Robert H. Barth, Jr.
The Mating Behavior of Gromphadorphina potentosa (Schaum) (Blattaria, Blaberoidea, Blaberidae; Oxyhaloinae): an Anomalous Pattern for a Cockroach.
Psyche 75:124-131, 1968.

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19681 Earth -Behavior of Gromphadorhina 127 within a few seconds, and the pair remained in the opposed position throughout the period of copulation.
The activities of the first courting pair aroused the other males sufficiently so that only brief contact with a female was necessary for release of the walking and hissing activities of the male. During this period of heightened activity, a number of males walked about hissing even though not in contact with females. If such a male did contact a female he immediately attempted to copulate. Females
usually fled from such males.
Observations on four successful copulations suggest that several seconds may normally be required to achieve a satisfactory genital connection. During copulation attempts, the female may be pushed forward several inches by the male's vigorous backing movements. Copulating pairs remain quiescent even though other animals may crawl over them. Males wave their antennae quite actively during copulation. Females wave theirs much less actively. In one case a copulating pair was disturbed by the observer after about 15 min- utes and the female ran off dragging the male behind her as in other cockroach species, even though in this case the male was much the heavier of the two. Copulation last a remarka'bly short time for so large a species, no more than twenty to thirty minutes (data from three copulations). These females were found to contain spermato- phores at the termination of copulation so it may be concluded that these were normal copulations,
All successful copulations were preceded by long periods of gentle mutual antennal fencing and body stroking. Thus it seems highly probable that tactile stimulation resulting from antennal contact with a female is the normal releaser of the male's courting activities in quiescent males, and that antennal fencing promotes sexual re- ceptivity in females. No male-male courtship sequences were ob- served. It seems quite possible that differences in the type of tactile stimulation resulting from antennal fencing between two individuals may alone be sufficient to determine subsequent behavior. As we have seen, the vigorous antennal fencing following male-male contacts is invariably followed by aggressive behavior, whereas the more gentle antennal fencing following male-female contacts is followed by court- ship behavior. The long-laterally projecting sensory hairs on the antennae of the male may indeed be tactile receptors specialized for precisely this purpose, i.e., the detection of different intensities of tactile stimulation. This hypothesis might be tested by subjecting males to various types of artificial tactile stimulation. Whether con- tact chemoreception plays a role in sex recognition is unknown.



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I 28 Psyche [June
M ATI N G BE H AV I 0 R of GROMPHADORH/NA P ORTENTOSA ? BEHAVIOR ' RELEASER CT BEHAVIOR
MOTIONLESS
~~ctile Slim- TOUCHES 9 with
ANTENNAL / ANTENNAE
FENCING -
(if receptive)
ANTENNAL
~oclile.Sliffi---~ FENCING
ANTENNAL
FENCING +
STROKING of
d 5 BODY
WALKING AROUND 9
with HISSING
d a 9 ABDOMINAL TIPS OPPOSED
^'^'SliffTÌÔ- BACKING +
COPULATORY THRUSTS
(Genital Connection)
OPPOSED POSITION
Figure 1. A summary of the mating behavior of Gromfhadorhma $or- tcntosa, indicating the possible releasers for each step in the sequence. The possible role of chemical stimuli in the release of subsequent steps in the courtship sequence is also uncertain. As noted above, there is a characteristic odor associated with the hissing male. This odor is apparent in all situations in which hissing occurs but this fact does not necessarily preclude the possibility that the odor has some effect on the level of sexual receptivity and subsequent behavior of the female, thus qualifying as a true male sex pheromone. Du- mortier (1965) reports that no odor perceptable to human olfactory receptors was associated with hissing in the closely related species, G. brunneri. A schematic representation of the mating behavior of G. portentosa indicating the possible releasers of the various stages in the sequence is shown in Figure I.




