PSYCHE.

ON SOME GLANDS WHICH OPEN EXTERNALLY ON .

BY GEORGE DIMMOCK, CAMBRIDGE, MASS.

The following paper consists, for the yellow or red,-- contains the odorous most part, of compiled material 1)rought fluid. The red tubercl(!s are seen, in into connected form in consequence of sections cut with the microtome, to be ideas suggested to me in studying 0(101'- divided into compartments, the cavities iferous glands of the hrvae of Attacus of each spine opening into a compartment cecropia, to which I have already called at its basal end. :['he spines themselves the attention of the Calnbridge Entomo- are quite rigid and very brittle, so that logical Club, at its meeting of 13 Oct. they break away at a slight touch and 1882. Since that time I have made sec- leave a hole in the tubercle, out of which tions of the above-mentioned glands of the odorous fluid pours, pushed by inter- Attact.ts cecrol)ia, and of those which I hal pressure. This fluid, which I have found later in tile larvae of a pterophorid, not examined carefully, but which I hepe Aciptihts lobidcctylus; the glands of the later to study chemically, is strongly acid larvae of these two species have furnished to litmus paper, but causes a purple the original descriptive matter of this precipitate in carmin solutions. Larvae paper. of Attacus cecropia are provided with The peculiar odor of tile larvae of Atta- these glands and the odorous fluid as cus cecropia, when they are roughly han- early as the third la.rval stage--perhaps dled, has probably escaped the notice of earlier--and apparently shed the glands but few persons who have reared these in the tubercles when moulting the last through their larval stages. If a larval skin in order to enter the pupal larva be examined carefully the black state. spines upon its red, blue, and yellow The odor given out by the glands of knobs, o1: tubercles, will be seen to break the larvae of Attacus cecropic suggests at easily from the tubercle, and a clear yel- once their protective function, and, after low fluid of disagreeable odor to ooze having watched a sparrow (Passer domes- from each opening left by the injury. ticus) drag a sphingid larva about, seiz- By crushing the tubercle with a pair of ing it usually by the horn, it seemed forceps the same strong odor is very likely to me that the disagreeable acid noticeable, and by this mode of treat- fluid in the tubercles of the larva of ment one has no difficulty in proving that Attacus cecropia was a protection to the each tubercle, small or large,blue, larva fi'om similar rough treatment. 388 PS T'CHE.

Having found these interesting glands American species.of which on the larvae of .Attacus cecropia, glands are known to sting. Lintner has experi- concerning which I can find no mention mented further upon the stinging power in entomological literature, my attention of the larvae of Lagoa crispata, and Miss was drawn further to the subject of exter- Murtfeldt upon that of the larva of naI glands of insects, many of wlfich are Lagoa opercularis. That the sting of protective or defensive in function. some of these larvae can do lasting in- Glands similar to those of tile larwt jury is certain, for my mother, when of Attacus ceero;pia, in that they have no twenty-seven years old, received so outlet until one is produced by external severe a sting in the middle finger of one agency, are not rare in the larvae of hand in brushing away a larva from her bombycidae. The severe poisoning pro- neck that tile distal joint, healing only duced by tile hairs of certain larvae of after several months, remains somewhat bombycidae, of which the so-called pro- stiffened and slightly deformed, now thir- cessionary caterpillar of Europe is an ty-seven years. For a time the stinging example and observed, according to of these bombycid larvae was attributed Moufet, by Dioscorides and other early to the action of the hairs in entering writers, is caused by tile secretion fi'om a and wandering about in tile flesh, and minute gland at the base of each hair. even as late as 1881, long after tile dis- The secretion of these glands fills the covery of the glands at the base of the hollow central portion of the hair, and hairs, Goossens advances the idea that when the sharp, often barbed, hairs are the poison of the proeessionary caterpil- broken in the flesh of attacking , lar of Europe comes from other glands the broken parts carry with them the which I shall mention more in detail poisonous secretion. This secretion is, later. Keller, in 1883, discusses the perhaps, formic acid or a formate in solu- mode of urtication in the processionary tion. Karsten, in 1848, described the caterpillars (larwxe of Gcstropacha) and anatomy of the poison glands at the base figures the glmds at the bases o' their of the hairs of an American species of hairs. Sturnia. Fine illustrations of this kind Still another tbrm of gland without of gland are to be found in the stinging any outlet until broken open, but a gland hairs of the larvae of Ityperchiria io and which can scarcel}" be classed witll those Hemileuca maia, both common insects previously mentioned, is that at the an- in parts of the United States. Lintner terior end of certai bombycid pupae, and Riley have recorded their experi- which breaks when the imago springs ments on the stinging power of these the chitinous pupal skin, and leaves its two species of larvae, and the latter secretion, which has been termed bomby- writer has given a list of the larvae of cic acid, on the head of the , the ltttter using the secretion to moisten tile 1For