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THE EVOLUTIONARY DIFFERENTIATION of STRIDULATORY SIGNALS in BEETLES (Insecta : Coleoptera)

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THE EVOLUTIONARY DIFFERENTIATION of STRIDULATORY SIGNALS in BEETLES (Insecta : Coleoptera) THE EVOLUTIONARY DIFFERENTIATION OF STRIDULATORY SIGNALS IN BEETLES (Insecta : Coleoptera) BY RICHARD D . ALEXANDER AND THOMAS E . MOORE Museum of Zoology and Department of Zoology, The University of Michigan, Ann Arbor AND ROBERT E. WOODRUFF Florida Department of Agriculture, Division of Plant Industry, Gainesville Introduction "Spontaneous" Stridulation in Trogidae Stridulatory noises have been discussed for While collecting at night with headlamps on many different families of beetles, and presumed 16th May, 1959, in Ripley County, Missouri, stridulatory organs have been described for we heard chirping coming from the ground hundreds of species that have not actually been around a dead cat lying in a sparsely vegetated heard stridulating. (cf. Arrow, 1904, and biblio- area along the roadside. Using a portable tape graphy by Frings & Frings 1960). So far, recorder, earphones, and a microphone to neither auditory organs nor acoustical functions amplify and localize the sounds, we found that have been experimentally demonstrated in any chirping came from several burrows, each about beetle. There are a few records of beetles stridu- 8 mm. diameter, which were scattered within a lating upon contact with other individuals of few feet of the carcass . The burrows were about their own species (Liles, 1956 ; Alexander, 1957), 10 centimetres deep and nearly vertical ; each but otherwise, only gross disturbances, e .g. of several excavated contained one adult female handling or restraint, have been observed to of Trox suberosus Fabricius at its lower end . elicit stridulation. One burrow contained a male and a female ; As a more or less incidental adjunct to in- it is not known whether sound was coming from vestigations on the acoustical behaviour of this particular burrow. Orthoptera, Homoptera, and Hemiptera by The sound was a continual irregular, repeating Alexander & Moore, and as a result of the of the phrases shown in Fig. 1 . Each phrase was particular efforts and interest of Woodruff, produced by one stroke of a small ridge (pleg- we have accumulated tape-recordings of about matium) on the dorsal surface of the abdomen 40 beetle species in eight families ; Passalidae, across two tooth-like ridges (files) on the under- Trogidae, Scarabaeidae, Chrysomelidae, Tene- side of the elytra. The file ,(Figs. 18 and 19) brionidae, Carabidae, Curculionidae, and Cer- covers approximately the posterior one-third of ambycidae . In this paper we compare disturb- the length of the elytra and is composed of over ance stridulation among congeneric, sympatric 150 teeth in a very narrow strip running parallel species in two subsocial families (Trogidae and to the suture on a raised carina . The teeth in the Scarabaeidae) and one non-social family (Cer- middle of the ridge are slightly more widely ambycidae), and discuss two new kinds of beetle separated than those on either end but they are stridulations : a "spontaneous" stridulation in all minute and difficult to detect except under isolated, undisturbed Trogidae and a distinctive high magnification (60X) . Sharp (1897) described aggressive stridulation in the social Passalidae . similar stridulatory apparatus in several Trox All of these sounds taken together appear to species. During stridulation the abdomen is illustrate three stages in the evolutionary elabor- moved up and down and slightly forward and ation of communicative signals : (1) a single backward to engage the plegmatium (Fig. 16) structural kind of signal utilized in a single with the file. This movement is not extensive but functional context (disturbance-actual function is easily observed. Apparently only a portion of not proven) ; (2) a single structural kind of signal the narrow file is struck on each movement, since utilized in two different functional contexts a maximum of twenty tooth strikes is eindicated (disturbance and calling, disturbance and con- on the audiospectrograph. gregation, and possibly disturbance and some The beetles (both sexes) produced essentially sort of "recognition") ; and (3) two structurally the same sound when captured (Fig. 2), different signals utilized in two different func- and when isolated overnight in vials and left tional contexts (disturbance and aggression) . undisturbed . Some individuals produced this 111 112 ANIMAL BEHAVIOUR, XI, 1 sound continually for very long periods of time proprioceptive feedback, or through rhythms while lying otherwise motionless on their backs . inherent in the make-up, of the central nervous Using the amplifier of a tape recorder as a system . Trox suberosus may represent an inter- monitor, we found that some beetles were still mediate stage in the divorcement of a sound stridulating hours after being isolated in padded from its old stimulus situation (probably dis- chambers completely separated from any reason- turbance) as a result of its having acquired a able possibility of external disturbances.