet, one area of were (1) two residential farms in Kula, Maui the ecology of this fly that is least understood and (2) a coffee plantation in Kona, Hawaii. but of vital importance to control programs, especially those involving the SIRM, is the KULA , MAUL Over a 2-yr period, 10 field mating behavior. trips (2-3 days each) were made to the two Earlier descriptions of the mating behavior farms. Observations of C. capitata mating Lek Behaviorof Ceratitis capitata-ARITA AND KANESHIRO 137 behavior activity at each of the farms were and allowed a 24-hr acclimation period to the made throughout each day of the field trips. cage conditions. On the following morning, a female (reared from the same substrate) was KONA, HAWAII. Over the same 2-yr period, placed into the cage and all courtship en­ five trips (2-3 days each) were made to a counters between the pair were recorded onto coffee plantation on the Kona coast above 3/4-in. (l9-mm) videotape with a JVe camera Kealakekua Bay. A portion of the plantation and a Sony recorder. This procedure was re­ was chosen as the study site. We surveyed a peated for 18 pairs reared from coffee and 28 plot that contained 53 coffee trees. Each tree pairs reared from Jerusalem cherry. The tapes was individually identified by row and col­ were later reviewed and analyzed for mating umn. Throughout each day, each tree within behavior actions. the transect was monitored for mating behav­ ior activity. RESULTS Rearing Methods of Wild Flies for Laboratory Observations Previous authors such as Feron (1962) divide the mating behavior of C. capitata Flies used for this research were reared into different stages. However, the results of from two different host plants collected on our field and laboratory observations of the island of Hawaii. One population of flies C. capitata activity indicate that the mating was reared from Jerusalem cherry, Solanum behavior is separated into two basic phases pseudocapsicum L., collected from Kipuka Ki, that lead to copulation: (1) Jek behavior and Volcanoes National Park. The other popula­ (2) courtship. Each phase is described below. tion was reared from Arabian coffee, Coffea arabica L., collected from Kona, Hawaii. Infested fruits from these two areas were Lek Behavior brought into the laboratory, set into rearing The first phase in the mating behavior of C. containers, and maintained in separate rooms capitata involves the acquisition ofa territory at a temperature of 22°e. The rearing con­ by the male. In C. capitata, aggregations of tainers consisted of a wooden-framed screen these territories, which are referred to as leks (26 x 37 em) placed 10 em deep inside a plastic or lek sites (Wilson 1975), were formed within container (IS x 50 x 32 ern).
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