literature referred to throughout this paper threads of the cocoon so that they can the end of the article. PS 389 be cut or pushed aside to allow the es- Second are hairs (from 0.08 to 0.14 ram. cape of tbe imago within. I have never long) more or less dumb-bell or club studied this gland and will refer for fur- ibrmed which are filled with granular ther notice to tile easily accessible papers matter, and seem to be set usually only of Trouvelot, Packard McLaren1 and upon the surface of the chitinous cover- Worthington, wherein references can ing of the larva. Third are the longer be found to earlier European writers on hairs (from 0.8 to 1.3 mm. long) linear this subject. or slightly clavate usually burst at the It is an easy transition from the glands tip or sometimes along the sides, and of the larvae of Attacus, Hyperchiria where burst surrounded by a drop of and Hemileuca, closed by brittle hollow exuded gummy matter. These last hairs spines or hairs, to the glandular hairs are mounted by a kind of joint such as of certain larvae ofpterophoridae where is often present at the base of the hairs are apparently burst open at hairs, upon or near the summit of little their tips by the pressure of the secretion conical elevations which rise about 0.2 within them the liquid then oozing out mm. above the surface of tile dorsal and to tbrm a dew-like drop upon each hair. lateral parts of the larvae. These Zeller 1: mentions glandular hairs ('dr- hairs are arranged systematically and senh/irchen") on the larvae of Mimeseo- synmetridally upon the different seg- ptilus phaeodactylus and M. mictodactylus ments of the larva, tile most prominent but says nothing of the structure or use of them being a pair upon a conical of these hairs. Miss Murtfeldt writes elevation just at each side of the median of the larva of Leioptilus sericidactylus dorsal line of each segment. A com- 'Dorsal hairs proceeding fl'om prominent parison of the arrangement of these hairs tubercles and of two sizes in each tuft and prominences with the arrangement each of the shorter ones tipped with a of hairs and warts upon other lepido- minute pellucid bead of viscid fluid to pterous larvae, especially of those upon which pollen and bits of leaves often ad- the larvae of tortricidae would be an here." I have found the larva of Aci.pti- interesting study. The interior of hairs lus lobidactylus to be covered in like of this third form opens at the base into manner, with glandular hairs. the conical proninence or wart on which Upon making transverse sections of the hair is situated. The prominence is the larva of Aciptilus lobidactylus, its ex- probably entirely filled by the gland ternal surface is ibund to bear three which secretes the viscid matter that kinds of appendages. First are the very finds outlet through the hair. minute but obtuse spines (about 0.01 The specimens of the larva of which mm. long) which clothe most softer and I made microtomic sections were not more flexible portions of its external quite well enough preserved to admit of covering, and which are found on many carefully studying the gland at the bas larvae of difirent orders of insects. of the hairs. The hairs of the second 390 PS 2CttE. and third form seem to be modifications lobidactylus, reared from larvae taken of each other, for, in the longer and when nearly full-grown, while I have al- more clawtte hairs of the second form, ways obtained, in rearing a much less the granular matter in the hair extended number of Oxyptilus periscelidactylus without interruption into the larva, under similar conditions, several para- and these hairs often burst and give out sites. It is noticeable that the viscid viscid fluid. The larger hairs of the secretion upon the larvae of .Aciptilus second form are often upon the sides of lobidactylus is alkaline to litmus paper, the conical warts which bear at their while the fluid from the tubercles of the summits hairs of the third form. Hairs larvae of Attacus cecropia is acid. of similar nature, but smaller and with- Turning from the study of insect-hairs out viscid fluid, clothe the larva of Oxy- which furnish more or less temporary ef- .ptilees periscelidactylus and Ptero2)horus ferent ducts for glands, there are forms monodactylus. of hairs--leaving out of account, as un- Mimeseoptils phaeodactylus feeds on proved, Weismann's curious, but not Ononis repe.ns, M. mictodactylus on Saxi- improbable, view* that scales of lepi- fi'aga granulata, Lioptilus sericidactylus doptera may be sometimes ducts for an on Vernonia noveboracensis and Acgtilus odorous fluid secreted by cells at their lobidactylus on Solidago ?canadensis bases--which are regularly and perma- thus, as will be noticed, all the species nently the outlets for glands at their of pterophoridae mentioned above feed bases. An interesting example of this on plant., clothed with glandular or long kind of open hair duct is furnished by hairs. The glandular hairs of the larva the hairs upon the foot of the common protect it, probably, by causing it to re- house-fly (Musca domestica), through semble the surface of the plants on which which hairs, as recently shown by De- it feeds, a kind of resemblance not rare witz, a sticky secretion is poured, the in insects; but it is also probable that fly being enabled by the adhesion of this the secretion of the glandular hairs also secretion to rest upon smooth vertical protects the larva, to some extent, from surfaces. Leydig, in 1859, describes the attacks of ichneumons and of other and figures glands in the tarsi of several parasites, for I obtained no parasites coleoptera West, v in 1862, describes from over fifty specimens of Aci))tilus and figures the tai'sal hairs of the fly and of many other insects; and Dewitz, s in *Miss Murtfeldt writes follows: "I had not 1882, discusses the function in locomo- thought to mention it in connection wth my description tion of the viscid secretion which is dis- of Leio2MHus sercdactylus but there 's a very close imitation in the dermal clothing of the larva to that of charged by the glands of the foot through the young leaves of Vernon which the spring and the tarsal hairs, both in flies and in early broods feed."