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Sexual Behavior of Acanthocephala Declivis Guatemalana (Hemiptera: Coreidae) and the Allometric Scaling of Their Modified Hind Legs

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Sexual Behavior of Acanthocephala Declivis Guatemalana (Hemiptera: Coreidae) and the Allometric Scaling of Their Modified Hind Legs BEHAVIOR Sexual Behavior of Acanthocephala declivis guatemalana (Hemiptera: Coreidae) and the Allometric Scaling of their Modified Hind Legs WILLIAM G. EBERHARD Smithsonian Tropical Research Institute, and Escuela de Biologfa, Universidad de Costa Rica, Ciudad Universitaria, Costa Rica Ann. Entomol. Soc. Am. 91(6): 863-871 (1998) ABSTRACT The sexually dimorphic hind legs of Acanthocephala declivis guatemalana are used for different functions in males and females, and they scale differently on body size. The hind legs of males are used to deliver powerful squeezes to other males in battles at feeding and mating sites, and the diameter of the hind femur and the length of its large tooth are proportionally larger in large males compared with other body characters. The largest tooth on the female hind femur probably functions in the less forceful female-female battles, and it is also proportionally larger in large females. However, neither the female hind tibial plate, which is used as an aggressive signaling device, nor the diameter of the female hind femur are proportionally larger in large females. Wing wear and breakage of hind leg spines, combined with behavioral observations of marked bugs, indicate that the breeding system combines elements of resource defense polygyny and lek polygyny. Females mate more than once, despite the attendant cost of reduced feeding opportunities. KEY WORDS Acanthocephala, Coreidae, sexual behavior, weapons, visual display traits IT IS GENERALLY thought that both visual display struc- Materials and Methods tures and weapons under sexual selection have high All observations were made between 24 and 31 July allometric values (slopes of >1.0 in log-log regressions 1997 at the Chamela field station of the Universidad on indicators of body size) (Huxley 1972; Alatalo 1988; Autonoma de Mexico in Jalisco, Mexico (elevation =»50 Petrie 1988,1992; Green 1992; Burkhardt et al. 1994). m). Bugs were observed on the trunks and branches of The data supporting this idea are relatively sparse, young (=«20-30 cm diameter at breast height [dbh]) however (see summary in Eberhard et al. 1998, also Cordiaelaeagroidei (Borraginaceae) trees planted on the Wilkinson 1993, Goddard and Mathis 1997, Wilkinson station grounds. Bugs were given individually recognize- and Dodson 1997). In the case of visual display char- able marks on the pronotum with dots of colored fast- acters, the observations are largely limited to verte- drying paint while they rested unrestrained on the lower brates (mostly birds). It is thus of general interest to portions of trees. The trees were 10-15 m tall and bugs determine the allometric values of display structures sighted in the upper portions could not be checked for and weapons in other animal groups. marks (and sometimes could not be sexed). A. declivis is Many coreid bugs have modified hind legs that are large (body length «*3 cm) and moves relatively slowly thought to function in interspecific interactions. The and tolerates slow approaches of an observer to within most typical modification is an expansion of the hind ^30 cm. It was thus possible to observe many behavioral tibia into a more or less flattened and often brightly details in the field. colored plate or flag. The hind femora are also some- Surveys of bugs present on 4 adjacent trees (*=«5 m times thickened and adorned with spines (Schuh and apart) were made at irregular intervals separated by at Slater 1995) and serve as weapons in male-male com- least 2 h. There were 15-20 other trees on the station bat (Fujisaki 1980,1981; Mitchell 1980; Miyatake 1993, grounds; only 5 of these other trees had any bugs on 1995,1997). Males and females of some Acanthoceph- them during the study. The populations on the 4 study ala Lap. offer an interesting chance to study these trees were much larger than those on any other tree types of modifications: females have prominent tibial at the station. plates but relatively thin, toothed femora; whereas All measurements (Fig. 1) were of bugs collected at males have swollen hind femora with prominent teeth, the station, and include both specimens in the collec- but reduced tibial plates. The current study describes tion at the station and additional voucher specimens. the sexual behavior of Acanthocephala declivis guate- The ages of specimens were estimated according to malana Distant as it relates to hind leg use, and the the amount of forewing membrane torn by making a scaling of hind leg structures on body size. drawing of the outline of each front