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Theless Remain Significantly Elevated for Periods theless remain significantly elevated for subfamilies of Acrididae: Acridinae, species. A male was approached and periods of at least 1 year. This postre- Oedipodinae, and Cyrtacanthacridinae watched as he made the noise; his man­ gression plateau is in most cases within (7, 2). In most cases the sounds appear dibles could be seen moving in time the 2000 to 6000 unit level, which cor­ to be no more than relatively non­ with the sound. Individuals spaced a responds quantitatively to the first host- specific reactions to disturbances, pro­ few feet apart seemed to be responding response plateau seen shortly after duced by nymphs and adults of both to one another by repeating series of tumor implantation but prior to the sexes when they contact one another ticks in rapid succession, each one be­ appearance of the tumor. or when they are disturbed by the ginning about a second after his nearest Since the values for lactic dehydro­ activities of other animals. However, neighbor had finished. My attempts to genase activity in plasma have re­ in Calliptamus italicus (L.) (Cyrta­ get a response by tapping various metal sponded to the successful treatment of canthacridinae), several mandibular objects together were unsuccessful until the established tumors which we have noises occur as significant signals in finally a nearby male delivered a series tested, and since they have also re­ situations similar to those which are of ticks immediately following an imi­ flected tumor inhibition prior to visible regulated by tegmino-femoral stridula- tation made by striking a metal ther­ or measurable changes in the tumor tion in the Acridinae and Oedipodinae. mometer case against a brass belt buck­ mass, these enzyme methods are being Mandibular sounds are produced by le. In each of many subsequent trials, explored to determine their potential adults of both sexes when they are dis­ the insect responded to the imitation usefulness as additional indices in the turbed, and by males during aggressive after an interval of about 1 second (0.9 screening of antitumor compounds encounters with other males, during to 1.0 second in five tape-recorded and of extending the understanding of courtship, when courtship is interrupted, trials). This was the same interval as tumor-host-enzyme relationships (7). and during copulation (2,3). The single was occurring between successive series VERNON RILEY finding that keeps this series of situations of ticks by neighboring grasshoppers, FELIX WROBLEWSKI from paralleling those in which tegmino- and an irregular juggling of the time of Divisions of Experimental femoral stidulation is significant in Acri­ delivery of the imitation left no doubt Chemotherapy and Clinical dinae and Oedipodinae is that C. italicus that the grasshopper was responding Investigation, Sloan-Kettering has no calling sound—no signal pro­ to it. Institute for Cancer Research, duced by lone males in the absence of The mandible-snapping of Paratylo­ New York, New York other individuals which results in the tropidia brunneri is a simple sound, re­ coming together of the sexes or has sembling a low-intensity abbreviated References and Notes any of the other side effects of this sig­ version of the ticking song of the katy­ 1. B. R. Hill, and C. Levi, Cancer Research 14, nal in various Orthoptera and Homop- did, Microcentrum rhombifolium (Saus­ 513 (1954); K. M. Hsieh, V. Suntzeff, E. V. tera (see 4). Cowdry, ibid. 16, 237 (1956); C. Manso, K. sure) (4). It is audible from a distance Sugiura, F. Wroblewski, ibid. 18, 682 (1958); On 20 June 1959 I tape-recorded a of several yards. Audiospectrographic V. Riley and F. Wroblewski, Federation Proc. 18, 310 (1959). mandible-snapping noise made by Para­ analysis shows that the ticks have a 2. F. Wroblewski, Cancer 12, 27 (1959). tylotropidia brunneri Scudder (Cyrta­ nearly continuous frequency spectrum 3. A. E. Moore and F. Wroblewski, Proc. Soc. canthacridinae), which is not only the up to at least 15 key/sec, with intensity Exptl. Biol. Med. 98, 782 (1958). 4 V. Riley, in preparation. first sound recorded for this species but peaks at about 3, 5, and 8 key/sec. The 5. F. Wroblewski and J. S. LaDue, Proc. Soc. also appears to represent a close paral­ ticks are delivered at rates of 6 to 7 Exptl. Biol. Med. 90, 210 (1955); O. Warburg, Die Wasserstojfubertragenden Fermente (Saen- lel of the calling sound in other Orthop­ per second (7); of 16 tape-recorded ger, Berlin, 1948). tera and in Homoptera. This large grass­ series, 12 series were comprised of 4 6. V. Riley, ibid. 100, 155 (1959); "The Melano­ hopper was abundant in a hill prairie ticks each, and the other four, of 2, 3, ma as a Model in a Rational Chemotherapy Study," in Pigment Cell Biology (Academic along the crest of the Mississippi River 5, and 6 ticks each (8). Press, New York, 1959), pp. 389-433. bluff south of Valmeyer, 