Pacific Science (1989), vol.43, no. 2 © 1989 by University of HawaiiPress. Allrights reserved

Sexual Selection and Lek Behavior in the Mediterranean Fruit , capitata (Diptera: )!

LORNA H. ARITA 2 AND KENNETH Y. KANESHIR0 3

ABSTRACT: Field and laboratory studies were conducted to describe the complex system of the Mediterranean fruit fly, (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 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 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 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 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 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 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). The fruits were a section ofa host tree. In Kealakekua, Kona, placed onto the framed screen, which allowed host trees were coffee trees and in Kula, Maui, the mature larvae emerging from the fruits to the host trees were lemon and plum trees. fall through the mesh onto a layer of vermic­ Each male attempted to secure a territory ulite on the bottom of the plastic container. within the lek, which he utilized as a courting After 7 days, the vermiculite was sifted through site for receptive females cueing into this a O.I-mm mesh screen, and the collected pu­ communal display area. paria were placed into holding containers until The territories chosen by males both in the adult emergence. Within 24 hr of emergence, field and in the laboratory shared similar sex of the flies was determined and they were characteristics (Figure 1). A territory was placed into separate holding containers pro­ identified as a leaf of a host tree that was vided with food (honey, sugar, protein hy­ partially shaded by the overlying canopy of drolysate mixture) and water. The flies were the tree. Males were consistently observed then held until sexually mature (Arita 1982). occupying the portion ofthe leaves that was in direct sunlight. Another distinct feature of a Laboratory Observations ofMating Behavior territory was that only the underside of the leafconstituted the male's domain. Males reared out of the field-collected sub­ Depending on favorable environmental strates were placed into individual 3-gallon parameters, the lek begins to form in the (II.4-liter) aquariums when sexually mature morning at about 0900 hours. At that time, 138 PACIFIC SCIENCE, Volume 43, April 1989

FIGURE I. C. capitata male in pheromone calling position on a territory within the lek. one or two males arrive and occupy territo­ also be important in attracting females to the ries. Within the next 10-15 min, other males specific trees on which the leks are formed. begin to arrive at the lek. The number ofmales The effect of wind direction was also clearly that occupied territories within the leks varied evident in Kona where, because of the land­ from as few as 2 males to as many as 12 males sea breeze system, the wind blows onto shore in the field. This abrupt gathering of males at from the sea. Thus, the leks were always lo­ specific sites on the host tree leads to numer­ cated on the south side of the trees. ous encounters between the males as they The second parameter identified as impor­ jockey for territorial position. These males tant in lek formation was light intensity. As remained on their territories until about 1500 indicated above, a territory of a male was on hours, at which time they disbanded. the underside of partially shaded leaves, with There appeared to be two important envi­ the male occupying that portion ofthe leaf in ronmental parameters that influence lek for­ direct sunlight. The position of the male ap­ mation. The first parameter was wind direc­ to facilitate the detection of a potential tion. On Maui, the leks were always located mate arriving on the top surface ofthe leaf by on the host tree with reference to the trade­ the silhouette transmitted through the leaf. To winds. These tradewinds are prevalent 80 to detect this silhouette, the male must be sit­ 95% of the time, generally blowing from uated on the portion of the leaf receiving di­ the east-northeasterly direction (Blumenstock rect sunlight, since no silhouette is produced and Price 1972).The leks were usually located on the shaded portion of the leaf. It is pos­ upwind ofthe tradewinds so that the chemical tulated that the shaded portion ofthe leafmay cues (pheromones) being dispensed by the lek­ provide an important reference for light dif­ king males would be dispersed through the ferential that aids the male in perceiving a tree rather than away from the tree. This silhouette, thus reinforcing the notion that the orientation may serve not only to attract fe­ relative amount of sunlight penetrating the males to the lek site within the tree, but may leaf is important in selecting a territory. Lek Behavior of Ceratitis capitata-ARITA AND KANESHIRO 139

