Sound production and reception by stored products insect pests—A review of present knowledge

J. stored Prod. Res., 1971, Vol. 7,

Sound Stored

pp. 153-162. Pergamon Press. Printed in Great Britain.

Production Products

and Reception

Insect

of Present HUBERT

Pests-A

by

Review

Knowledge

FRINGS

and MABLE

FRINGS

Department of Zoology, University of Oklahoma, Norman, Oklahoma, U.S.A. (First received 1 December, 1970, and injnalform

20 June, 1971)

Abstract-The published literature on production and reception of sounds and acoustical behavior of beetles of eleven families and moths of two families that include important stored products pests is reviewed. Little is known directly on the pest species, but studies on related species suggest that the pests may have acoustical reactions that could form the basis for control. Except for one promising study on Plodia interpunctella, no critical attempts have been made to control stored products pests with sound. INTRODUCTION THE

of substituting physical for chemica1 methods in control of stored insect pests is quite attractive. So far, however, there has been no routine

POSSIBILITY

products

use of such easily monitored of sounds cidental

for practical discovery-can

of the pest species. pertinent literature search this

come

as electricity,

products

through

and reception

knowledge,

acoustical controls. For this review,

only

of energy

of stored

light

or sound.

pests-barring

knowledge

some

of the

The

use

totally

ac-

acoustical

behavior

With this in mind, the present review and bibliography is presented in the hope that it will suggest and facilitate

on production

new

forms

control

we may we shall

of sounds

be able

consider

by stored

to predict

that

and

the major

products

insect pests. With

critically

insect

pests

evaluate

possible

of stored

products

belong to the Coleoptera (Families: Anobiidae, Bostrichidae, Bruchidae, tidae, Cucujidae, Curculionidae, Dermestidae, Nitidulidae, Ostomatidae, Silvanidae, Pyralididae). pests might papers

and

Tenebrionidae),

Because

be suggestive,

on death-watch

references

without

and

work on species

the Lepidoptera in these families

this is also briefly

beetles

particular

(Anobiidae)

reviewed.

as irrelevant;