* new function . The situation here suggests that To our knowledge there is no previously re- the continual chirping may attract males into corded instance of a beetle producing a stridulat- the female's burrow, or deter other females from ory sound in the absence of disturbance by entering occupied burrows, or do both. Each other animals . In other disturbed beetles that burrow was undoubtedly destined to become we have recorded, the stridulation always the home of one pair of these sub-social beetles stopped a few seconds after the disturbance was and their off spring. discontinued. In the particular case of Trox suberosus, even if stridulation in the burrows Aggressive and Other Stridulations in Passalidae was initiated by our footsteps-unlikely because There is little doubt that sound has acquired we were several yards away and motionless a congregating function in the social Passalidae, when the chirps were first noticed-it continued where larvae have evolved the ability to produce during several long periods when we were pro- a sound similar to that of the adults by means of viding no disturbance at all, and it was delivered a strictly juvenile apparatus on the larval legs at thd' same tempo regardless of our activities . (Figs. 11-13). Passalids live in dense colonies, The "spontaneous" stridulation of Trox and stridulate almost continually . The adults suberosus can be analogized with the calling and guard, feed, and otherwise care for their off- congregating noises of male Orthoptera and spring from egg to adult (cf. Miller, 1931 ; Homoptera, often termed spontaneous, not so Ohaus, 1900). Colonies develop from one or much because of the kind of initiating stimulus, two adults isolated with 5-20 juveniles that are but because once started they continue for long kept by the adult in a compact group . Passalid periods unless disturbances cause their cessation . sounds have evolved somewhat farther than Although the thresholds for the production of other beetle noises known, as is revealed by the such sounds can be adjusted by different events distinct aggressive sound of Popilius disjunctus -for example, changes in light intensity (Alex- (Illiger) (Fig. 12), which Mr. William Hebb and ander, 1960) and the presence or absence of a Dr. Thomas J. Walker at the University of spermatophore in the spermatophore pouch in Florida (personal communication) first heard Orthoptera (Huber, 1960)-it is not known and tape-recorded . Subsequently Alexander re- whether any of them are ever initiated in a norm- corded it from Florida and North Carolina al animal without external stimulation ; in fact, beetles just brought in from the field . Hebb without external acoustical stimulation of one and Walker heard it produced by an adult sort or another. We have not yet satisfactorily brought into the laboratory during spring (1960) answered the ancient question : "What makes when individual broods of young larvae were the cricket chirp?" present in the field with one or both parents . "Spontaneous" or long-continued sound pro- We heard it in fall (1960) when only adults and duction could evolve from any kind of signal pupae were present, caged together in large requiring a specific and more or less continual numbers. Always it is produced by an aggressor stimulation through.: (1) a lowering of threshold during fighting . The stridulator advances, biting with respect to the required intensity or specific- at the other beetle, rooting him about with the ity of the external initiating stimuli and (2) a head, and sometimes lifting him into the air tendency to continue the reaction after the in- after seizing some appendage with the man- itiating stimulus has disappeared. The latter can dibles. We have seen reciprocal aggression with be effected through auditory feedback, through aggressive stridulation only once, although the fights were sometimes violent, and once a non- *All sounds were recorded with a Magnemite Portable reciprocating beetle lost an appendage . One Tape Recorder, Model 610-E, or a Magnecorder, of the aggressive beetles in our cages returned Model PT63AH, and a D-33 American Dynamic Microphone, at a tape speed of 15 inches per second . three times to the same crevice and took up Specimens and tape recordings are located in the the same stance after attacking beetles passing University of Michigan Museum of Zoology . near it, suggesting some form of territoriality . ALEXANDER et al. : EVOLUTIONARY DIFFERENTIATION OF STRIDULATORY SIGNALS IN BEETLES PLATE IX Figs. 1-5 . Audiospectrographs of sounds of Trox species (Trogidae) : Fig . 1 . T. suberosus Fabricius, "spontaneous" stridulation of an adult in a burrow, 61°F . ; Fig . 2. Tsuberosus, disturbed adult, 70°F . ; Fig. 3 . T. variolatus Melsheimer, disturbed adult, 80°-85°F . ; Fig . 4 . T. unistriatus Beauvois, disturbed adult, 80 °-85 ° F. ; Fig . 5 . T.capillaris Say, disturbed adult, 80°-85 °F . Anim. Behav., 11, 1 ANIMAL BEHAVIOUR, XI, I PLATE X Figs. 6-10. Audiospectrographs of disturbance sounds of adult Scarabaeidae (Geotrupinae) and Cerambycidae : Fig . 6, Geotrupes splendidus (Fabricius) (Geotrupinae), 80 ° -85 ° F . ; Fig. 7, G . black- burnii (Fabricius), 80°-85°F . ; Fig . 8, G . serniopacus Jekel, 80°-85°F . ; Fig . 9, Tetraopes tetraophthalmus (Forster) (Cerambycidae), 83°F .
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