... 'Later in the season, when feeding chiefly the flowers, the larva acquires coleoptera. purplish tinge which, vith the particles of the flowers Defensive glands of another kind, the that adhere to its glandular hairs, is sufficient disguise from any but the eye practised in its detection." ducts of which open into spines, are the PS rCttE. 391 glands which furnish the poisonous se- European bombycid Hart)yia vinula, cretion for the sting of bees and wasps. raises its head and jerks it from side to I have not discussed these glands further side throwing a strongly acid fluid from in this paper, because they are rather a gland, the opening of which is on the internal than external glands; for like ventral side of the first thoracic segment. reasons I have not referred to the silk- The propulsion is here partly produced glands of lepidopterous and hymenopte- by the rapid jerking of the head, and rous larvae. The literature of impor- partly by pressure upon the receptaculum tance upon the subject of the silk-glands or lumen of the gland within the body. of lepidoptera can be found in a disser- This gland, or at least its secretion was tation by Helm, 19 in 1876; and papers noticed by de Geer in 1750 and more by Dewitz, 2-21 in 1875 and 1877, and fully in 1755 by Bonnet2 who described by Forel,2 in 1878 give a clue by which the secretion as a true acid, sharp, sour to trace the literature upon the stinging and biting. Later besides many brief organs of hymenoptera. notes on this gland, may be mentioned The glands of the larva of Attacus more extended descriptions by Mtiller, :v cecropia and Hyperchiri io, sealed Amoreux :s JSrdens9 and Rengger. a until broken open by some externa! I have not observed whether the larvae agency, may be termed passive glands, of the common American species of while those glands which pour out their Harpyia (Cerura) possess this power of secretion in direct response to some vol- throwing a defensive fluid or not and untary or reflex action of the insects as Lintner as well as French,a in descrip- do the foot-glands of Musca, the sting tions of the American larvae nake no of the bees and many other glands mention of such power. The soft skin may be termed active glands. But there of the larva of Hrpyia vinula evagin- are glands characterized by still greater ates itself about the orifiice, on the first activity than those already mentioned. thoracic segment, where the gland dis- If tile larvae of certain species of Cim- charges its fluid, forming four points bex are suddenly disturbed, they will jet which are well figured by Miller and out, as observed by Bonnet:a (v. 1, p. JSrdens, in the books cited above. 470 473, 485), a clear yellow fluid of Within the duct of the gland passes to disagreeable odor, fi'om openings along the right of the displaced central nervous their sides. In the case of a German system as described and figured by Mrs. species of Cimbex I have seen this fluid Dimmock, in PSCHE V. 3, p. 340-341. thrown by the larva to a distance of I pass by, with mention only, the several centimetres, altho Frisch, : who openings which pour out an odorous first mentions this secretion, in 1736, yellow fluid from the joints of the legs of apparently only saw it flow iom the species of coccinellidae chrysomelidae and openings. meloidae, for further notice of which I Upon disturbance, the larva of the refer to papers by Leydig (p. 37-38) 392 PS T'CtlE. who thought the fluid to be only the to some papers which will guide further blood of the insect; the glands opening to the literature of the subject, in order between the prothorax and head and to come directly to forms of which the between the mesothorax and metathorax morphology is more interesting in this in dytiscidae, of which the nature of the connection, or of which the function is secretion has been somewhat studied by not so well understood. Plateau8; the anal glands of Brachinus An interesting economical provision is and other carnivorous coleoptera, the evident in the odoriferous tubercles of the explosive mode of whose discharge in larvae of Attacus cecropia and in the urti- Brachinus has given rise to its popular caring spines of Hyperchiria io. There English name of bombardier, br the is no waste of the secreted material by literature of which I refer to Leydig16 either of these larvae; it is sealed up (p. 46-49), and to a late paper by until its use is necessary and then it is Rougement; the cornicles of the sixth applied exactly where it will accomplish abdominal segment of aphidae, which mos. But what shall one say of the are supposed to pour out a sugary mate- novel mode of avoiding waste, to which rial sought by ants, but which material attention was called by Claus, *a in the W itlaczila writes, in a paper to which larva of the European chrysomelid, Lina I will refer for further literature of the populi? Along the dorsal side of this anatomy of aphidae, comes from the larva are rows of short