In addition to its importance in territory substrate and perpendicular to the longitudi­ selection, light was an important factor in the nal aspect ofthe body. The tibia is positioned formation and location of the leks. At one of perpendicular to the substrate. The midfemur the farms in Maui two leks were observed in is also parallel to the substrate and perpendic­ plum trees between 0900 and 1000 hours. ular to the longitudinal aspect ofhis body with Shortly after 1000 hours, the lek began to the tibia positioned such that the inner angle disband and within a period of 15 min from formed by the femur and tibia is 120°. The the disappearance of the last male from the third pair of legs are set in the same position plum trees, males began to arrive and form as the forelegs. The male remains in this three leks on a lemon tree located approxi­ pheromone calling position on the underside mately 20 m from the plum trees. Given that of the leaf until he perceives the presence ofa the lemon leaves were thicker than the plum potential mate as a silhouette through the leaf. leaves, it appeared that the males were relocat­ The male then initiates courtship actions (see ing leks based on the amount ofsunlight filter­ below) while orienting toward the approach­ ing through the leaves that served as territo­ ing individual on the top surface ofthe leaf. As ries. Light as an important factor in the loca­ the individual reaches the edge of the leaf, tion of the lek became even more evident in walks onto the bottom side of the leaf, and Kona, Hawaii. Although the leks were found approaches the courting male, the male must in the same coffee trees during the investiga­ discern whether the individual is a potential tion period, the location of the leks on the mate or another male who is challenging the trees moved relative to the position ofthe sun resident for territorial rights. Ifthe approach­ so that.the sunlight was directly on the leaves ingindividual is a receptive female, the male that served as territories. continues his courtship actions. If the ap­ The apparent influence of these two envi­ proaching individual is another male , how­ ronmental parameters in lek formation was ever, the resident initiates defense actions. consistent over the 2-yr observation period. The intruder male may approach the terri­ Leks were always located in positions that tory from any direction, landing within or appeared to optimize the amount of sunlight near the territory. Males were observed ap­ penetrating leaves that served as mating terri­ proaching directly onto the territory (bottom tories while referencing wind direction for the surface of the leaf) or approaching from the facilitation ofpheromone dispersal. top of the leaf and then walking to the lower A distinct series of actions is performed by surface. They were also observed hopping males in acquiring and maintaining territories from bottom surface to bottom surface of within a lek. Once a male occupies a territory, neighboring leaves onto the resident's terri­ he assumes a characteristic position referred tory. When a resident male perceives an in­ to as "pheromone calling" (Prokopy and truder male, there are two sets of actions that Hendrichs 1979). In this position, the male the resident can initiate. These defensive curls his abdomen upward and extrudes the actions have been termed (I) aggressive de­ terminal end of his rectal epithelium, which fense actions and (2) passive defense actions. forms a bubble-like structure presumably for the release of pheromone (Lloste and Roche AGGRESSIVE DEFENSE ACTIONS. Immediately 1960, Arita and Kaneshiro 1986). In addition, upon recognition ofan intruder male, the resi­ there are two pouches located in the pleural dent male terminates his calling position, region ofthe abdomen that are displayed only returns his abdomen to a more normal posi­ during pheromone calling and courtship tion in line with his thorax, and folds his wings (Arita and Kaneshiro 1983). The wings are over his abdomen. Then in a lunging motion, extended laterally during pheromone calling the resident rushes toward the intruder and with the under surface of the wings facing physically pushes the intruder with his head. anteriorly. The male's legs are set in a charac­ If the push is not met with an equal challenge teristic position during pheromone calling by the intruder, the resident will succeed in with the forefemur positioned parallel to the pushing the intruder off his territory. In addi- 140 PACIFIC SCIENCE, Volume 43, April 1989 tion to the "head butt," the resident brings his dicates that there is a distinct pattern during wings forward in a repetitive slashing motion this phase of the male's mating behavior. It in what appe ars to be even more forceful ac­ appears that wing vibrations may be facili­ tions against the intruder. The intruder can tating the dispersal of the pheromone toward also meet the resident's actions with similar the direction of the approaching stimulus actions of his own and indeed , on several oc­ (silhouette) on the top surface ofthe leaf. The casions, the intruder male was observed to resident male is constantly reorienting his po­ displace the resident. sition such that when the individual on the top of the leaf walks over the edge and onto the PASSIVE DEFENSE ACTIONS . The resident underside ofthe leaf, it will be facing the resi­ male initiates this type of defense in the same dent male head-on. Given that the wing vibra­ manner as the aggressive defense actions by tions are apparently facilitating the movement folding his wings over his abdomen. However, of pheromone toward the female, facing her as he approaches the intruder, rather than would allow for maximal reception of the attacking the intruder, the resident male will chemical stimulus. engage in a "face-off" where the two males If the individual that approaches the resi­ contact heads without any visible force and dent male is a female, a series of signals cu­ remain in this position for as long as 5 min. mulatively known as courtship will be ex­ Eventually, one ofthe two males will turn and changed between them. Courtship actions are leave the territory. As far as we have been able performed by the male upon the arrival ofthe to detect , there does not seem to be any distin­ female onto his territory and when the male guishable pattern as to which ofthe two kinds - has discerned that she is a female. During the of defense actions occur-during itiiy.one will entire period that the female is approaching, particular encounter. the male has his abdomen tucked under his body with the rectal epithelium still everted. His