George Gamboa and John Alcock.
The Mating Behavior of Brochymena quadrapustulata (Fabricius) (Heraiptera, Pentatomidae).
Psyche 80:265-269, 1973.

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PSYCHE
Vol. 80 December, I 973 No. 4
THE MATING BEHAVIOR OF
BROCtIYMENA Q lJAIIRAPlJSTlJLArTA ( FAHRICIUS)* ~ 3 3 ~ GEORGE GAMBOA AND JOHN ALCOCK
Department of Zoology,
Arizona State University,
Tempe, Arizona 85281
l'entato~nid reproductive behavior has been the subject of a number of papers (e.g. I<ullenberg, 1947 ; Teysovsky, I 949; Southwood and Hine,
1950; Leston, 1955 ; Kaufmann, 1966 ; IIitchell and illa~i~ 1969; Tostowaryk, 1971 ; Alcock,
1971 ; Fish and Alcock, 1973).
These studies have revealed considerable diversity and complexity in the courtship activities of male pentatomids, raismg questions about the evolution and ecoIogica1 significance of these behaviors. Answei-s to these questions will require additional comparative data. We present infornlation here on Brochymeno quodra~ustuiata; loose aggregations of this cryptic species were obsei\ed on grapefruit (Citrus parrzdisi) and desert bsoon~ (Bmchoris sar?throides) in suburban Tempe, Arizona, from 25 February to 17 Ifay 1973. ln addition to written records of field observations of eight courtship, super-8 films of three separate courtships leading to copulation were utilized for detailed analysis of the mating behavior of B. quodra- RESULTS
Eight complete and three incomplete courtships were observed between I 145 and 1540 hrs. The components of mating are out- lined chronologically below and illustrated in Fig. I. (I) The male appioaches the female (she may be ~noving 01- in- nob bile at the he) and touches her with his antennae. If
moving, the female may freeze with her abdomen held close to the branch on which she was walking or escape bl running away.
*Manziscript receiw8d by ~/IL tditor Octobet 9, 1973



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266 Psyche [December
Figure 1. A diagram of the courtship of Brochymma quadrapustulata. A. The male's crab-like movements on the anterior dorsum of the female. B. The male moves to the rear of the female. C. The view from above and
from the side of the male antennating the venter of the female's abdomen, R. The view from above and from the side of the male about to achieve genital linkage.
E. Genital linkage in an end-to-end position. F. The male drumming on the side of the female with its hindlegs.



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The male seizes the anterior or posterior dorsum of the female with his legs. Unreceptive females respond by breaking away from the male or by moving off with their partner clinging to them.
If the female is immobile, the male moves so as to face his mate while continuing to grasp the dorsum of her body. He then begins a rapid crablike walk from side to side over the anterior female dorsum. Receptive females gradually raise their abdomen in response to this activity. The male's back and forth movements varied from 20-34 in number and lasted from 23-43 seconds in four filmed courtships.
At some point the male moves off the anterior dorsum of the female and continues out along the side of his mate while con- stantly antennating her lateral surface. The male, upon reaching the posterior of the female, sweeps his head under her elevated abdomen; his antennae move rapidly up and down in alternating strokes touching the female's abdominal venter. The male may prod and lift the body of unresponsive females which have not voluntarily raised their abdomens.
As the male's head passes under the body of the female he begins a tight 180å turn that brings him into the end-to-end precopulatory position. Unreceptive females may dash off down a branch as the male executes this maneuver. The male elevates his abdomen while sweeping the aedeagus in a zig-zag pattern against the venter of the female7s abdomen. Each sweep brings the male aedeagus pi-ogressively closer to the female genitalia; linkage occurs when mutual genital con- tact occurs.
Upon linkage, the insects' bodies jerk violently from side to side for several minutes. The coupled pair may move a short distance during this time.
The male rapidly drums the sides of the female's abdomen with his hindlegs (also abserved by Ruckes, 1938, for B. sdcata). Spells of drumming are interrupted by pauses during which the male rests its hind legs on the dorsun~ of the female's abdomen. This continues as long as the partners are linked. One pair of stinkbugs remained in copula for at least 75 rnin but 11ad sepa- rated when observed 45 rnin later.
'I'his report provides additional evidence that ~ N I I ~ the Penta-