black spines, not anus of these insects ;* the sacs, proba- noticeably different from the spines often bly glandular in function, found by found on larvae of coleoptera, but if' the IIagena6 upon the larvae of certain neu- larva is disturbed it will suddenly press roptera the odoriferous glands of hem- out upon the tip of each of the spines a iptera mentioned by Landois and others spherical drop of milk)" liquid. Watch the scent-glands which Aub4 mentions these drops for a moment. As the dis- on the elytra and thorax of Eumolpus turbance which has caused their appear- pretiosus; the wax-secreting glands found ance subsides, they become smaller, on certain insects, treated of by Claus, sometimes suddenly, sometimes gradu- in 167; and the odoriferous glands of ally, but always in unison, until finally, phasmidae described by Scudder, in when the larva no longer feels itself in PSYCHE, V. 1, p. 137-140:all these danger, the drops entirely disappear" glands I pass by, givang reference only the fluid has been drawn back into the spines to be used again in case of need. *Buckton,:7 in vol. I, of his Monograph of British aphides, discusses (p. 39"47) "honey dew" of aphYdae Claus has suspected that salicylic acid well (p. 37-38) various waxy and silky secretions with exists in the secretion fl'om the larva of which hemiptera themselves. may add here that Gentry's3s observation, in 874 that young a/h'- Lina populi, which is very probable, since dae suck the secretion of the cornicles of older is the larva feeds upon leaves of Salix and anticipated by Morren,a9 in i836, and Morren adds to his statement "fait observ ddj par Bonnet"fact Populus, that contain salicin, of which already observed by Bonnet. salicylic acid is an oxidation-product, in other insects. Little search reveals Claus sa3"s that the larva of Linapoluli numerous forms of glands in which has been used as a source of salicylic part or the whole of the duct or the acid in small quantities. The odorous glandular surtace itself is evaginable. secretion of tile glands of the larva of This form of gland is not confined to Lia protect not only tile larva but as any one order of insects; altho first Lyonet noticed, also tile pupa from the found in lepidopterous larvae glands of attacks of birds. Claus shows how this this form have since been found in ima- protection of the pupa is accomplished: gos oflepidoptera, coleoptera and ortho- the glands, with their contents, are shed ptera. The latest writer who has dealt with the last larval skin, which remains especially with this kind of glands is around the point of attachment of the Klemensiewic, in 1883. abdomen of the pupa and, when the In this group of glands or gland-like latt,.;r wriggles about on account of any organs ma.y be classed the evaginable disturbance, the odorous fluid is squeezed osmateria of the larvae of Paipilio, the from the molted glands. De Geer4 de- protrusile tails or modified anal legs of scribes and figures the spines of the the larva of Harpyia (Cerura), tile larva of Lirta ppztli in 1775. Lyonet,4 lateral appendages of Mt.lctchbts, cer- probably somewhat earlier, describes and tain evaginaMe appendages upon the oures the larvae of Li,tapopttli and L. eleventh and twelfth segments of the doralis, and mentions the protrusion larvae of some species of Lycaena, and retraction of the milky drops, lateral evaginable appendages in a Ratzeburg mentions briefly the glan- species of Corydia, tile red protrusile dular secretion of L. pcpttli, and West- warts upon the dorsum of the larva of wood collects together notes on the O'gyia, the evaginable warts of some of larvae of different species of Lia. the stinging larvae of bmbycidae the Chal)uis and CandSze 4s write that the evaginable appendages of various tbrms larva of Lina (Plagiodera) scripga from near the anus of certain imagos of lepi- the United States is similar to that of doptera and coleoptera the organs on L. poFuli, the ventral side of the first thoracic Claus 4a was the first to study the in- segment of many lepidopterous and of ternal anatomy of tile glands and spines a few phryganeid larvae, and the pro- of Lina 'populi, and he shows that the trusile organs near the anus of larvae of liquid is pushed ont of the spines by Myrmeleon. a contraction, which I may be allowed to Organs of the sort now under eonsid- term an incipient evagination, of tile eration were first mentioned in 1602 by walls of the gland. This leads one to Aldrovandus,0 who observed and roughly search further in order to find if this figured osmateria on the larva of principle of evagination of glandtflar Popilio. Friseh (Theil 2, p. 41-42), walls is not carried to a greater extent in 1721, describes the osmateria of the 394 PS TCttE. larva of Papilio machaon, which he says seeming often to whip the whole surface "protect it from its enemies, not by their of its body. These threads or filaments strength, for they are hollow and delicate are p,'obably evaginated by internal lobules, but by the stench which comes pressure of the fluids of the body, but are from them. Madame Merian calls it a retracted by muscles within them. There pleasant odor (2d ed., pt. 1, n. 38) but is some doubt whether these threads I find it intolerable." Since the time of are really glands, and Frisch (1740, Frisch many authors have described the Theil 6, p. 18-19), who expected to find osmateria of different species of Ppil,;o, them odorous, writes that the odor among them Karsten, who described, in "must be vetS subtile and be only per- considerable detail, the anatomy of these