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19681 Barth -Behavior of Gromphadorhina 129 Dumortier (1965) reports briefly on the mating behavior of G. brunneri which from his description appears to be very similar to that of G. portentosa, and includes in sequence such elements as antennal contact, antennal fencing and mutual body stroking with the antennae, male circling the female with hissing (the individual hisses being both softer and shorter than those associated with ter- ritorial defense), and finally assumption of the opposed position by the male backing directly into the female. The mating behavior of species of this genus represents a marked departure from the typical pattern of cockroach mating behavior as described by Barth ( I 961, I 964, I 96th). Variation in the male's preliminary courtship activities and in the form of the male's wing raising display (present in all winged species studied with the ex- ception of Pycnoscelus indicus and two species of the genus Panchlora - Barth, 1968a) is frequently encountered in various phylogenetic lines of the Blattaria, but the absence of the female's mounting and feeding response is quite unusual and has been observed to occur in only three subfamilies of the Panchloroid complex of the family Blaberidae (classification according to McKittrick, 1964), the Pycnoscelinae (Pycnoscelus) , the Panchlorinae (Panchlora) , and the Oxyhaloinae (Gr07?zphadorlzina). In Pycnoscelus indicus, there is a reversal of the typical pattern in that the male mounts the female with very little preliminary courship (Roth and Barth, 1967). Panchlora nivea and P. irrorata resemble Gromphadorhina in that the male after a much reduced preliminary courtship achieves genital connection merely by backing into the abdominal tip of the female (Roth and Willis, 1958 ; Willis, I 966). Within the Oxyhaloinae, mating behavior follows the typical cockroach pattern quite closely in Leucophaea maderae and Nauphoeta ckerea, the only other spe- cies which have been studied (Roth and Barth, 1967). Similarly within the Panchlorinae, the only other species for which informa- tion is available, Capucina patula, shows a fairly typical mating be- havior pattern with a male wing-raising display, a female mounting and feeding response, etc. (Barth, unpublished observations). On the basis of present evidence these two subfamilies, the Oxyhaloinae and the Panchlorinae, appear to be the most interesting for study of the evolution of aberrant mating behavior patterns in the Blat- tasia. Investigation of additional species might reveal intermediate stages and perhaps permit some conclusion as to whether we are really dealing with the parallel evolutionary development of similar aberrant mating behavior patterns in related phylogenetic lines as currently seems to be the case.




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Psyche
SUMMARY
The mating behavior of Gromphadorhina portenosa represents a marked departure from the typical pattern for cockroaches. The male recognizes the female upon contact and if the female is re- ceptive engages her in antenna1 fencing and mutual stroking of the body with the antennae. Then the male proceeds to walk around the female several times hissing softly in short bursts. Finally the male orients so that his abdominal tip is opposed to that of the female and achieves genital connection merely by backing into the female. Thus the opposed copulatory position is assumed directly without the preliminary maneuvering observed in most species of cockroaches. Also included in this communication are some observations on alarm behavior and aggressive behavior both of which also employ the curious hissing (produced by the expulsion of air through the second abdominal spiracle) noted in connection with courtship be- havior.
EARTH, ROBERT H., JR.
1961. Comparative and Experimental Studies on Mating Behavior in Cockroaches, Ph.D. Thesis, Harvard University, Cambridge, Massachusetts. 274 pages.
1964. The Mating Behavior of Byrsofria fumiga.ta (Gutrin) (Blat- tidae, Blaberinae). Behaviour 23: 1-30.
1968a The comparative
physiology of reproductive processes in cock- roaches. Part I. Mating behavior and its endocrine control. Advances in Reproductive Physiology 3: in press. 1968b. The mating behavior of Periplaneta amerlcana (Linnaeus) and Blatta orientalis Linnaeus (Blattaria, Blattoidea, Blattidae, Blat- tinae) with notes on the mating behavior of three additional species of Periplaneta. Manuscript in preparation. DUMORTIER, BERNARD
1965. L'Emission sonore dans Ie genre Gromphadorhina Brunner (Blattodea, Perisphaeriidae) Etude morphologique et biologique. Bull. Soc. 2001. France 90: 89-101.
MCKITTRICK, I?. A.
1964. Evolutionary Studies of Cockroaches. Cornell Univ. Agric. Exp. Station Memoire 389, 197 pp.
ROTH, Louis M. AND R, H. BARTH, JR.
1967. The sense organs employed by cockroaches in mating behavior. Behaviour 28 : 58-94.




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19681 Barth - Behavior of Gro7nphadorhina 131 ROTH, LOUIS M. AND E. R. WILLIS
1958. The biology of Panchlora nivea with observations on the eggs of other Blattaria. Trans. Am. Ent Soc. 83: 195-207. 1960. The Biotic Associations of Cockroaches. Smithson. Misc. Coll. 141, 470 pp.
WILLIS, E. R.
1966. Biology and behavior of Panchlora irrorata, a cockroach ad- ventive on bananas (Blattaria; Blaberidae). Ann. Ent. SOC. Amer. 59: 514-516.