(Families that

CorynePtinidae,

: Gelechiidae

are not stored

We have

of re-

omitted

and

products the many

they would add many more

utility. COLEOPTERA

Sound production In our bibliography sound-producing species

about 50 families (1960), are listed. Of these, about 153

of beetles

that

include

25 are well known

known

to entomolo-

151

HUBERTFRINGSand MABLE FRINGS

gists; a few (e.g. Cerambycidae, Elateridae) were known even to the ancients. (LANDOIS, 1874; SWINTON, 1880). As HASKELL (1961, p. 41) notes: “It may be surprising to some to discovTer the tremendous range of stridulatory mechanisms in beetles. They have been almost completely overlooked in recent research on insect acoustics, and yet they were paradoxically one of the first groups in which much concentrated work was carried out on the stridulatory mechanisms. .. Darwin, in the Descent of Man, dilated at some length on the meaning and significance of sounds produced by beetles.” Of the eight major methods of sound production by insects (FKINGS and FRINGS, 1958) only two (use of tymbals and expulsion of air over a vibrating membrane) are not found in some beetles (ARROW, 1942). Among the stored products pests, only Dermestes Iardarius L. (BAILEY and LEMON, 1968) has been found to make sounds, in spite of the extended efforts reported by WOJCIK (1968, 1969a, b). Two families that include important stored products pests-Curculionidae and Tenebrionidae-have many species that are well known as sound producers. For weevils, a number of authors have described presumed stridulatory structures on the elytra and abdomen of many genera: PATTON (1884), GAHAN (1900), SIMM (1911), KLEINE (1913-31), DUDICH (1920, 1923), and MARCU (1930-1939). Other authors have reported hearing weevils stridulate and have usually also indicated the mechanism of sound production : WESTRING ( 1847)) WOLLASTON ( 1860)) RILEY (1871), MORLEY (1902), TEMPERE (1937), KORSAKOFF (1942), GIBSON (1967) and HARMAN and KRANZLER (1969). TEMPERE ( 1937) states categorically that he has found no weevil that does not stridulate, and that one needs just put himself to listening to discover the sounds. However, WOJCIK found (1968, 1969a, b) that a stored products infesting weevil, Sitophilus granarius (L.), is not a sound producer. A number of tenebrionids likewise produce sounds, as described by GOUREAU (1837, 1838), SOLIER (1837), DUDICH (1920), REMY (1935), PRIESNER (1949), and LILES (1956). For Ptinidae, HOUGHTON ( 1906) reports stridulation in only one species. The presence of stridulatory organs suggests that the sounds produced are significant in the lives of beetles, but few studies have been made. RILEY (187 1) suggested that sounds made by the plum curculio, Conotrachelus nenuphar (Herbst), might be important in communication. Conversely, KLEINE (1919a) working on curcuconcluded that stridulatory sounds in lionids, and REMY (1935) on tenebrionids, beetles have no significance. However, the studies of GRAY (1946) on acoustical signalling in passalids, of ALEXANDER (1957) and ALEXANDER et al. (1963) on trogids, passalids and cerambycids, of VAN TASSELL (1965) on a hydrophilid, and of MICHELSEN (1963, 1966a, 6) on cerambycids demonstrate that-for these beetles at least-stridulatory sounds have biological functions. MAMPE and NEUNZIG (1966) and GIBSON (1967), furthermore, have experimentally confirmed Riley’s hypothesis for C. nenuphar by demonstrating acoustic signalling. PRIESNER (1949) noted that a tenebrionid he found stridulating answered imitations of its sounds. And the socalled death-watch beetles (Anobiidae) have been known for a long time to answer tapping that is similar to their sounds. For the beech weevil, Rhynchaenus.fagi (L.), CLARIDGE (1968) has shown that four identifiable sounds with different uses by the insect are produced. Similar, but often distinguishable, signals are produced by three other species of the same genus, suggesting that the sounds are important in behavioral isolation.

Sound

Production

and Reception

by Stored

Products

Insect

Pests

155

Summarizing, then, sound production by a wide variety of methods is widespread in beetles. Except for Dermestes lardarius, however, specific sounds produced by pests of stored products have not been observed. The importance of sounds in behavior of the few weevils that have been studied suggests that these beetles, at least, should be more closely examined in this regard. Sound reception Some sensory physiologists have argued fruitlessly that true hearing does not occur in animals other than vertebrates. The argument is entirely semantic (cf. FRINGS, 1964, for discussion). If sound is defined as physicists define it: periodic compressional waves in an elastic medium, then so-called vibrations are merely solid-borne sounds. Reception of vibrations, therefore, is just as truly phonoreception as is reception of air-borne or liquid-borne sounds. Because sound is a mechanical agitation of a medium, any mechanoreceptor capable of responding to alternating waves of pressure or particle movement is a potential phonoreceptor. Even man receives air-borne sounds not only through the tympanum of the ear, but also through bone conduction and particularly at low frequencies through cutaneous receptors. For insects, therefore, we must consider any mechanoreceptor as a potential receptor for sounds. The following mechanoreceptors, at least, are probably involved in phonoreception in some insects: tactile hairs, spines, or hair plates; campaniform sensilla; muscle receptor organs; and chordotonal organs, either in simple forms in appendages, or as parts of complex sensilla, such as tympanal organs and Johnston’s organs. Studies upon the general morphology and locations on the body of these are numerous enough (cf. FRINCS and FRINGS, 1960 for references), but critical experiments are very few and have involved only a few species from scattered orders. For Coleoptera, even counting strictly morphological studies with no data on function, as in taxonomic papers, representatives of only about 25 families have been surveyed. For no species has there been any study in depth. We shall not attempt a general review of mechanoreception in insects, for BUSNEL ( 1963)) HASKELL ( 196 1)) DETHIER (1963) and BELTON (1962) give good surveys. For pests of stored products, morphological studies include CROMBIE’S (1944) brief note on various sensilla, mostly chemoreceptive, on legs and antennae of Rhyzopertha dominica (F.) , and SUSTER’S papers (1930, 1933, 1936) on Johnston’s organs of Tenebrio molitor L. and Dermestes lardarius. HAYES et al. (1967) and SOKOLOF et al. (1967) have recently reported that Tribolium confusum Duv. can live for some time in the vacuum needed for scanning electron micrography. This allowed them to get highly magnified pictures of exoskeletal structures in the living animals. Reactions of stored products insects to environmental tactile cues or to agitation of the medium in which they live have been studied by BORELL DU VERNAY (1942) in Tenebrio molitor, BROWNING (1946) in Sitophilus granarius (L.), ADAMS et al. (1953, 1954) in Sitoflhilus oryzae (L.), HOLLIS, (1963) in T. molitor, GRAHAM and WATERHOUSE ( 1964) in Tribolium spp., AMOS (1965) in Carpophilus spp., MALLMANN(1965) in T. molitor, LEVINSON and BAR ILAN (1970) in Trogoderma granarium Everts, and ARBOGAST and CARTHON ( 1970) in Oryzaephilus surinamensis (L.). In none of these cases were the actual receptors discovered or their physiology studied. For other Coleoptera, studies on potential sound receptors, not to say sound