ceptible, for the most part, to insects organs in Papilio asterias. He sass hostile to this larva" but, if these fila- they discharge "a substance, strong- ments are not glands, they have the smelling like butyric acid, by which they same coarser anatomy and should be scare away their enemies this substance considered here. reddens litmus readily, produces white The earliest description of the evagi- clouds when brought near strong ammo- nable caudal appendages of Harpyia that nia, has a somewhat biting acid but not I have seen is that by Goedart, who unpleasant taste. Mixed with water, sass of them This larva has two tails oil-like drops are seen under the mi- and when it is vexed, it will throw out croscope; neutralized with ammonia fi'om both tails a red rod or sting, which it crsstallizes." The crsstals "are it bends back, and, as if mad, vibrates not volatile upon raising the tempera- formidably, and draws it back in again." ture but decompose with disengagement Since Goedart's time these appendages of empsreumatically smelling vapors. have been described by Raumur, Caustic baryta dissolves the acid, the Frisch (Theil 6, p. 18-19), de Geer salt crystallizes in groups of very fine (Bd. 1, tab. 23, fig. 12), Bonnet,: needles." For the anatomy and his- Miiller (p. 28-30), JSrdens, 9 Meckel, tology of the osmateria of the larvae of Klemensiewicz,9 and others. Some of Pa2oilio I will refer to Karsten and Kle- these authors also describe the gland mensiewicz. previously mentioned (1). 391), as open- Each of the two caudal appendages of ing upon he under side of the first tho- the larva of Itarpyia consists of a basal racic segment of the same larva, and tube which projects backward from the Schtffer, in 1754, first shows the inti- posterior end of the larva, but which mate relationship of this gland to evagi- can be turned somewhat upward and nable glands found in the same position forward. Out of the tip of each of on larvae of other species, to the osma- these basal tubes a reddish filament can teria of the larvae of Popilio, and in be evaginated, which the larva, upon the general mode of action to the evaginable approach of danger, curves above the glands of Malachius. body and waves with a trembling motion The evaginable organs of the imago of PS ?'C/Y/?. 395

Malachius consist of a pair of reddish, Guen4e, in 1867, describes and figures trilobed organs, one on each side, be- these organs in Lycaena boetica. W. H. tween the head and the first thoracic Edwards, -' in 1878., described and fig- s(:gment, and a similar, but somewhat ured similar organs on Lyca.ena pseu- larger pair of bilobed organs, one on dargiolus and .L. comyntas, American each side, betwee'n tim metathortx and species. Scudder, a in 1881, repeats first abdominal segment. These organs Edwards' figures, referring however to are protruded by the fluids of the body- Guene as the discoverer of these organs. cavity, but have muscles for their retrac- The finction and structure of the evagin- tiou. These organs were mentioned by able organs of the larva of Lycaena are Schtffer: in 1754, by Sulzer5 in 1761, not yet settled. and since that time more or less dis- Evaginable organs were described cussed by Kirby and Spence (in whose from a blattid, Corydia caruculigera, by work may be/bun(l much that is impor- Gerstaecker, in 1861. Tile organs of tant and interesting on the glands and Corydia are yellowish white, covered odors of insects), by Westwood,v La- with hairs, and there are two of them on boulbine, Liegel, s Klemensiewicz, each side of the insect, evaginable from and by others whose papers I cannot at between the dorsal and ventral plates of present cite, as I have not collected the the first and second abdominal segments. literature of this part of my subject with These organs are present in both nale much thoroughness. The function of and female, but are not found in the these organs of Malachius is not deter- larva of Corydia. Gerstaecker suspected mined with certainty. them to be of glandular nature. Brunner The larvae of certain species of Zy- yon Wattenwyl, in 1879, described an caena have been found to attract ants, evaginable organ ou the femora of ac)'i- on account of an opening upon the dor- didae; Burgess gives an abstract of his sum of the eleventh segment, which paper in PsYCln V. 3, p. 32. These gives out a liquid apparently containing organs in acrididae and those of Corydia sugar. Upon the twelfth segment, and are the only evaginable organs of which evidently connected in function with the I have ibund mention in orthoptera. opening above mentioned, are two pro- The larvae of European species of Or- trusile organs covered with fine hairs. gyia (e. g., of O. auriflua) have long been Tile fact of ants being attracted to these known to have evaginable warts upon larvae was first observed, so far as I can their dorsum. Brief notice of these warts learn, by Esper Petzhold, in 1793, will be found in works of Schwarz, figured, and described with considerable JSrdens, Klemensiewicz, and other minuteness, the appearance of the pro- writers, but I have found only tile slight- trusile organs on tile twelfth segment of est allusions to them in American species. Zycaena biton, and nentions their occur- Fitch in describing the larva of O. nova, rence on the larvae of other species. mentions them; Coleman, s in describ- 396 PS ng the larva of 0. leucostigma: terms In quite a number of staphylinidae, them 'coral-red retractile warts"; and and more rarely in other beetles, there Coquillett, 69 in describing the larva of is a pair of evaginable organs, one on Parorgyia clinton,ii, speaks of them as each side of the anal opening, which 'reddish warts" without adding that they give off a disagreable odor when pro- are retractile. truded, and which must have attracted Evaginable warts similar to those up- the attention of most collectors of cole- on the dorsum of Orgyia are present on optera in capturing the larger species of different parts of various bombycid lar- stophylinidae. Tlaey have been men- vae, and Klemensiewicz 9 has described tioned or described by Schtffer, Du- their structure in _Leucoma salicis. I four, and Leydig TM (p. 52-53). The have mentioned their occurrence in Hy- function of these organs is evidently perchiria io on p. 352-353 of PSYCHn, V. defensive. 