abdominal pouches are still inflated as he Courtship continues performing wing vibrations. Should There is a series ofactions that serve as the the female change direction while approach­ transitional phase between lek behavior and ing the male, the male will make a correspond­ courtship and are essentially the initial steps in ing directional change so that he is facing her. courtship. These actions are referred to as When the female is within 3-5 mm of the "orientation and fanning motions" (Arita and male, he begins a series of head movements, Kaneshiro 1983). During pheromone calling oscillating his head in both directions to a by the male from the underside of the leaf maximum plane of 30°. Within 1-2 sec of (territory), a visual stimulus produced by the initiating these head movements, the male ini­ silhouette of an individual on the top of the leaf tiates a second set of wing actions that are causes the male to immediately orient toward superimposed on the wing vibrations. While the direction of the stimulus. As the male continuing wing vibrations, the male simulta­ orients, he terminates the pheromone calling neously begins rhythmic wing fannings, bring­ position and tucks his abdomen under his ing his wings from a backward position with body with the rectal epithelium still everted the broad surface ofthe wings facing ventrally and begins to vibrate his wings. He continues to a forward position with the wings facing wing vibrations with his abdomen tucked anteriorly. He then moves closer to the fe­ under his body as long as the stimulus is pres­ male, who remains motionless except for ent. The abdominal pouches described ear­ some cleaning behavior. The male then leaps lier are inflated during wing vibrations and onto the female's back while still vibrating his appear to form a physical channel through wings and with his hindlegs clasps the female which the male can direct air currents carrying near the base of her wings. He then reorients the pheromone. Analysis of wing motion himself so that he is facing in the same direc­ using a variable frequency stroboscope (Arita tion as the female and copulation ensues. At and Kaneshiro, unpublished observations) in- any point, the female is able to reject the Lek Behavior of Ceratitis capitata-ARITA AND KANESHIRO 141 male's courtship advances simply by walking 1981) as well as in a variety of other away from the male and leaving the leaf. Or, if such as the wildebeest (Wilson 1975), fruit­ the male has already leaped onto her back in eating bats (Booth 1960, Bradbury 1975), and an attempt to mount, the female can still reject in endemic species of Hawaiian the male by releasing her hold on the leaf (Spieth 1968). From these studies, it is clear surface. that the lek mating system plays an important Although it has been suggested that acous­ role in regulating the effective reproductive tical signals may also be important in mating population. success in other tephritid species (see, for ex­ In laboratory studies of C. capitata, Arita ample, Webb et al. 1983) and although such and Kaneshiro (1985) showed that approxi­ signals may also playa role in the courtship of mately 85% ofthe "census population" (total C. capitata, we believe that they are of only number of males in the population) acquired secondary importance in this species. Our and defended territories and participated in observations indicate that males begin to the laboratory lek system. Thus, 15% of the vibrate their wings, during which time air­ census population was not considered as part borne sounds are probably being produced, of the viable reproductive community. They only after visual contact is made with the fe­ also showed that even among the males that male (or male) that has arrived on the top of participated in lek formation, only 70% (or the leaf (i.e., the silhouette of the individual; 60% ofthe census population) actually mated see discussion above). So that at the very least, with females that were introduced into the lek even if sounds are involved in the mating sys­ system. It is proposed that these males formed tem, they dQ D-ot seem to beimportant in the "effective male population." Further­ attracting females. Recently , Kuba and Sokei more, even among the effective male popula­ (1988) reported on the emission ofpheromone tion there was a certain percentage of males in the melon fly, Dacus cucurbitae Coquillett. (15%) that were more proficient maters. Earlier studies by Keiser et al. (1973)suggested These superior males accounted for nearly that wing vibration during courtship displays 46% of all matings. Because acquisition and in this species, as well as in several other Dacus successful defense ofa territory only provides species, serve as an acoustical signal impor­ a male with a courting site, the ability to per­ tant in mating success. It was postulated that form courtship actions to the satisfaction of sexually dimorphic structures on the wings the female is still a necessary component for and abdomen of males evolved as parts of a mating to occur in this species. In addition, it stridulating mechanism for sound produc­ has been observed that there are certain terri­ tion. However, using high-speed movie pho­ tories within the lek that are frequented more tography, Kuba and Sokei (1988) convinc­ often in comparison to others. Consequently, ingly showed that these structures are, in fact, it is on these territories that a majority ofmale used specifically for producing a "cloud" of battles occur . Ultimately, it is the male who pheromone that is dispersed toward the fe­ can secure one of these prized territories and male during courtship. These data provide who can perform the courtship ritual to the supporting evidence that, at least in some satisfaction of the female who will have a tephritid species, even if acoustical signals larger percentage of matings. These results are may have some biological significance in the strikingly similar to those of studies of other mating system of the species, they are only of species in which only a small percentage of secondary importance. We suggest this to be males are found to be responsible for most of the case in C. capitata. the matings that occur in the population. What of the remaining 15% of the census population that were unable to secure a terri­ tory within the lek? These males may be prime DISCUSSION candidates for adoption of the alternate mat­ Lek behavior has been well studied in bird ing strategy of approaching females at ovi­ species (e.g., Wiley 1973, Lill 1974, LeCro y positional areas that has been proposed for C. 142 PACIFIC SCIENCE, Volume 43, April 1989