156

HUBERTFRINGSand MABLEFRINGS

reception-even mechanoreception in general-are amazingly scarce. Beetles lack any obviously specialized tympanal organs, although Johnston’s organs are almost universal in the group. However, CROWSON (1963) has reported what he interpretswithout tests-as a tympanal organ in a staphylinid, Philonthus sp. Since this organ was not reported earlier, in spite of the many morphological studies on staphylinids, it suggests that other structures of possible acoustic significance might be discovered with careful search. MCINDOO (1926) and D~NGES (1954) have described Johnston’s organs in two species of weevil, and MALLMANN (1965) has studied the tactile reactions of a nitidulid and curculionid. There is also the indirect evidence from studies on communication mentioned previously (studies on death-watches, the works of Alexander, Alexander et al., Van Tassell, Michelsen, Priesner, Claridge, Mampe and Neunzig, and Gibson). Experimentally, CORBI$RE (1967, 1968) has demonstrated that hairs of the larva of a cavernicolous silphid are truly tactile, the first such demonstration for Coleoptera. AUTRUM and SCHNEIDER (1948) and SCHNEIDER (1950) studied electrophysiologically chordotonal organs of the legs of various insects, including a few species of beetles. They found the beetles variously sensitive : Melolontha melolontha L. and Geotrupes sylvaticus Panz. very sensitive to substrate vibration, and tvio species of carabids (Pterostichus vulgaris L. and Pseudophonus sp.) and a silphid (Silpha sp.) quite insensitive. Obviously knowledge of the physiology, even the morphology, of phonoreceptors in Coleoptera is in a sub-rudimentary state, This is undoubtedly due to the fact that Coleoptera have no recognized phonoreceptive organs (the one described by Crowson still needs proof of function). Johnston’s organs, apparently present in most beetles, have been studied carefully only in mosquitoes, where they are definitely involved in reception of air-borne sounds. For beetles, the only experimental studies on Johnston’s organs have been on gyrinids (EGGERS, 1926, 1927, DE WILDE, 1941). In these beetles, the organs are receptors for two dimensional surface waves on water. So studies on potential and actual sound receptors in beetles are greatly needed. First should be demonstration of the actuality of reception of sounds by the \:arious mechanoreceptors present on the bodies of beetles. Then fruitful morphological and physiological studies can be essayed. The presence of many specialized types of stridulatory structures in beetles certainly indicates that reception of sounds is important in social and sexual behavior of these insects. The presence of a thick exoskeleton, a good conductor of sound, indicates that solid-borne sound (vibrations) might be of particular importance (MARKL, 1969). Altogether beetles offer a tremendous challenge for future research in bio-acoustics. LEPIDOPTERA Sound production in Lepidoptera has been much less studied than in Coleoptera. The death’s head moth (Sphingidae) has been the subject of a sizeable literature. Otherwise only about 40 families have been reported as having sound producing species, many of these reported in one study on pupal sound production (HINTON, 1948). As far as stored products pests are concerned, WOJCIK (1968, 19693) reported no sounds other than incidental from the two pyralidids (Ephestia elutella (Htibner) and Cadra cautella (Walker)) he monitored for long periods.

