Author Institution: Museum of Zoology and Department of Zoology, The University of Michigan, Ann Arbo

Moore, Thomas E.

The Ohio Journal of Science

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THE OHIO JOURNAL OF SCIENCEVol. 62 MAY, 1962 No. 3ACOUSTICAL BEHAVIOR OF THE CICADAFIDICINA PRONOE (WALKER)(HOMOPTERA: CICADIDAE)THOMAS E. MOOREMuseum of Zoology and Department of Zoology, The University of Michigan, Ann ArborOn 4 April 1959, while descending from the cloud forests on the eastern slopesof the Sierra Madre de Oaxaca into the Valle Nacional, Oaxaca, Mexico, I graduallybecame aware of a chorus of bell-like sounds above the roar of our Jeep pick-uptruck. The chorus was continuous in the tropical, broad-leaved, semi-deciduousforest bordering the Rio Valle Nacional along the road to Tuxtepec, Veracruz,from below Ciudad Valle Nacional (60 km roadpost) to the roadpost 50 km westof Ciudad Miguel Aleman, Veracruz. The cicadas were singing in trees andshrubs along the river and along the border of the very tall, heavy forests coveringthe steep slopes along the valley sides. Many of the trees on these steep hill-sides were 80 to 90 ft tall.I stopped to tape-record the song at about the 55 km roadpost at 10:00 am,and discovered that nearby individuals did sound bell-like, but instead of theapparent two-part sound as heard from a distance above the truck's roar, therepetitious, musical phrases of their calling songs resembled the ". . . —" of"V" in Morse code, the "V's" separated by short buzzes of lower intensity. Theindividuals recorded were singing in the tops of trees that reached about 20 ftabove the rank vegetation along the steep bank of the river. This was theapproximate limit of heavy forest which descends onto the coastal and riverplains. Cleared fields and orchards were scattered along the road.The cicadas sang for at least an hour, and during this time the sky was heavilyovercast—almost as dark as at dusk because of the very heavy and very lowcloud cover. Between 10:30 and 11:00 am the clouds lifted, and the males rapidlydropped out of the chorus, one by one. During chorusing, individual males sangonce or twice, then flew a short distance and repeated the performance. A fewrelatively isolated males singing in the tops of very tall trees were seen in flightbetween calling songs, but no females were observed flying toward them. Despiteour efforts, no specimens were collected here and no nymphal skins were seenamong the dense vegetation. This was the only area in which cicadas were heardsinging all the way from Cd. Valle Nacional to Cd. Aleman.About 6:30 that evening I heard the same cicada song start up suddenly atthe edge of Cd. Aleman. Two small choruses of males were singing in isolatedoak trees near a jai alai fronton (court) at the edge of a lime orchard borderingthe Visitor's Cottage of the Comision del Papaloapan. These trees were about50 ft tall and relatively small at the base. I climbed one of the trees and shot 7individuals (5 males and 2 females) that were still out of reach but silhouettedagainst the twilight sky. All individuals shot fell into the thick grass and othervegetation below. In the failing light, only 3 of the individuals, all males, couldbe retrieved.THE OHIO JOURNAL OF SCIENCE 62(3): 113, May, 1962.114 THOMAS E. MOORE Vol. 62During the last 5 min of singing activity, about 15 individuals, both males andfemales, flew into an area of approximately 24 ft3 where two males were the lastones singing. They were about 25 ft above the ground on the west or sunsetside and near the perimeter of the crown of an isolated oak tree. Some individualsflew to this tree from smaller trees and shrubs at least 100 ft away. No noisesother than calling songs were heard and no copulations were observed, althoughseveral males and females were within a few inches of each other. Here chorusinglasted from about 6:30 to 7:00 pm. beginning and ending abruptly. It seemedabout as dark as 6:30 at Cd. Aleman as it had along the Rio Valle Nacional at 10:00am. On the morning and evening of March 29, and on the morning of April 5,the weather was calm, clear, and warm, and I could hear no cicadas singing inCd. Aleman. It was evidently early in the period of adult activity, perhaps tooearly to expect copulation. The two locations at which I heard this speciesranged from approximately 100 ft in elevation at Cd. Aleman in Veracruz to about600 ft in the Valle Nacional in Oaxaca, and were both near tributaries of the RioPapaloapan.Each call has three parts: 1) a short buzz, swelling