1:) S yC I-I F.

THE SCALES OF COLEOPTERA.

BY GEORGE DIMMOCK, CAMBRIDGE MASS. Altho the following paper is mainly scales of lepidoptera in their works. a description of a few forms of scales During the preselt century the literature of coleoptera, on some of which the of this subject has increased rapidly, scales have not before been noticed, and and anaong the authors of leading on others of which they have been more papers which treat mainly or consider- or less fully described, a brief outline ably of the scales of lepidoptera are, in of the history and bibliography of the chronological.order, Deschamps (I 835) , knowledge of the scales of in Bowerbank (838), Craig (I839), general and of coleoptera in particular Ratzeburg (i84o), de la Rue (I852), may not be amiss as an introduction to Semper (I857) Kettelhoit (i86o), these descriptions. According to Mayer and Schneider Deschamps, B. Recherches microsco- the scales of lepidoptera were first piques sur l'organisation des ailes des ldpi- mentioned by Fabricius, in 16oo, were doptres. (Ann. sci. nat., 1835, s. 2, v. 3, P- later mentioned by Malpighi, in I65o 111-137.) and since then by many other observers. Bowerbank, J. S. On the structure of Up to the beginning of the present the scales on the wings of lepidopterous in- sects. (Entom. mag., v. century the literature of this subject is 1838, 5, P. 3o0-304.) E. the of little Craig, On configuration of the importance except historically, scale of butterflies' wings, as exhibited in and I will cite only the names of the microscope. (Edinb. philos, mag., 1839 Bonanni, Ledermfiller, R6aumur, RSsel s. 2, v. 15, p. 279-282, fig.) and Swammerdamm, who made men- Ratzeburg, J. T. C. Die Forstinsekten... tion, to a greater or less extent, of the Bd. 2, 184o. de la Rue, W. On the markings on the Mayer, F.J.C. Ueber den Staub der scales of Amalkusia korsfleldii. (Trans. mi- Schmetterlingsflfiigel. (Allgem. reed. Cen- cros. soc. Lond., 1852, v. 3, P. 36-4o, pl. 2.) tral-Zeit., 186o, jahrg. 9, P. 772-774 -) /5"a- Semper, C. Beobachtung fiber die Bildung en, Bibl. entom. der Fltigel, Schuppen und Haare bei den Schneider, R. Die Schuppen an den ver- Lepidopteren. (Zeitschr. f, wiss. Zool., schiedenen Fltigel- nd K6rpertheilen der 185;7, v. 8, p. 326-339, pl. 13. Lepidopteren. Dissertatio... Halls Saxo- Kettelhoit, T. De squamis lepidoptero- num, 1878. Also (Zeitschr. f. ges. Naturw., rum. Dissertatio...Bonnae, 186o. I878.) PS T'CttE. [Jam--Feb. 8S3.

Mayer (I86O) ,1 Landois ( 87 ),1 and (878) and Aurivellius (x88o). =1 Schneider (878). Even special Weismann believes that it is not impos- modifications of scales, called by sible that these scales give out an etherial Scuddern androconia, have been found oil secreted by the cells at their bases. on the males of a large number of Without discussing the correctness butterflies, and have given rise to con- Weismann's view, the extensive litera- siderable discussion in regard to their ture devoted to the scales of lepidoptera, function. Androconia were first discov- of which I have given only the outline, ered about 8z 5 by Baillif, who termed shows how broad the subject is. But them plumulae. They have been how is it with the scales of insects discussed since in papers by numerous other than lepidoptera? Are forms as writers, among whom may be mer- interesting as androconia waiting the tioned Deschamps8, Schneider, Watson search of thoro investigators? (I865-869),2-6 Wonfor 1868-1869) Leeuwenhoek, in 68o, and Swam- Anthony (87z), >x8 Fritz Mfiller merdamm, in the next century, figure (877),1 Scudder (877),n Weisnqann the scales of the wings and body of Clex, and Weismann, '2a in 1864, speaks 10 Landois, H. Beitrige zur Ent,vicklungs- of scales on Sarc@haga car,caria. geschichte der Schmetterlingsfliigel in der These are all the references which I Raupe und Puppe. (Zeitschr. f. wiss. Zool., know to diptera having scales and I i871, v. 2I, p. 3o5-316, pl. 3.) Scudder, S. H. Antigeny, or sexual v dimorphism in butterflies. (Proc. Amer. Anthony, J. The markings on the battle- acad. arts and sciences, 877, v. 12, p. dore scales of some of the lepidoptera. (Mo. 58.) micros, journ., I872 v. 7, P. I-3, pl. -2.) s Watson, J. On certain scales of some Anthony, J. Structure of battledore diurnal lepidoptera. (Mem. Lit. and phil. scales. (Mo. micros, journ., 1872 v. 7, P. soc. Manchester, 1865, s. 3, v. 2, p. 63-7o. 250.) a.Watson, J. On the microscopical exam- Mfiller, Fritz. Ueber Haarpinsel, F'ilz- ination of plumules... (Entom. mo. mag., fleck und hnliche Gebilde auf den Flfigeln miinnlicher Zeitschr. I865, v. 2, p. -2, fig.) Schmetterlinge. (Jena. f. Naturw., bd. x, p. Watson, On the plumules or battle- 877, 99-I4.) J. 0 dore scales of lycaenidae. (Mere. Lit. and Weismann, A. Ueber Duftschuppen. phil. soc. Manchester, 1869, s. 3, v. 3, P. I28- (Zool. Anzeiger, 879, Jahrg. 