Sound Production and Reception by Stored Products Insect Pests

157

In contrast with the Coleoptera, however, studies on sound reception, at least by phalaenid moths, are relatively numerous. Tympanal organs are present on the metathorax in 13 families of moths, including Phalaenidae, and on the abdomen in 7 families, including Pyralididae (EGGERS, 1928). The stored products insects in this last family have well developed tympana, but morphological studies have been published only on Gall&z mellonella (L.) (MULLEN and TSAE, 1971). The tympanal organs of pyralidids have been described grossly for a few species, none stored products pests, by KENNEL (1912), KENNEL and EGGERS (1933), and EGGERS ( 1928, 1937), but without experiments. KIRKPATRICK and HAREIN (1965) showed that adults of Plodia interpunctella (Hiibner) apparently respond to air-borne sound by reduced oviposition. TREAT (1955) tested reactions to sounds in a variety of moths, including two species of day flying pyralidids. BELTON (1962a) electrophysiologically demonstrated reception of sounds by the tympanum of Ephestia kuehniella (Zeller) and Galleria mellonella (L.). These moths are sensitive to ultrasonic sounds, as are other tympanate species, but the mechanism of action of their tympanal organs is essentially unknown. Other mechanoreceptors, as in beetles, are potentially involved in reception of sounds, as was shown for Ctenucha virginica (Charpentier) (FRINGS and FRINGS, 195 7)) and for larvae of caterpillars by MINNICH (1936). Chordotonal organs, other than those associated with the tympana, have been described for larvae of pyralidids (HENIG, 1930) and in the wings of certain gelechiids (VOGEL, 1910). Johnston’s organs are present, but their importance is uninvestigated. Summarizing, the situation with respect to Lepidoptera is only better than that for Coleoptera insofar as studies on the specialized tympana of phalaenids are concerned. Otherwise the location, morphology, and function of phonoreceptors remain mostly to be discovered. POSSIBLE

USE OF SOUND FOR CONTROL

OF STORED PRODUCTS

PESTS

Some efforts have been made to use sound or other mechanical agitation to control pests in stored products. Among the earliest, were studies of RoBERTsoN ( 1944), who proposed a device for removing pests from stored products by combining mechanical agitation with light and heat. BROWNING (1946) studied pullulation of Sitophilus granarius with a similar idea in mind, but did not reach any practical conclusion. Agitation and stirring seriously reduce oviposition or development of insects in grains, as reported by COOMBS (1956), LOSCHIAVO ( 1960), JOFFE (1963) and JOFFE and CLARKE (1963). BAILEY (1962) projected grains against a hard surface killing larvae inside, but this also had deleterious effects on the grains. Later BAILEY (1969) used methods like those of Joffe and Clarke and found that even small disturbances prevented development of the pests. None of these workers used scund, as such, but KIRKPATRICK and HAREIN (1965) did, finding that submission of ovipositing Plodia interpunctella populations to sound fields reduced emergence in the next generation. The sounds (120-2000 Hz) were not ultrasonic, although these moths are most sensitive in the ultrasonic range (BELTON, 1962a). Possibly extension of tests into the ultrasonic would improve control. Larvae feeding and adults moving in stored products make incidental sounds. Consequently detection of infestation should be possible by acoustical monitoring.