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THE LARVA OF MICROSTIGMUS COMES,
WITH COMMENTS ON ITS RELATIONSHIP TO
OTHER PEMPHREDONINE GENERA
(HYMENOPTERA, SPHECIDAE)*
BY HOWARD E. EVANS AND ROBERT W. MATTHEWS Museum of Comparative Zoology
Compared to other Sphecidae, members of the Pemphredoninae exhibit an unusually wide spectrum of morphological features. Lar- val characters have suggested that simple division of the subfamily into the two tribes Psenini and Pemphredonini may not be ideal. (For discussion and for previous descriptions of the larvae of Pemphredoninae or references thereto, see Evans, 1958, pp. 128- 136; Evans, 1959, pp. 139-145, 167-168; and Evans, 1964, pp. 245- 253.) Recently, larvae of another member of this group were obtained in the course of studies on the nesting biology of Microstigmus comes Krombein (see Alatthews, 1967). Because, in addition, the genus exhibits many unique behavioral features, consideration of the larva and possible relationships of Microstigmus seem particu- larly appropriate. No previous larval descriptions exist for the genus; Myers'
(1934) drawings of the body and mandible of the larva of M. theridii fail to illustrate critical characten and hence are of little use to this discussion, although the features shown essentially agree with those of Ad. comes larvae. Description of mature larva of Alicrostig7nus comes Krombein (Figures I to 3)
Length 4 mm; narrowly fusiform, gradually tapered ~osteriorly, apical segment conically produced above the anus. Body slightly constricted at the intersegmental lines, more noticeably dorsally. Integument smooth, visible spines or setae absent even under high magnification. Spiracles minute, atrial walls completely smooth, entrance into the very slender subatrium unarmed. Head 47 mm wide, 44 mm long measured to apical margin of clypeus (about as long as wide when measured to apical margin of labsum) ; head devoid of setae and completely unpigmented except for the three brown apical mandibular teeth; parietal bands absent. Antenna1 orbits large, subcircular, about 55p in diameter ; antenna1 papillae well developed, about 25p long. Labrum short, its apical margin *Manuscript received by the editor April 6, 1968 132




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THE MATING BEHAVIOR OF
GROAfFHADORHINA PORTENTOSA (SCHAUM)
(BLATTARIA, BLABEROIDEA, BLABERIDAE,
OXYHALOINAE) : AN ANOMALOUS PATTERN
FOR A COCKROACH1
Department of Zoology
The University of Texas at Austin
This communication is the fourth in a series of largely descriptive papers dealing with the mating behavior of cockroaches. (see Barth, 1961 ; Barth, 1964 ; Roth and Barth, 1967 ; and Barth, ms in prep- aration). The aim of this series is twofold: first to provide back- ground information for experimental studies, and second to provide the detailed comparative information necessary for a study of the evolution of mating behavior within the Blattaria. A more general introduction to the series may be found in Earth (1964). The anomalous mating behavior of the Madagascar cockroach, Grompha- dorhina portentosa, (Schaum) forms the subject of this communi- cation.
MATERIALS AND METHODS
Stock cultures of G. portentosa were maintained as described by Bart11 (1964) for firsotria fumigata. The observations on mating behavior were made in the evening (the normal active period for these animals) under red illumination is specially designed obser- vation chambers (for details, see Barth, 1964). The ethological terms employed in the description of the behavior patterns have been previously defined by Barth ( I 964).
RESULTS AND CONCLUSIONS
Gromphadorhma portentosa is a large, heavy-bodied wingless spe- cies found under debris on forest floors in Madagascar (see Plate 6 Wo. 4 in a series of papers entitled "The Mating Behavior of Cock- roaches".
'Much of this work was carried out at the Biological Laboratories Har- vard University. Financial support from National Science Foundation Predoctoral Fellowships and N.S.F. Grant G 19962 is gratefully acknowl- edged.
Manuscript received by the editor 5 April, 1968



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19681 Barth - Behavior of Gromphadorhina 125 in Roth and Willis, 1960). There is considerable size variation in the adults of this species, large males sometimes reaching a length of 8 to 10 cm. Males tend to be larger than females and differ from females in having antennae with many long laterally projecting sensory hairs, while the latter possess the simple filiform type of antennae characteristic of cockroaches. Males also differ from fe- males in that their pronotal shields are greatly thickened and pro- vided with a pair of large heavy knobs.
Alarm Behavior and Aggressive Behavior of Males. The most notable features of the general behavior of this species are the aggressive behavior of the males and the production, by expulsion of air through the second pair of abdominal spiracles, of a loud hissing noise when alarmed. The former is briefly described below. Two males approaching each other while foraging, raise their bodies considerably off the substratum, curving their abdomens up- wards at the tip. The rate of antenna1 waving or twitching increases markedly. When the antennae of the two come into contact they are twitched back and forth very rapidly and vigorously. Both animals then lower their foreparts so that their pronotal shields are directed towards the opponent. Rhythmic hissing may occur at this st?ge. Then they charge, their kno<bbed pronotal shields coming together frequently with an audible sound. They vigorously push each other back and forth, the winner being the one which can push his opponent backwards until it takes flight. Sometimes the victor slaps his abdomen vigorously against the substratum, and may turn and slap his abdomen against the opponent. A retreating animal may be vigorously chased for some distance by the victorous animal. The largest male of a group of males is the most frequent victor in these disputes and there is some evidence (Engelmann, pers. con^.,) that dominance hierarchies among caged animals are set up in this way.
Dumortier ( 1965) discusses the hissing behavior of a closely re- lated species, G. brunneri, and describes in some detail the mech- anism of sound production. He reports that in addition to its role in alarm behavior, males of G. brunneri also employ hissing in ag- gressive behavior. According to Dumortier the aggressive behavior of G. brunneri males (which seems to be very similar to that of G. portentosa males), appears in connection with territorial defense. Mating Behavior.
There appear to be no previous accounts of the mating behavior of G. portentosa in the literature. The present description is based



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