wing's margin and 864 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 91, no. 6 wear on the teeth on each hind femur was estimated by counting the numbers of teeth in which (1) the extreme tip was broken off, (2) a more sizeable por- tion of the tip was gone, or (3) at least half of the tooth was gone. A 3rd estimate of wear involved the per- centage of the small spines at the tips of the tibiae that were at least half missing. Four counts were made for each animal (prolateral and retrolateral views of each of the 2 tibiae), and the average of these 4 percentages Female was used as the hind tibia spine value. Results Behavior. Bugs were found only on the trunks and branches of the C. elaeagroidei trees or on leaves and branches (but not trunks) of nearby vegetation. Sex- ual activity occurred only on C. elaeagroidei. Copu- lating pairs were noted at sunup (0730 hours), during the day, and up to 3 h after sundown (at 2400 hours). Fig. 1. Hind legs of a male and a female A. declivis, showing sexual dimorphism and the portions that were measured. Two copulations were observed to begin at 2230 hours. Copulation lasted up to ^90 min, so mating (and probably also fighting and feeding) continued rips in the membrane, labeling the backs of the draw- throughout the day and night. There was no obvious ings with specimen numbers, cutting out and then tendency for copulation to be especially common at mixing all the drawings, and then sorting them into any particular time of day. categories of wing damage (Fig. 2). The mean of the Both males and females inserted their mouthparts values for the 2 wings was calculated for each speci- for up to 30 min into the trunk of C. elaeagroidei trees. men. The estimates of forewing damage were thus In some cases a 1- to 2-mm-diameter circular area of obtained blind with respect to other traits. A femoral the bark around the point of insertion became wet spine wear value was also assigned to each hind leg, (Fig. 3), and often ants and occasionally vespid wasps and these were combined for each specimen. The clustered around such a feeding site (both while the Fig. 2. Typical representatives of the 6 categories of wing damage of male A. declivis (black portions were missing). November 1998 EBERHARD: SEXUAL BEHAVIOR OF Acanthocephala declivis guatemalana 865 Females that had been resighted once were also less likely to remain for a further day after being resighted than were males. None of the 4 females was resighted on any further days, whereas 11 of the 17 resighted males were sighted at least 1 additional day (P = 0.0351, Fisher exact test). Summing all sightings of bugs during 28 surveys on 7 d, there were 659 sightings of males, and 552 of females on the 4 study trees (an additional 99 sightings involved bugs whose sex could not be determined because they were too high in the trees). Assuming that the bugs whose sex was not determined were a random sample of males of and females, the ratio of males to females was 1.19:1.0. Of the males and females sighted 31.0 ± 11.5% and 36.7 ± 10.4%, respectively, were copulating. The distribution of bugs on the study trees was strongly clumped. Thus, on 24 July there were at least 22 on 1 tree, and only 2 and 3 on the 2 adjacent trees (/ = 32.7, df = 2, P < 0.001). Relative population sizes on different trees varied over time. For instance, 4 d later the same 3 trees had 6, 23 and 1 bug respectively (G = 31.2, df = 2, P < 0.001 with respect to 1st counts). Courtship and Copulation. In 6 cases I saw a male perform series of brief (M3.5- to 1.0-s) bouts of quivering or body vibration while standing alongside but not touching a female, or facing her just before attempting to Fig. 3. A feeding female A. declivis raises her near hind mount. Bouts of vibration occurred both in rapid suc- leg in the warning posture as a male approaches her (drawn cession, and up to >30 s apart. In other cases a male from a photograph). mounted without any obvious preliminaries. Females often resisted mounting, by raising their hind legs, at- bug's proboscis was inserted and afterward). Several tempting to push the male away, or walking away. One "wet" feeding sites were used repeatedly by several male succeeded in squeezing past the female's raised different bugs over several days. Some feeding sites hind leg and onto her dorsum, but in 17 other cases in were at apparently undamaged sites of the trunk, but which female resistance was observed carefully, she suc- most occurred at fissures in the bark, or where the ceeded in leaving a male who attempted to mount her. outer bark had been recently scraped away. The bark Males sometimes pursued retreating females for up to of C. elaeagroidei is very thin, with <1 mm of outer 20-50 cm along the trunk, but none of these mounting layer covering the green inner layer which is also <1 attempts was successful.
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