HUBERT FRINCS and MABLE FRINCS

158

been reported by ADAMS et al. ( 1953, 1954) and BAILEY and MCCABE Unfortunately, such high amplifications are needed to detect these sounds that practical applications are not now feasible. So far, attempts to use sound in insect control have been few, and usually disappointing (FRINGS and FRINGS, 1962a, b, 1965, 1968; FRINGS, 1969; ALEXANDER, 1967; WATTERS, 1965). The studies of BELToN ( 19626) and BELTON and KEMPSTER (1962) on protection of corn from corn borer attack, using ultrasonic frequencies of sound, and those of PAYNE and SHOREY (1968) on cabbage looper moths are promising and lead one to believe that sounds may ultimately have some value in insect control. However, barring some lucky chance, it certainly seems that we must get much more fundamental information about phonoreception and acoustical behavior of insects than we now have if we are to project possibly fruitful techniques (BELTON, 1962a). At least as far as pests of stored products are concerned, our ignorance of acoustical physiology is almost complete. Possibly if we had the necessary knowledge, we could plan equipment and techniques for use of sound in pest control that might have a reasonable chance of success. Without this knowledge, the attempts are mere shots in the dark. This

has

(1965).

Acknowledgements-This work was supported by Agricultural Research Service, U.S. Department of Agriculture, Grant No. 12-14-100-9196 (51), Stored-Product Insects Research Branch, Market Quality Research Division. REFERENCES ADAMS, R. E., WOLFE, J. E., MILNER, M. and SHELLENBERGER,J. A. (1953) Aural detection of grain infested internally with insects. Science, N.Y. 118, 163-164. ADAMS, R. E., WOLFE, J. E., MILNER, M. and SHELLENBERGER,J. A. (1954) Detection of internal insect infestation in grain by sound amplification. Cereal Cizem. 31, 271-276. ALEXANDER, R. D. (1957) Sound production and associated behavior in insects. Ohio J. Sci. 57, lOl113. ALEXANDER, R. D. (1967) Acoustical communication in arthropods. A. Rev. Ent. 12, 495-526. ALEXANDER, R. D., MOORE, T. E. and WOODRUFF, R. E. (1963) The evolutionary differentiation of stridulatory signals in beetles. Anim. Behav. 11, 11 l-l 15. AMOS, T. G. (1965) Kinesis. Nature, Lond. 208, 908-909. ARBOGAST, R. J. and CARTHON, M. (1970) Light, tactile and humidity responses of larvae of OryCucujidae). Entomologia exp. a&l. 13, 395-402. zaephilus surinamensis (Coleoptera: ARROW, G. J. (1942) The origin of stridulation in beetles. PZOC.R. ent. Sot. Land. A17, 83-86. AUTRUM, H. und SCHNEIDER,W. (1948) Vergleichende Untersuchungen uber den Erschtitterungssinn der Insekten. 2. vergL. Physiot. 31, 77-88. BAILEY, S. W. (1962) The effects of percussion on insect pests of grain. J. econ. Ent. 55, 301-304. BAILEY, S. W. (1969) The effects of physical stress in the grain weevil Sitophilus granarius. J. stored Prod. Re.T. 5, 31 l-324. BAILEY, S. W. and LEMON, R. W. (1968) Sound production by the larder beetle, Dermestes lardarius Linnaeus. J. stored Prod. Res. 4, 271-273. BAILEY, S. W. and MCCABE, J. B. (1965) The detection of immature stages of insects within grains of wheat. J. stored Prod. Res. 1, 201-202. BELTON, P. (1962) The physiology of sound reception in insects. PYOC.ent. Sot. Ontario 92, 20-26. BELTOW, P. (1962a) Responses to sound in Pyralid moths. Nature, Lund. 196, 1188-l 189. BELTON, P. (19626) Effects of sound on insect behaviour. PYOG. ent. Sot. Manitoba 18, l-9. BELTON, P. and KEMPSTER, R. H. (1962) A field test on the use of sound to repel the European corn borer. Entomologia exp. appl. 5, 281-288. BORELL DU VERNAY, W. v. (1942) Assoziationsbildung und Sensibilisierung bei Tenebrio molitor L. 2. z:ergl. Phpiol. 30, 84-116.