3, and they seem especially abundant on Similar appendages, but often more stinging larvae. Goossens regards the striking on account of their greater size, spines of the larvae of Cnethocampa to have been found in several lepidoptera, b poisonous because of a powder pro- in most cases American species, altho duced by the drying of the secretion the first mention of these organs in lepi- given out by the evaginable glands upon doptera also dates back to Schi.flir, in the dorsum of these larvae. This view 1754. I have given an outline of the seems inacceptible after Karsten,: Kel- bibliography of this subject, as far as it ler and others have so clearly proved concerns the species of lepidoptera found the presence of glands at the bases of in this vicinity, in PSYCHE, V. 4, p. 59, the spines of these stinging larvae. and need not repeat it here. Fritz The structure of the evaginable warts Miiller, v-v has contributed several pa- and their glands in Leucoma salicis is pers (in 1874, 1877, and 1883) to this well described by Klemensiewicz,9 and subject. Doubleday (1846-1850) called wil! answer in a general way for the attention to the presence of such organs similar warts of Orgyia and of the Euro- in butterflies of the genus Acraea. pean ,L(paris. The wart is protruded These organs are sometimes naked and by pressure o.f the fluids within the body sometimes clothed with hairs or scales. and retracted by muscles; at or near its The long yellow organs protrnded by centre open the ducts of one or two Spilosona acrea are good examples of glands which are situated beneath the the hair-clothed form while the organs warts. The position and general struc- described by Fritz Miiller in Heliconius ture of these glands, as well as their and allied genera are covered with scales. motion when the larvae are disturbed, The function of all these organs is not indicate that they are defensive in func- yet determined, but it is certain that tion, but exact observations are still a part of them at least are for the dis- lacking on this subject. tribution of odor. /'S 397

Mention has been made already twice of Brauer a protrusile organ beneath the of the gland which opens beneath tim first segment of a phryganeid larva. first segment of the larva of Harpyia In dealing with the protrusile organs vinula. In the same position on many of different insects above I have given larvae, some of those of Vanessa, Meli- more citations of early authors than I taea, .Argynnis, Bryophila, Cucullia, should have done had I found them Habrostola, and Cleophane, according to brought together elsewhere. Klemen- RogeuhoferT and of ,4pori, many siewicz, altho not claiming to give the tyrids, .Aplecta and Leucania according fifll literature of the subject, certainly to Goossens--is an evaginable portion omits and apparently has not seen which iu some cases, probably in some of the more important papers on is glandular, but the flnction of which tim subject of which he treats. I have is entirely undetet:mincd except in the mentioned only the more important pa- case of tta'pyia winula where it is an pers on the glands of European insects organ of defense. These organs were but I have sought to refer to all papers compared to the osmateria of Papilio which have dealt originally with evagi- larvae by Sch,tffer, and have been more nable glands ()f American insects. or less discussed by Rogenhofer,7 La- I have not examined carefully the pa- cordaireS GoossensT.79 and Klemensie- pers published upon the subject of femoral wicz. 9 Bonnets discovered these organs tufts of lepidoptera, or tufts upon other in 1739, altho he published nothing about parts of lepidopterous insects; which them, as far us I know until 1755, tufts I presume may well come into con- year af'ter the paper mentioned above sideration at this point. Fritz Mfillers was published by Scldtffer. Bonnet gives treated of them at some length in 1877. a list of thirty-one caterpillars which They have been suspected o/ten to be possess these organs, and he tried exper- organs for the distribution of odors, and iments to see if cutting off these organs Bertkaus: has shown, in one case--that in larvae affected in any way the imago of the male of Hepialus hecta-- that the produced .fi'cm them proving that ima- analogous tilaial organ is filled with gos from larvae thus treated were per- glands, and is used, in connection with fect. Bonneta (p. 503-504) also found organs on the first abdominal segment, two evginable fleshy organs near the to diffuse an odor for sexual purposes. posterior extremity of the abdomen in These organs of Hepialus are noticed the larva of the ant-lion (Myrmeleon). more fully than here, by Burgess, in Raumur: (1737, tome 3, p. 165), PS'CHE, V. 3, p. 32. notices an organ, probably similar to A few considerations upon the use to that under