1, p. 98-99 .) 33, pl. 1-3.) 1 Aurivellius, C. Ueber sekundire Gesch- Watson, J. Further remarks on the lechtscharaktere nordischer Tagfalter. (Bi- plumules or battledore scales of some of the hang till k. Svenska vet.-akad, handlingar, lepidoptera. (Mem. Lit. and phil. soc. Man- 188o, bd. 5, n :o 25. chester, 869, s. 3, v. 3, P. 259"269, p1"5-7') Swammerdamm, J. Buch der Natur 6 Wonfor, F.W. On certain butterfly Leipzig, x752. scales characteristic of sex. (Qmrt. journ. Weismann, A. Die nachembryonale micros, sci., I868, n. s., v. 8, p. 8o-83, pl. 1; Entwickelung der Musciden (Zeitschr. 1869, v. 9, P" x9-2, P" 426"428") t: wiss. Zool., 1864, bd. 14, p. 87-336.) an.mFeb. 1883.] PS NCHE. will add here to their number, that I coleoptera, the literature of which I have found scales upon the legs of a have, as far as possible, seen and species of 2aloas from Germany. studied. Scales have been examined, but not The earliest mention that I have extensively studied, which were ob- found of scales on coleoptera is in 1762 tained fi'om thysanura (Le2bisma Ma- by Geoffroy,8 who not only mentioned 24 chilis and todura). L. Landois scales on several curculioidae, but speaks of scales in Phthirius, but it is also noticed those of dermestidae and evident fi'om his description that they scarahaeidae. The next notice of are not homologically and structurally scales of coleoptera is in 1773, by Dru- like the scales of lepidoptera. Leydig, 25 ry, in his description of Elimus to whose paper I shall have occasion imperialis, where he alludes to the to refer later, mentions scales resem- scales upon this species of czrculion- bling those of lepidoptera on spiders idae. In 777, Lindenberg8 figured of the genus Salticus. Claus6 says and briefly described Entimus imperi- the 2bhryganidae are with hairy or alis and the scales which render it so scaly wings," but I know of no special brilliant. In 178o the same author8 studies made upon the scales of these gave quite an extended description, ac- insects. As far as I have been able to companied by colored figures, of the discover, scales have only been recorded scales of Enlimus. Lindenberg's last on hemipterous insects in the case of paper is partly devoted to a curious the curious dimorphic form of A2bhis consideration of the question why in- aceris (originally described by Thorn- sects and small , some of them ton, in 85z. as Phyll@horus testudi- requiring a microscope to reveal their natus) which is figured and briefly beauty, were made even more beautiful described by Packard, and have never than larger animals. Since the above- been recorded fi'om hymenoptera. mentioned papers were published, many I come now to what is more strictly the subject of this paper, the scales of e8 Geoffroy, E. L. Histoire abreg6e des in- sectes qui se trouvent aux environs de Paris 4 Landois, L. Untersuchungen tiber die v. I, 1762. [See especially p. 69, 78-79, auf dem Menschen schmarotzenden Pedicu- 114, I 5, 277, 282-83, 288, 289, 293, 293 295 linen. Anatomie des Pktkirius inguinalis and 299. Leach. (Zeitschr. f. wiss. Zool., 1864, v. 14, Drury. Illustrations of natural history p. 1-41, pl. I-5.) v. 1, 1773. Leydig, F'. Zum feineren Bau der Ar- 0 Lindenberg. Beschreibung eines bra- thropoden. (Mtiller's Archiv, 1855, p. 376- silischen Rtisselkfers. (Der Naturforscher, 48o, pl. I-I8.) lotes Sttick, Halle, 777, P. 86-87. fig.) * Claus, C. Grundzifge der Zoologic. 4re sl Lindenberg. Ausffihrlichere Beschrei- Aufl. 188o. bung des.., brasilischen Riisselkifers, nebst Packard, A. S. Guide to the study of einigen Betrachtungen. (Der Naturforscher, insects... Salem, 1869 p. 5o-51. I4tes Stfick, Halle, 178o, p. 211-22o, fig.) PS 'CttE. [Jan.