capitata by Prokopy and Hendrichs (1979) LITERATURE CITED and by Burk (1983) for the Caribbean fruit fly. Anastrepha suspensa (Loew). ARITA, L. H. 1982. Reproductive and sexual Clearly, lek systems represent an intensifi­ maturity in the Mediterranean fruit fly, cation of the sexual selection process. Leks Ceratitis capitata (Wiedemann). Proc. seem to serve as a giant " sieve" through which Hawaii. Entomol. Soc. 24(1):25-29. males are assayed for their genetic quality via ARITA, L. H., and K. Y. KANESHIRO. 1983. sexual selection. Although intrasexual selec­ Pseudomale courtship behavior of the fe­ tio~ (i.e., male-mal.einteraction) seems to play male Mediterranean fruit fly, Ceratitis capi­ tata (Wiedemann). Proc. Hawaii. Entomol. an Important role In "sorting out" the "best" males in the population through the establish­ Soc. 24(2-3): 205-210. ment of a dominance hierarchy (Arita and - - -. 1985. The dynamics ofthe lek system Kane~hiro 1985), female choice (i.e., epigamic and mating success in males ofthe Mediter­ s~lectIon) based on courtship ability still pro­ ranean fruit fly, Ceratitis capitata (Wiede­ vides the ultimate criterion for mating success. mann). Proc. Hawaii. Entomol. Soc. 25: The importance offemale choice in tephrit­ 39-47. ids has apparently led to an array of court­ - - -. 1986. Structure and function of the s?ip rituals that are performed by males, par­ rect~1 epithel~um. and anal glands during ticularly among lekking species. In Dacus mating behavior In the Mediterranean fruit tryoni (Froggatt), though males actively de­ fly male. Proc. Hawaii. Entomol. Soc. 26: fended territories and produced acoustical 27-30. and chemical signals .for.the attraction of fe­ BACK, E. A., and C. E. PEMBERTON. 1918. The males, mating occurred soon after the arrival - Mediterranean fruit fly in Hawaii. U.S. of the female, with little or no courtship dis­ Dep. Agric. Bull. 536. play by the male before copulation (Tychesen BLUMENSTOCK, D. I. , and S. PRICE. 1972. Cli­ 1977).On the other hand, A. suspensa appears mates ofthe states: Hawaii. Pages 155-204 to have a mating behavior strikingly similar to in E. A. Kay, ed. A natural history of the that of C. capitata. Males of A. suspensa not Hawaiian Islands. University of Hawaii, only defend territories and produce an air­ Honolulu. borne sex pheromone for attracting females BOOTH, A. H. 1960. Small mammals of West but for successful mating to occur, females are . Longmans, Green, London. a.dditionally stimulated through courtship ac­ BRADBURY, J. 1977. Social organization and tions (Burk 1983). These studies on closely comI?unication. Pages 1-73 in W. Wimsatt, related groups of lekking species provide the ed. BIOlogy ofbats. Vol. 3. Academic Press foundation for understanding the role of the New York. ' lek system and courtship in mating success. BRADBURY, J., and R. GIBSON. 1983. Leks and We are continuing our studies to determine mate choice. Pages 109-138 in P. Bateson, which components are of critical importance ed. Mate choice. Cambridge University in mating success. Press, Cambridge. BURK, T. 1983. Caribfly studies aimed at im­ proving medfly control. Citrograph 68(4): 73,86. ACKNOWLEDGMENTS BURK,.T., and s:. O. CALKINS. 1983. Medfly matmg behavior and control strategies. Fla . We thank Ted Burk for reviewing early Entomol. 66: 3-18. drafts of the manuscript and Arlene Mihara CAUSSE, R., and M. FERON. 1967. Influence for manuscript preparation. We also express du rhythme photoperiodque sur l'activite our appreciation to the John Hashimoto and sexuelle do la mouche mediterraneene des Earl Fujitani families in Kula Maui for fru.its: Ceratitis. capitata (Wiedemann) allowing us to conduct field studies on 'their (Diptera Trypetidae). Ann. Epiphyt. 18: farms. 175-192. Lek Behavior of Ceratitis capitata-ARITA AND KANESHIRO 143

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