the first thoracic segment of which the different forms of glands, so certain lepidopterous larvae, in the same briefly noticed in this paper, especially locality, on a phryganeid larva and Ro- adapt themselves may be appropriate in genhofer, mentions, on the authority closing this paper. The simple glandu- 398 lar opening is not economical for the invagination of odor-distributing appen- application of odorous, ill-tasting or ir- dages is a double one; first when inva- ritating fluids. If it is provided with ginated their necessarily delicate surface means for throwing its secretion, as it is is less liable to injury, and, second, their in Cimbex and tIarpyia, it is somewhat surface can be kept constantly moistened more efficient at long range. The chanc- and ready for use without loss of the es, however, of a waste of the secreted odorous fluid. Besides this, too, is the matter are still great, and, as J6rdens general rule that the appendages of loco- has observed, the larva of Harpyia uses notory animals are, as far as is possible its stream-throwing gland only when in attaining the purpose for which they much disturbed. For the application of are developed, normally invaginated or an ill-tasting fluid the modified form of retracted to avoid hindrance to locomo- gland found in the knobs of the larva of tion, while the correspondbg organs of Attacus cecropia seems to be the best. sessile animals and plants attain their The secretion is set free by the very ac- surface-development by evagination. tion of the enemy of the larwt. The use Secretions which are intended to pro- of a strongly irritating fluid attains its tect by their odor usually protect by highest development iu means of injec- their taste also, for taste and smell are so ring the fluid into the flesh of the attack- nearly related senses iu higher animals, ing as is done by Hyperchiria while such glands as secrete ill-tasting and by many hymenoptera. The great- fluids, if these fluids are volatile, also est economy iu the use of an odorous usually protect by their odor. Still it fluid is attained by exposing suddenly a may be safely asserted, as a general rule, large surface moistened with the fluid to that odor protection is accomplished by the surrounding air. This is accom- evaginable organs. plished by osmateria, as in the larvae of The chemical substances proved by Papilio, and by various evaginable ap- actual test to be secreted by external pendages, in other insects. Clothed glands of insects are few in number, and, with hair these appendages expose still as I hope sometime to make this the sub- more surface to the air, and thus give ject of a special paper, I should be glad still better results in rendering the sur- to have my attention called to any secre- roundin air odorous, whether for pro- tions of insects, especially to such as tective or for sexual purposes. For sex- can be obtained in sufficient quantity ual purposes a less penetrating, often to for chemical examination. us tt less disagreeable, odor, and less of In the preceding notes I have sought it, is necessary than for protective pur- to call attention to glands of insects, in poses. It is perhaps for this reason that hope that some American students of the evaginable appendages of Spilosoma, insects, who have not enlisted ah'eady in and of some other lepidoptera, appear that army of species-describers, which to us always dry. The reason for the represents so-called American entomo- PS 399 logy by a probable majority, may be in- aLintner, J. A. Entomological contribu- duced to turn their attention to the anat- tions, 872: no. , p. x-2; no. 2, p. 36-37. *Riley, C: V. Fifth report state entom. omy, biology or physiology of American Missouri, 873, p. 25-26 3 , 34- insects. The tudy of the general anat- aMurtfeldt, Alfss M.E. An experiment Nov. can in with a stinging larva. (Can. entom., omy of insects be pursued any 876, v. 8, p. o-.o.) country, but there always remain special Goossens, Th. Des clfenilles urticantes et points in the anatomy of insects, of which quelques considdrations sur l'utilitd des ceufs pour la classification. (Ann. soc. entom. points glands and similar modifications France, I88t, s. 6, t. x, p. 23-236. of parts of insects for special purposes Keller, C. Die brennenden eigenschaften form no small part, wherein insects of der processionsraupen. (Kosmos, I883, bd. 3, P. 302"3o6, fig.) one country difier from those of any STrouvelot, L: The Amenican silk worm. other.. In such fields of study as these (Amer. nat., 867, v. , p. 30-38, 85-94, 45" American can com- 49, il.) p. 33. young entomologists Packard, A. S., jr. The mode of extrica- pet most successfully with European tion of silkworm moths from their cocoons. anatomists and can make important dis- (Amer. nat., June 878, v. ta, p. 379-383, fig.) coveries, and to researches of this kind McLaren, D. C. The mode of extrica- future entomologists who have aspira- tion of the American silk-worm moth. tions to become more than locally known (Amer. nat., July 878, v. , p. 454-456. Worthington, C: E. On the emergence must turn their attention. Leave the of lepidoptera from their cocoons. (Can. creation of scientific nmnes, quarrelling entom., Aug. 878, v. o, p. x58-59.) over synonyms among and search Zeller, P. C. Revision der pterophoriden. them (Linnaea entom., 85, v. 6, p. 39-46.)p. for new speies, to the antiquated type 356. of naturalists who have becmne so ha- aMurtfeldt, Miss M. E. New plume moths, (Amer. entom., Oct. bituated to the exterior of their tero2horgclae. sudying 88o, v. 3, P. 35"236") insects