--Feb. 883, popular descriptions and brief notes viously noted, between hairs and scales upon the scales of coleoptera have been of coleoptera. printed, especially in handbooks for microscopists, but the important con- tributions to the subject are in Dujar- HAIRS OF CICINDELA DORSAL1S. din's Manual for the microscope, 82 in The white hairs which clothe the Deschamps' Researches on the elytra sides of the thorax of C. of coleoptera8 and lastly in Fischer's dorsalis and are abundant all somewhat extensive dissertation84 on the upon nearly parts of the under side of scales of coleoptera, published in this , even upon 846. its and some of its mouth- Fischer's dissertation was based upon legs upon parts, owe their white color to the a large collection of coleoptera of Eu- pres- ence of air rope, which he examined and consid- in their interior. In ered by families. He classified the transverse section these hairs for scales of curculion[dae into four groups they are scarcely flat enough to be termed scalesare and made a fifth group of the kind of circular, ellipsoi- dal in or with a scales found on AntTreus. These (as fig. , d) slight to be divisions will be considered later, in tendency triangular. The central connection with the form and structure of the scales. I will begin the descriptive part of my paper with an explanation of the hairs of Cicgndela dorsalis, for I wish to say a good deal about scale-like hairs in this paper, since the scales of coleoptera are simply flattened hairs of a more or less complex nature. In the progress of this paper I hope to be able to point out aflqnities, not pre- Fig. Hairs of Cdcddela dorsalfs: a from thorax; partly deprived of air; c, apical end of same; a Dujardin, F. Nouveau manuel complet transverse section of same; e basal end of same; gr, hair from antenna. F_.nlargement: a, and gr, l'observateur au Avec atlas, de microscope. dam. c, d and e, 3o0 diam. Paris, 1843. aa Deschamps, B. Recherches microsco- portion is a canal, about one-fourth the piques sur l'organisation des dlytres des diameter of the whole hair, and is filled coldoptres. (Ann. sci. nat., 1845, s. 3, v. with air. This canal is surrounded by 3, 354-363.) P" very minute cavities forming a sort of a4 Fischer, L. H. Microscopische Unter- substance with air suchungen tiber die Kiiferschuppen. Disser- pith-like filled and tation Freiburg, 1846, fig. Reprint (Isis, extending outward to the outer chiti- 1846, v. 6, p. 4o,-41, fig.) nous covering of the hair. The cavi- Jan.--Feb. 883.] PS ICtIF_.. ties or interstices of the pith-like por- The expulsion of the air fl'om these tion open into the central canal of hairs, when they were broken at any the hair, but are prevented from direct point, was readily effected by chloro- communication with the outer air by form or alcohol, and scarcely less readily the thin sheath of chitin which forms by glycerin, by turpentin, or even by the outer covering of each hair. Each water, but if the hair had not been in- hair is closed at the apex (fig. I, c) jured at any point the air was not driven by this outer, impervious chitin layer out by any of these reagents, even after and at its basal end (fig. I, e) by a thick- several days action. When the air has ening of tle chitin sheath to such an been expelled from a hair, the latter be- extent as to entirely close the central comes transparent, and is colorless with canal, with its surrounding pith-like directly transmitted light in all parts substance, from the outer air. This except where the chitin is thickened to structure, together with the mode of close the base; this chitin is slightly closing of the basal end, leaves the brown. By obliquely transmitted light, nature of the development of the hair obtained by Aub6's illuminating appar- very apparent. It is a closed sacciform atus, the hairs which have been deprived appendage of the external chitinous cov- of air exhibit a slightly bluish shade. ering of the insect, as are the scales of Fig. , , represents a hair of which the lepidoptera, coleoptera and diptera. distal end has been deprived of air, the The knob formed by the basal end (tg. part fl'om x to y is partly deprived of , e)of the hair is inserted into a pit air and the basal portion is still filled in the chitin covering of the