Sound

Production

and Reception

by Stored

Products

Insect

Pests

159

BROWNING, H. C. (1946) Mechanical disturbance and light as factors influencing pullulation of Calandra granaria Linn. Proc. zool. Sot. Lond. 116, 675-691. BUSNEL, R. G. (1963) Acoustic Behaviour of Animals. Elsevier, Amsterdam. CLARIDGE, L. C. (1968) Sound production in species of Rhynchaenus (=Orchestes). Trans. R. ent. Sot. Lond. 120,287-296. COOMBS, C. W. (1956) Stability of grain as a factor influencing the oviposition rate of the grain weevil, Calandra granaria. Bull. ent. Res. 47, 737-740. CORBI~RE, G. (1967) Mise en evidence tlectrophysiologique de la function tactile des sensilles tricho’ides sit&es sur l’antenne de la larve de S’eophyes lucidulus Delarouzee. C.r. hebd. Seam. Acad. Sci. Paris (D) 265, 1327-1329. CORBI~RE, G. (1968) Determination tlectrophysiologique de la function tactile des grandes sensilles trichoides des segments abdominaux chez la larve du Speophyes lucidulus (Delar.). C.r. hebd. Seam. Aced. Sk., Paris (D) 266, 18751878. CROMBIE, A. C. (1944) Sensillae of the adults of larvae of the beetle Rhizojertha dominica Fab. Proc. R. ent. Sot. Lond. A19, 131-132. CROWSON, R. A. (1963) On a supposed tympanal organ in the staphylinid genus Philonthus. Entomologist’s mon. Msg. 98, 197. DETHIER, V. G. (1963) The Physiology of Insect Senses. Wiley, New York. DBNC:ES, J. (1954) Der Kopf von Cionus scrophulariae L. Zool. Jb., (Anat.) 74, l-76. DUDICH, E. (1920) Uber den Stridulationsapparat einiger KIfer. Ent. Bl. Biol. Syst. Kiifer. 16, 146-161. DUDICH, E. (1923) Uber die Stridulation von Lepyrus capucinus &hall. Ent. Bl. Biol. Syst. Ka$r 19,

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Sci. 66, 131-135. HOUGHTON, C. 0. (1906) Notes on Caenocara oculata Say. Ent. News 17, 114-l 15. JOFFE, A. (1963) The effect of physical disturbance or ‘turning’ of stored maize on the development of insect infestations-I. Grain elevator studies. S. Afr. J. agric. Sci. 6, 55-64. JOFFE, A. and CLARKE, B. (1963) The effect of physical disturbance or ‘turning’ of stored maize on the development of insect infestations-II. Laboratory studies with Sitophilus oryzae (L.). S. Afr. J. agric.Sci. 6, 65-84. KENNEL. .J. v. (1912) Uber Tympanalorgane im Abdomen der Spanner und Zunsler. Zool. Ant. 39, 163-170. KENNEL, J.

v. and EGGERS, F.

(1933)

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moths,

Plodia interjnmctella, by exposure

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Zeit. 6, 357-359. KLEINE,

R.

(1917)

Der

Stridulationsapparat

der Gattung

Li’arus

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54-68. KLEINE, R. (1918)

Der Stridulationsapparat

der Gattung

Lepyrus Germar.

Ent. BI. Biol. Syst. Kc@.

14,257-274. KLEINE, R.

(1919a)

Hat der Stridulationsapparat

der Rhynchophoren

einen

Zweck?

Ent. Bl. Biol.

Syst. Kiifer 15, 193-202. KLEINE, R. (1919b)

Der Stridulationsapparat

der Gattung

Anisorrhynchus Schiinh.

Ent. Bl. Biol. Syst.

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