in volume, followed by2) a series of 50 to 60 repeated units described below, and 3) a more or less con-tinuous terminal whine lasting 4 to 5 sec. An entire call lasts about 50 to 60sec, and gradually increases in intensity to the terminal whine. The repeatedunits consist of three short, musical phrases (whines), followed by a longer whine,which is followed by a short buzz containing usually 9 short phrases (fig. 1). Twoor three buzzy phrases are normally produced between whines in each unit. Theseunits are repeated at a rate of 1 to 1.5 per sec. At the end of a call only twodistinct, short whines are produced, while the third is run together with the finalwhine of the preceding phrase (fig. 2). Figure 3 shows the concluding portion ofthe terminal whine of one calling song.These calling songs were recorded at 78°F, 50 or more ft from the singingindividual. No complete calling songs were tape-recorded, but one of the tworecordings was of a nearly complete song (UMMZ Tape No. 543). Figure 4shows the irregular buzzing of a disturbance squawk (UMMZ Tape No. 544),recorded at 83°F with the microphone 2 in. from a hand-held Cd. Aleman malethat apparently had only wing damage after being shot with a .22 caliber dustshot load. Both calling songs and disturbance squawks contain a relatively nar-row band of frequencies at about 4 kc/sec. All recordings and analyses weremade with the equipment and techniques described by Moore (1961), except thatan aluminum parabolic reflector 2 ft in diameter with a focal length of 6 in. (madeby C. W. Torngren Co., 236 Pearl Street, Somerville 45, Massachusetts) was usedin recording the calling songs.The audiospectrographs in figures 5 to 8 are made at different analyticalsettings in order to show more detail of structure. It is evident from these audio-spectrographs that rate and duration of timbal-muscle contraction and relaxationis the same in all types of calls. Each pulse of sound (represented by a darkvertical mark) is apparently produced by either an in- or an out-popping of thesynchronized pair of timbals; the closest pairs of pulses evident in figure 8 un-doubtedly represent reciprocal timbal pops. In this species, pulses produced inin- and out-pops are usually of similar intensity. Of particular interest is thefact that there appear to be two contraction rates. The complicated phrase-program of calling songs has been superimposed on one contraction rate. Thisrate of contraction, shown in the regular and rapid pulses of the whining partsof calling songs, indicates a timbal-muscle contraction every 0.0046 sec (220 to230 contractions per sec) to produce 440 to 460 pulses per sec (fig. 6). At thisrhythm, apparently the maximum possible at this temperature in F. pronoe, theintervals between contraction and relaxation and between relaxation and con-traction are the same. This same rate is evident in the terminal whine (fig. 7),and in the grouped pulses of the buzzy portions of calls (fig. 5). The secondNo. 3 ACOUSTICAL BEHAVIOR OF FIDICINA PRONOE 115pulse rate is 340 to 350 pulses (170 to 175 contractions) per sec as seen in theirregularly produced phrases of disturbance squawks (fig. 8). Here the intervalbetween in- and out-pops of the timbal remains the same, but the interval betweentimbal-muscle contractions is more variable and is increased by about x/?> (0.0054to 0.0062 sec).Phrase-programs involve fairly regular repetition of similar units. The con-secutive buzzy phrases of the repeated units of calling songs, each containing 4to 6 pulses (fig. 5), apparently depend on a regular delay in timbal-muscle con-traction following two or three contractions at the normal rate. It is this delay,FIGURES 1 to 4, Audiospectrographs of sounds of Fidicina pronoe: 1) calling song, part ofan early phrase; 2) calling song, beginning of the terminal whine; 3) calling song, conclusion ofthe terminal whine; 4) disturbance squawk. All analyses at 3^2 HL BW PN (Moore, 1961).plus the lowered intensity and the associated lowered abdominal position (chang-ing the timbal tension and resonating properties of the abdomen), that gives thebuzzy quality to this part of the call. The time-lapse between the final con-traction of one buzzy phrase and the first contraction of the next varies from 0.011to 0.019 sec, buzzy phrases being produced at this temperature at a rate of 40 to50 per sec. Short whines contain 14 to 30 pulses and are repeated at a rate of3.9 to 4.2 per sec, while the longer whines contain around 90 pulses and are re-peated at a rate of 1.3 to 1.4 per sec in