insect, with air. Sometimes, especially when almost exactly as the corresponding glycerin or turpentin is used to expel the part of the scales of other insects are air, a part of the air will remain col- attached. lected in the central canal (fig. , x to y) The structural points above described quite a while af:er it has left the cavities- were obtained in two ways;first, by of the pith-like portion. The clearly sectioning the hairs with the microtome, defined outline of this column of air led and, second, by carefully observing to the suspicion of an open canal, the under the microscope the action of dif- presence of which was later proved by ferent reagents on the hairs. I obtained sections. a few good transverse sections of hairs The hairs from different parts of C. taken from the sides of the thorax, and, dorsalis vary little in form and size. by examination of these sections, veri- Those from the thorax and from the un- fied the existence of an open central der side of the abdomen are club-shaped canal. Fig. , d, shows one of these (fig. , a and b), are from o.z8 to o.35 sections which was about o.oi ram. in ram. in length, and from o.o5 to o.oz 5 thickness and about o.oz ram. in aver- ram. in diameter. They taper gradually age diameter, drawn with the camera. and slightly from the middle toward PS 2"CITE. [Jan.--Feb. I883. both ends the apex is usually truncated white portions of C. dorsaNs owe their (fig. 1, c). The basal part forms a creamy whiteness to the hairs described slight neck just above the point of inser- above; these hairs are set on shining, tion (fig. I, e). These hairs are but cupreous or green-bronze surfaces. slightly curved, but are so inserted as to The hairs upon the sides of the thorax lie nearly flat on the surface of the insect. of C. vulgaris and C. Surilaca are The hairs (fig. I, g) from the anten- similar in structure and general form to nae are the smallest of the white hairs hairs from the same region in C. dor- on C. dorsalis. They are only about salis, but they are smaller in C. 2uritana o.o7 ram. long and scarcely o.oI ram. (o. I8 ram. long by o.oI ram. in diana- in diameter near the base, fi'om which eter) and slimmer in C. vu/garis (o.55 they taper to their acute tip. The long- mm. long and about o.oI mm. in diana- est hairs are those from the labial palpi, eter) in the latter species they are not of which the second joint only is densely so abundant. hairy. These hairs are from o.3o to Taking the families of coleoptera in o.5o ram. long, and about o.o 5 ram. in their systematic order, I examined next diameter near their base, fi'om which the scales of they taper gradually to a fine, acute apex. They are cofisiderably curved and slenderly filiform. ANTHRENUS SCROPHULARIAE. Between and upon the hairs of this are yellow, amorphous masses, The figuration of the whole body and quite transparent, and apparently secre- even of the legs of this insect is due to ted fi'om the surface of the insect, altho scales which are not imbricated as are they may be remnants of some rnatrix in which the hairs are packed during the pupal state of the insect. Fischer men- tions substances of apparently similar nature in his dissertation, * and Hagen, in I882, further discusses them. These Fig. Scales of Anthrenus: a, of A. scrotbhular/ae; masses on C. dorsalis often take beauti- b, arrangement of portion of elytron; c, fully clear impressions of the hairs scales of A. varius. Enlargement: and c, diam., themselves, and are insoluble in water, b, 50 diam. alcohol, turpentin, glycerin or chloro- the scales of lepidoptera. The basal form. end of each scale is inserted in a cavity With the exception of the elytra, lab- which is at the bottom of a funnel- of rum and parts the mandibles, all shaped deepening of the chitinous sur- face of the insect, and the scales are H. A. the Hagen, On color and the arranged to a certain extent, altho rather pattern of insects. (Proc. Amer. acad. arts irregularly, in lines. Fig. z, shows and sciences, 1882, v. 17, p. 234-267. Jam--Feb. 1883.] PS their order on a portion of an elytron, paring fig. 2, a with fig. 2, c, both of the little circle about the base of each which figures are equally enlarged. scale showing the limits of the funnel- At this point may be noted the