111., in Mon­ Every aspect of this observation sug­ 7. We are deeply indebted to Editha Huerto, Carol Ross, David Bardell, and Frank Lilly roe County. A similar prairie just south gests that the ticking of P. brunneri— for their assistance in these studies, and to of this one has been described and illus­ produced by lone males and elicited Henry Abrahams, Alan F. Arnold, Marie trated (5). The dominant plant is An- consistently by auditory stimuli—is Bunker, Eleanor Havesi, and Claude Arpels of the Sloan-Kettering Institute Volunteer De­ dropogon scoparius Michx.; there are functionally analogous to the calling partment. We also wish to thank C. Chester occasional clumps of A. gerardi Vit- songs already known for Acridoidea, Stock for support and counsel. man present, along with several other Tettigonioidea, and Auchenorhynchous 21 March 1960 native prairie plants and animals. Dur­ Homoptera. This is a significant addi­ ing the day in late spring and early sum­ tion to our knowledge of insect acous­ mer, the principal sounds in the prairie tics, representing another instance of are the calling songs of three Acridinae: parallel evolution in the development Communicative Mandible-Snapping Chloealtis conspersa Harris, Pseudopo- of long-range sound signals. Further ob­ mala brachyptera (Scudder), and Eritet- servation on this species is likely to re­ in Acrididae (Orthoptera) tix simplex (Scudder) (6). The tiny veal that mandibular sounds function in Abstract. Paratylotropidia brunneri grassland cicada, Beameria venosa (Uhl- several situations, as they do in Callip­ Scudder is the first insect known to er), and three largely nocturnal crick­ tamus italicus. possess a long-range mandibular sound ets, Acheta fultoni Alexander, Miogryl- Communicative mandible-snapping signal. This signal probably evolved lus verticalis (Serville), and Oecanthus has probably evolved in every case through a stage in which feeding noises argentinus Saussure, were the only other through a stage in which the noise were significant; it is believed to be a singing insects heard in the prairie on made by feeding grasshoppers was the functional analog of other insect calling three separate visits at this time of year. initial auditory stimulus. Visually sig­ sounds. This is perhaps the only habitat in nificant motion of the mandibles seems Many insects with chewing mouth- eastern North America in which slant- a less likely precursor, though it may parts make audible noises while feeding, faced grasshoppers are at any time the have appeared as an intermediate stage but only among the short-horned grass­ dominant noisemakers. in some cases; Acrididae are generally hoppers are cases known in which Several Paratylotropidia brunneri most active in bright sunlight, and vision sounds made by movements of the were collected before it was discovered is important in their close-range behav­ empty mandibles operate as intraspecifie that series of soft ticks heard almost ioral interactions. Lepiney (9) has communicative signals. This ability has continually here and there across the shown that the odor of crushed leaves appeared in scattered genera in three prairie were being produced by this acts as an attractant to migratory lo­ rn SCIENCE, VOL. 132 custs, and this could have been a pre- known to be at equilibrium with respect Table 1. Comparison of rates of dialysis of cursory stage to aggregation through to the alpha and beta anomers were certain sugars in water and alkaline solutions. The values shown are the rate of dialysis in response to feeding noises. dialyzed through cellophane (19 mm alkaline solution minus rate in water. The mandibles of Paratylotropidia diameter when round, No. 10886, Will brunneri show no special modification Corporation) into 70 ml of the indi- Alkaline concentration Sugar indicative of a role in sound produc- cated solvent at room temperature. 0.01N 0. ION tion. Because of the importance of man- Every effort was made throughout to dibular structure in feeding, it seems keep the dialyzing surface at approxi- Sodium hydroxide sohction D-Glucose 3.2 9.1 unlikely that mandibular sounds could mately 52 cm" The procedure outlined D-Galactose 4.1 10.4 ever become as extensively elaborated by Craig (2) was employed. The D-Mannose 11.4 29.0 as the tegminal and tegminofemoral selected time for dialysis, 45 minutes, D-Arabinose stridulations of other Orthoptera. It is L-Arabinose was the approxinlate half escape time D-Xylose probably significant that P,hrunneri oc- for most of the sugar solutions. All L-Xylose curs in a habitat where there are few analyses were by means of the o-amino- S~icrose other sound-producing insects, and biphenyl procedure (4). An analysis Maltose where a soft, simple sound is more was considered satisfactory when the Cellobiose likely to become an effective long-range total sugar calculated from the concen- hydroxide solutiorz 9.5 signal.
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