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tomidae it may not be w~commn for males to initiate cop~dation while facing directly away from the fen~ale. ?'his behavior is rare among the Heteroptera (Weber) 1930).
Below we have summarized
the major methods by which pentatomids achieve genital linkage. Genital linkage initiated in an end-to-end position Brochynzena (this paper) ) Euschistus (Alcock, I971 ) , Nezartz (Mitchell and Maul 1969)) ChZorochroa and CosnzopepZa (Fish and Alcock, 1973 ) ) Perillus (Esselbaugh, I 9481, Podisus (Olsen) 1910)
Genital linkage initiated with male above fernale, both facing in same direction
Bmchymena (Ruckes) 1938)) Ddycoris (Teyrovsky, 1949) ) CaZidca ( Kaufmann, I 966)
Genital linkage initiated with female above male, both facing in .same dir-ec tion
Po&sus (Tostowaryk, I 97 I )
\\7e. also have records of end-to-end rnatings ill Y1hy(~nt({ pdMo-
virms. Xlales of this species, upon ~naking contact with a female,
begin antennating the surface of her body while moving to the tip of her abdomen. There they may prod and lift the female's body with their head before turning away from her. The female) if receptive, raises her abdomen slightly.
Unlike other species ~vhicl~
initiate copulation in a dismounted position) the male's body often forms a right angle with the female instead of a 180å angle. 7'he male then kicks lightly at the rear of its partner's wingcovess and abdomen with its hind legs before inserting the aedeagus into the female's genital opening.
As Fish and Alcock ( 1973) have noted, species which employ t l ~ end-to-end method have highly similar courtship routines. Commo~~ characteristics include ( I) male antennation of the female, (2) at- tempts by the male to lift the female's abdomen with its head, (3) abdominal elevation by receptive females, and (4) tactile stim- ulation of the venter of the fen~ale's abdomen with the antennae and aedeagus of the male. The male's "goal" in courtship appears to be to induce the female to adopt a position that will make insertio~~ of the aedeagus relatively easy.
? 7
1 he unusual feature of the courtship of B. quadrapu~tzdata is the rapid crab-like movement of the male over the head and thorax of the female, a behavior that may have evolved from efforts of males in the past to prevent physically the escape of females. Now the action may serve as a releaser of abdo~ninal elevation by receptive



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19731 Gam boa & AZcock - Brochymena 269
females. Interestingly, very similar behavior has been reported for the distantly related African pentatomid Calidea dregii, a member ,of the Scutellerini (Kaufmann, I 966).
Finally the fact that different members of the same genus (e.g. Brochymena, Podisus) may exhibit basically different methods of initiating genital linkage suggests that this component of reproduc- tive behavior is evolutionarily labile. The reasons for evolutionary changes of this sort remain obscure.
We thank Elinor Lehto for the identification of the plants men- tioned in the paper. Alan R. Hardy of the California Department of Agriculture provided the identifications of Brochymena and Thyanta for which we are grateful. This research was supported in part by NSF grant GB-35269.
ALCOCK, J.
1971. The behavior of a stinkbug, Euschistus cons~ersus Uhler (Hemip- tera, Pentatomidae). Psyche 78 : 215-228. ESSELBAUGH? C. 0.
1948. Notes on the bionomics of some midwestern Pentatomidae, En- tomol. Americana 28: 1-73.
FISH, J. AND J. ALCOCK
1973. The behavior of Chlorochroa ligata (Say) and Cosmopepla bimaculata (Thomas) (fiemiptera, Pentatomidae) . In press, Entomol. News.
KAUFMANN, T.
1966. Notes on the life history and morphology of Calidea dregii (Hemiptera : Pentatomidae : Scutellerini) in Ghana? West Africa. Ann. Entomol. SOC. Amer. 59: 654-659.
KULLENBERG, B.
1947. Uber Morphologie und Funktion des Kopulationsapparats der Capsidm und Nabiden. 2001. Bidrag Fran Uppsala 24: 217-418. LESTON, D.
1955. The function of the conjunctiva in copulation of a shieldbug? Peizodorus lituratus (Fabricius) (Hemiptera, Pentatomidae) . J. SOC. Brit. Entomol. 5: 101-105.
MITCHELL? W. C. AND R. F. L. MAU
1969. Sexual activity and longevity of the southern green stinkbug Nezara viridula. Ann. Entomol, SOC. Amer. 62: 1246-1247. OLSEN, C. E.
1910.
Notes on breeding Hemiptera. J. N. Y. Entomol. Soc. 18: 39-42. RUCKES? H.
1938. Courtship and copulation in Brochymena sulcata Van D. Bull. Brook. Entomol. SOC. 3 3 : 89-90.




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270 Psyche
[December
SOUTHWOOD, T. R. E. AND D. J. HINE
1950. Further notes on the bio'logy of Sehirus bicolor (L.). Entomol. Mon. Mag. 86: 299-301.
TEYROVSKY, V.
1949. Praeconnubia and courtship in terrestrial bugs. Acta Acad. Nat. Hist. Moravo-Silesiacae, Brno 21 : 1-16. TOSTOWARYK, W.
1.971. Life history" and behavior of Podk modestus (Hemiptera: Pentatomidae) in a boreal forest in Quebec. Can. Entomol. 103: 662-674.
WEBER, H.
1930. Biologic der Hemipteren. Julius Springer, Berlin. PATTERNS OF ABDOMINAL FUSIONS IN MALE BOREUS (MECOP- TERA). - Since publication of my comments on the fusion of terminal tergal and sternal abdominal plates in male Boreus (Psyche, 79: 277,
1972), I have examined two males of Boreus vlasovi Martynova (determined + by Dr. Martynova) from Ashkhabad, Turkmeniya, U. S. S. R., through the kindness of Prof. I?. M. Carpenter. Contrary to my listing based on the original description, these males do not have the 9th tergum fused to its sternum. Both have their 8th and 9th terga and sterna free, namely (o,o), just as in the North American Boreus brevicaudus Byers, B. brumalis Fitch, B. nivoriundus Fitch, and B. notoperates Cooper. Byer's figure (Ann. Ent. Soc. Amer. 47: 491, fig. 15, 1954) shows the North berican B. reductus Carpenter also to be (o,o). - K. W. COOPER, Dept. of Biology, University of California, Riverside, Calif., 92502.



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