pres- shaped depression in which each scale ence of two sorts of hairs on the larvae is inserted. These scales are about of certain dermestidae, as described by o.o5 ram. long by o.o3 mm. wide. De Geer, 86 Dujardin8 andThevenet, 87.88 They are of three different colors-- and the existence of scales on the.larvae white, light brown and black--and they of Altagenus pellio, as described and all contain air. These scales were de- figured by Dujardin, 82 the latter species scribed by Fischer and I introduce and being, so far as I can discover, the only figure them here (fig. 2, a) only because coleopteron from the larva of which they furnish a good example of what scales are known. Fischer termed fibrous scales (Faser- The contain a number schuppen). of genera in which scales are the rule Of other species of derraestidae I rather than the exception, and, among only examined Dermestes lardarius and them, for the first time, comes the con- Mnthrenus varius, tbr I have had access sideration of brilliantly colored scales, to but a portion of my collection, vhich those of the genus ttopHa. I hope to examine more thoroughly later. The former species was clothed with hairs only. SCALES OF COERULEA.

This well-known European insect is SCALES OF ANTHRENUS VARIUS. light metallic blue above and silvery beneath, but when deprived of the scales The figuration of A. varius is due, to which its metallic coloration is due, like that of A. scr@hulariae, to scales it is brown. The scales of the elytra which do not inbricate. The scales of and upper surfice of the thorax are A. varius (fig. 2, c) are narrower than imbricated; those of the abdomen, legs those of A. scr@hulariae, being about and under side are not imbricated. o.o5 ram. long by O.Ol 5 ram. wide. In The scales of ]/. coeru.lea vary in form color they are either dark brown, yellow from round to ovate and lanceolate, most or white; and they are striate, but the striae, about six in number, are rather De Geer, C. M6moires pour servir i obscured by the presence of much air l'histoire des insectes v. 4, 1774, P.------in the scales until the latter are treated pl. 8, fig. 4 6. 7 with liquid reagents. The fine notching Thevenet, J. Note sur Ies polls de la larve de l'anthrenus verbascL (Ann. Soc. at the end of scales of A. varius apical entom. Fr., 874, s. 5, v. 4; Bull, p. 84, 97.) is not so evident as it is in those of A. Thevenet, J. Note sur les polls de la scrophulariae, as can be seen by corn- larve du megatoma serra. (Ann. Soc. en- tom. Ft., I874, s. S, v. 4; Bull., p. Xl.) 10 [Jan.--Feb. I883. of those from the upper side being be reflected from the scales is thus usually nearly round and smooth (fig'. brought from a different direction in 3, a), while those of the under side are regard to the scales, some of the parts more variable in shape and are always before dark become bright bluish while others just before blue become darkened, but in no case do parts of a scale which were reddish by transmitted light be- come bluish by reflected light. If the light is not excluded from above the stage of the microscope, when examin- .'"' (..L.... ing these scales by transmitted light, places will be discovered, where the scales are injured or where they are turned up at their margins, in which Fig. 3. Scales of H@l[a coerulea: a, from elytron" b, from under side of thorax; c, from femur; d, fine bright blue, or rarely green, will be structure to be in with high powers. Enlarge- seen. In all cases where I speak of ment a, and c, diam. d, 500 diam. transmitted light I have excluded the covered with spines or hairs (fig. 3, b light from above the stage of the micro- and c). The scales upon the legs are scope, and where I speak of reflected most variable of all in form; common light I have excluded light from below among them are lanceolate forms (fig. the stage of themicroscope. 