calling songs. The phrases of disturbancesquawks are quite irregular in duration and in repetition rate.116 THOMAS E. MOORE Vol. 62The bell-like effect of portions of the calls of Fidicina pronoe is apparentlycaused by sudden beginning and ending of intense multi-pulse phrases with anarrow band of frequencies. Pringle (1954) described the song of the Ceylonesecicada Purana campanula as bell-like, and discussed in some detail the factorsresponsible for its quality.The intense whine-like portions of calling songs are accompanied by a raisingof the abdomen, as has been described for other cicadas (Pringle, 1954). Thecontrol of rhythm, intensity, and quality depends upon the interaction betweenFIGURES 5 to 8, Audiospectrographs of sounds of Fidicina pronoe: 5) calling song, part of anearly phrase; 6) calling song, part of an early phrase; 7) calling song, part of terminal whine;8) disturbance squawk. All analyses at Y2 HH BW PX.the central nervous system and a poorly-understood complex of muscles thatcontrols: 1) timbal contractions, 2) shape, position, and resonance of the abdomenand its tracheal air sacs, and 3) tension of the associated timbals, tympana, andfolded membrane (Pringle, 1954). The relatively low rate of repetition of phrasessuggests that these may be under central nervous control, while the timbal-musclecontraction rate suggests that timbal vibration may be under myogenic control(Bullock, 1961). Voskresenskaya and Svidersky (1961) have suggested a neuro-genic control of "high-frequency" muscles in insects involving frequent rhythmicreactions in which the sympathetic nervous system participates.No. 3 ACOUSTICAL BEHAVIOR OF FIDICINA PRONOE 117Fidicina pronoe is a relatively large cicada with contrasting green and blackmarkings (fig. 9). In males, the body length is about 34 mm and the forewinglength is approximately 43 mm. The configuration of the tenth abdominalsegment (fig. 10 to 11), especially the caudal protuberance and the very short,stout setae on the caudo-ventral surface, and the form of the aedeagal apex shownin the same figures are diagnostic. The right timbal of F. pronoe is shown in fig.12; the shape and position of its sclerotized ribs are diagnostic.In our collection there are several specimens from other localities in Mexico,Guatemala, Colombia, and Costa Rica which closely resemble the Cd. Alemanspecimens in size, coloration, and structure of the tympana. However, there areFIGURES 9 to 12, Structures of Fidicina pronoe: 9) male, dorsal view of anterior portion;10) ninth and tenth abdominal segments and aedeagus, lateral view; 11) ninth and tenthabdominal segments, caudal view; 12) right timbal, posterior view, timbal cover removed.slight differences in the structure of the ninth and tenth abdominal segments, theaedeagus, and the timbals. One series of specimens from near Arriaga in Chiapas,Mexico, was so similar morphologically that I would have assumed it to be con-specific with F. pronoe were it not for the following notes made by Dr. I. J. Cantrallat the time of collecting: " . . . sings with a coarse, . . . hollow, tone—as teeezeee, teee zeee—each expression is given louder and louder until the insect sings a118 THOMAS E. MOORE Vol. 62continuous, ringing, but hollow zeee." This description would fit my first im-pression of F. pronoe singing in the distance as heard over the truck's roar, butis not at all similar to its four-part, Morse code-like "V" which is readily dis-tinguished at close range (as Cantrall was when collecting the specimens). Inthe light of the importance of calling songs as specific aggregating mechanisms incicadas (Alexander and Moore, 1958), these Arriaga specimens and those examinedfrom other localities in Mexico and Central and South America most likely representother species, albeit very similar siblings, whose description is best left until veri-fication of specific identity can be made from field studies of the interactions andacoustical behavior of these populations in nature.F. pronoe has previously been reported from numerous localities in surprisinglyremote regions by several authors. Walker (1850) described the species withoutlocality, and later (1858) described Cicada ( = Fidicina) compacta, also withoutlocality. Stal (1854) described Fidicina vinula from Brazil, which he later (1862)synonymized, along with Walker's F. compacta, under F. pronoe. Distant (1881)listed in addition Mexico, Guatemala, Costa Rica, Panama, Colombia, Trinidad,and Brazil. Uhler (1892) added Cordova and Orizaba, Veracruz, Mexico, andSouthern Texas, U.S.A. This latter "soiled specimen," which "seems tobelong to this species," is most likely Tibicen superba, a common, green,similar-appearing, summer species in Texas. F. pronoe in southern Texas wouldbe several hundred miles north of the northernmost record for any other specimensof what has been called "pronoe." Gibson and Carrillo (1959) listed onlyCotaxtla, Ver., Mexico, for F. pronoe. As there is no type locality for Walker'sF. pronoe, and as Mr. Izzard (in litt.) feels that the figures of dorsal colorationand of structure of the ninth and tenth abdominal segments and aedcagus pre-sented here agree with Walker's type specimen, I am applying the name F. pronoeto the specimens recorded and collected in Oaxaca and Veracruz near confluents ofof the Rio Papaloapan until such time as specimens of another species which moreclosely resembles the type are found. My drawings were made from specimenspreserved in 70 percent ethyl alcohol, and the posterior view of the right timbal(fig. 12) apparently does not resemble very closely what can be seen of the partiallyconcealed, and perhaps damaged, timbal of the pinned type (Izzard, in litt.).SUMMARYMales of this green tropical cicada sing primarily at dusk and during otherperiods of low light intensity. The ringing musical phrases of males suggest the". . . —" of "V" in Morse code, while distant songs sound very much like thetwo-part peals of small, hand-shaken bells. Of special interest are the two ratesof timbal-muscle contraction, one common to all calling song phrases (220 to 230per sec), and the other found in disturbance squawks (170 to 175 per sec). Atthe end of twilight chorusing males and females fly into close proximity to the lastmales to stop calling. What has been called one species, F. pronoe, is probablya complex of several species extending from central Mexico into northern SouthAmerica. This anomaly can only be unraveled through analysis of distribu-tion and of acoustical behavior of populations through field studies.ACKNOWLEDGMENTSI am particularly indebted to Sr. Jordi Julia Zertuche, Entomologo, and Ing.Jose Ramos Magafia, Vocal Ejecutivo, Comision del Papaloapan, Ciudad MiguelAleman, Veracruz, Mexico, for providing living quarters and transportation inMexico. Dr. William E. Duellman, Gottfried Hogh, and Gerald Wilson assistedin collecting and recording the cicadas, and Mr. R. J. Izzard kindly comparedcopies of the illustrations with Walker's type of Cicada ( = Fidicina) pronoe inthe British Museum of Natural History collections. Drs. Theodore H. Hubbell,Richard D. Alexander, and Laurence C. Stuart gave valuable criticisms of theNo. 3 ACOUSTICAL BEHAVIOR OF FIDICINA PRONOE 119manuscript. This research was aided by a Faculty Research Grant from theHorace H. Rackham School of Graduate Studies of The University of Michigan,Ann Arbor, Michigan.REFERENCESAlexander, R. D., and T. E. Moore. 1958. Studies on the acoustical behavior of seventeen-yearcicadas. (Homoptera: Cicadidae: Magicicada). Ohio J. Sci. 58: 107-127.Bullock, T. H. 1961. The origins of patterned nervous discharge. Behavior 17: 48-59.Distant, W. L. 1881. Family Cicadidae. Biologia Centrali-Americana 48: 16.Gibson, W. W., and J. L. Carrillo S. 1959. Lista de insectos en la Coleccion Entomologica dela Oficina de Estudios Especiales, S. A. G. Folleto Miscelaneo No. 9: 35.Moore, T. E. 1961. Audiospectrographic analysis of the sounds of Hemiptera and Homoptera.Ann. Ent. Soc. Am. 54: 272-291.Pringle, J. W. S. 1954. A physiological analysis of cicada song. J. Exptl. Biol. 31: 525-560,21 fig., 1 pi.Stal, C. 1854. Foredrag. I. Nya Hemiptera. Ofv. Vet.-Ak. Forh. 11:242.. 1862. Synonymiska och systematika anteckningar ofver Hemiptera. Ofv. Vet.-Ak.Forh. 19: 482, 500.Uhler, P. R. 1892. Preliminary survey of the Cicadidae of the United States, Antilles, andMexico. Trans. Md. Acad. Sci. l(n.s.): 148.Voskresenskaya, A. K., and V. L. Svidersky. 1961. The role of the central and sympatheticnervous system in the function of the tymbal muscles of cicadas. J. Insect Physiol. 6(1):26-35.Walker, F. 1850. Cicada pronoe. List of the specimens of homopterous insects in the collectionof the British Museum. 1: 144.. 1858. Cicada compacta. Insecta Saundersiana, Homoptera. p. 14. John VanVoorst, London, p. 14.

Acoustical Behavior of the Cicada Fidicina Pronoe (Walker) (Homoptera: Cicadidae)

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Acoustical Behavior of the Cicada Fidicina Pronoe (Walker) (Homoptera: Cicadidae)

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