3, c), covered with fine hairs. The If the scales are in any way injured average size of the scales is about o..o or cracked, as they usually are in re- 1TIITI. long by 0.05 mln. wide, and they moving them @ore the insect, water will are attached to the insect by a more or readily enter them and discharge the air less prolonged basal portion (fig. 3, fl'om them. Scales thus treated with a-c) they lie very flatly pressed upon water are, by transmitted light, sky blue, the surface of the insect. sometimes tinged with carmine red, the Most of the scales of the dorsal sur- latter color being in portions of the face of the thorax and of the elytra, scales which had not been thoroughly when viewed by transmitted light, are penetrated by the water, for while the bright canary yellow, but many of them water is entering the scales they become are tinged with carmine red. Viewed reddish for a time before changing to by reflected light, or upon a dark back- blue. Boiling the water a minute ground, the parts befbre yellow are causes all red to disappear but seems to bluish, or dark and nearly invisible, have no further action on the scales; while all the parts before reddish are their structure is not altered, as the now dark and more or less indistinct water inside such minute cavities is not (or rarely greenish if they were purplish- readily boiled. By reflected light many red before). If the stage of the tnicro- scales that have been treated with water scope is now revolved, and the light to are dark greenish. Jan.Feb. 883.] PS UCHt?,. 11

Glycerin produces nearly the same become brownish grey by transmitted effects in these scales as water does; its or reflected light; they are apparently action is, however, slower and gives charred sufficiently to lose their color more opportunity to examine the scales without having their structure greatly during the process of saturation. Light altered. transmitted through scales that are In structure these scales are readily treated with glycerin is nearly color- seen to be little flattened sacs;whereVer less, and reflected light from scales they are injured, especially if they are thoroughly penetrated with glycerin is, broken off near the base, the edges of for the most part, colorless; sometimes, the upper and lower sides appear dis- however, it is greenish in spots, and tinctly. The inner structure of these these spots are usually where a tinge of scales is not easily discoverable. They red remains, when viewed by trans- appear to be filled with a very delicate mitted light. network (fig. 3, d), which is always In absolute alcohol the scales show, reddish after the yellow has left the by transmitted light, a more reddish scales. If glycerin is used as a reagent tinge than they show in vcater; in in treating the scales their reddish net- chloroform the tinge is more purplish work, even in scales originally yellow, than in water, altho the purple is very remains long after the yellow has dis- pale; in turpentin and in oil of cloves appeared. The yellow occupies the the scales are transparent and colorless. interspaces of the network. The neg- Scales moistened with any of these work itself, under high powers, appears reagents and put over a dark or black as if it were caused by retiform designs surface are light metallic green;if dry in reddish oil between the layers of the scales are put over dark surfaces they scales, but the fact of the rea.ppearance are light metallic blue. Redried from of the color after treating the scales with water, alcohol or chloroform these scales solvents for oil, e. g'., chloroform, regain their original colors, showing shows that it is not oil. From the ap- that it is a coloration due to the struc- pearance of scales charred to different ture of the scales and not due to any extents I am inclined to think this net- pignent in them. In further proof of work to be formed by thickenings of this I exposed the scales to dry chlorin the chitin walls of the scales themselves. gas and the color remained unaltered. If this be the case, the thickenings pro- Solutions of chlorin or of hypochlorites ject only inward from these walls, and fail to destroy the coloration, for after are found in corresponding figures upon they are washed out and the scales both upper and under walls of the again dried the colors reappear as bright scales. The colors red and yellow by as at first. transmitted light may exist where only Dry scales heated slightly over flame the upper or under half of the scale is suddenly lose all their metallic colora- present. tion, and, while retaining their form, (To be couticcued.) International Journal of Peptides

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