that to "destroy" a rare speci- 4Weismann, A. Ueber duffschuppen. men by dissecting it has become a crime (Zool. anzeiger, 26 Aug. 878, jahrg. , p. 98-99 to them. In their regard for the exte- Dewitz, H. Wie ist es den stubenfliegen rior of aninals, coupled with their crude und vielen anderen insecten m6glich, an senkrechten glaswRnden emporzulaufen knowledge of internal anatomy, they re- (Sitzungs-ber. gesells, naturf, freunde zu mind me of Romans, who makes a grave Berlin, 7 Jan. I882, p. 5-7.) statement in his history of Florida (p. eLeydig, F. Zur anatomie der insecten. (Archiv. f. anat., phys. u. wiss. med., 859, P. 55), that 'Anatomy has taught us, that 33-89, 49-83, pl. 2-4.) p. 35 and 38. the bone of a negroe's skull, is always lWest, T. The foot of the fly; its struc- black ." ture and action:elucidated by comparison with the feet of other insects, &c. (Trans. Linn. soc. Lond., 86a, v. 23, p. 393-42, pl. LITERATURE. 4'43') Moufet, T: Insectorum sive minimorum SDewitz, H. Ueber den kletterapparat der animalium theatrum... Lond., I634. p. insekten. (Sitzungs-ber. gesells, naturf. 85-86. freunde zu Berlin, t8 July 88, p. m9-t3. Karsten, H. Bemerkungen fiber einige gHelm, F.E. Ueber die spinndrtisen der scharfe und brennende absonderungen ver- lepidopteren. (Zeits. f. wiss. zool., t876, bd. schiedener raupen. (Archiv. flir anat., phys. 26, p. 434-469, pl. 7-28.) u. wiss. reed. 848, p. 37-38a, pl. -a.) Dewitz, H. Ueber bau und entwickelung 400 PS ICItE. des stachels und der legescheide einiger (Arbeiten zool. inst. univers. Wien u. zool. hvmenopteren und der grfinen heuschrecke. stat. Trieste, I8c;2, t. 6, p. 1-45 (397-441), 3 (eits. f. wiss. zool., 1875, bd. pl.) p. 14-18. pl. 12-13.) a%Iagen, It. A. Beitrag zur kenntnis des lDevitz, H. Ueber bau und entwickelung tracheensystems der libellen-larven. (Zool. des stachels der ameisen. (Zeits. f. wiss. anzeiger, `5 April 188o, jahrg. 3, P. 157-161.) P" zool., 1877, bd. 28, p. 527-556, pl. 26.) I6I. Forel, A. Der giftapparat und die anal- TBuckton, G: B. Monograph of the Brit- drti el der ameisen. (Zeits. t: wiss. zool., ish aphides. Lond., 1876 v. I. 1878, bd. 3o, suppl., p. 28-68, pl. 3-4.) 3SGentry, T: G The use of aphis-excre- 3Bonnet, C: Observations diverses sur tion and benefit derived therefrom. (Can. les insectes (Collection compl6te des ceu- entom., Jan. 1874, v. 6, p. 5-8.) vves de C: Bonnet, t. 1, p. 259-574 [-.at- 2 p.], aaMorren, C: Mdmoire sur l'dnigration du pl. I-6.) puceron du pcher ((@his dersicae), et sur 4I'risch J. L. Beschreibung yon allerley les caractres et l'anatomie de cette espSce. insectenin Teutschland I736, th. 4, P. (Ann. sci. natur., Zool., I836. t. 6, p. 6-93, 44. pl. 6-7. ea Geer, C. Observation sur la propridtd 4Landois, L. Anatomic der bettwanze, singul;re qu'ont les grandes chenilles qua- cimex lectularius, mit berticksichtigung ver- torze jambes et . double queue, du saule, de wandter hemipterengeschlechter. (Zeitsch. seringuer de la liqueur. (Mdm. say. dtrang. f. wiss. zool., I868, bd. I8, p. 206-224, pl. 1I- Paris, 17..;o, t. I, p. `53o-53 I, pl.) 2.) p. 218-223. aBonnet, C: Mdmoire sur la grande che- 4XAub, C: [Note sur une scrtion ftide nille t queue fourchue du saule, dans leqnel d'eumo@us brelios'us. (Ann. soc. entom. on prouve, que la liqueur que cette chenille Ft., I837, s. 1, t. 6; Bull., p. ,58.) fail jaillir, est un vdritable acide, et un acide 4'Claus, C. Ueber die wachsbereitenden trs-actif. (Mdm. math. d. savants dtrang. hautdrfisen der insecten. (Sitzungsber. Paris, 1755, t. 2, p. 276-28e. (Collection gesells, z. bef6rd, d. gesammt, naturw, zu complete des oeuvres de C: Bonnet, 1779, t. Marburg, June I867, no. 8, p. 65-72.) , p. 17-4.) 4aClaus, C. Ueber die seitendrtisen der e:Mfiller, O.F. Pile-larven med dobbelt larve von chrysomela ]5op,li. (Zeitschr. t: hale, og dens phalaene Kjdbenhavn, wiss. zool., 186, bd. II, p. 3o9-314, pl. 25. 1772. P. 53-56, pl. 2, fig. 3-5" 44Lyonet, P. Recherches sur l'anatomie et eSAmoreux, P.J. Notice des insectes de la les metamorphoses de diff6rentes espces France, rfiputs venimeux,... Paris, 1789 d'insectes. Ouvrage posthume, public par p. 282-285. M. W. deHaan, Paris, 1832. J6rdens, J. H. Entomologie undhelmin- 4IvGeer, K: Abhandlungen zur ge- tholoie des menschlichen k6vpers Hof, schichte der insekten aus dem franz6sischen 8Ol, bd. I, p. lOI-lO2, pl. 12, fig, 6-8. iibersetzt Nfirnberg, I78I, bd. 4, P. 386- Rengger, J. R. Physiologische unter- 387, pl. 8, fig. 18-22. [-The first edition in suchungen fiber die thieriscbe haushaltung French was in 1755.1] der insecten. Tfibingen, I8I 7, p. 3,5-36. 4Ratzeburg, J. T. C. Die forst-insekten Lintner, J. A. Entomological contribu- Theil I, Die ktf'er... 1837 p. 246. tions, 878, no. 4, P. 85 (I97). Westwood, J O. An introduction to the 3l'rench, G: H. Larvae of cerura occf- modern classification of insects. In 2 vol. dezlalis Lint., and c. borealis, Bd. (Can. Lond. I839, v. 1, p. 388-389 entom., July I88I, v. I3, p. I44-I45.) 4SChapuis et CandSze. Catalogue des Plateau, F. Note sur une scrtion larves des col4opStres (M6m. soc. sci. de propre aux coloptves dytiscides. (Ann. Li6ge, 853, t. 8, p. 351-653, pl. 1-9. p. 6- soc. entom., Belg., 1876, v. 19, p. i-lO.) 612. 4Rougement, P. Observations sur l'or- 4OKlemensiewicz, S. Zur niiheren kennt- gane d4tonant du brachius crepzTas Oliv. niss dechautdrtisen bei den raupen und bei (Bull. soc. sci. nat. Neuchtel, I879, v. malachius. (Verh. k.-k. zool.-bot, gesells. P. 47-478, pl.) Wien., 1882 [-I883], bd. 32; Abh., p. 459-474, Witlaczil, E. Zur anatomic der aphiden. pl. 21-22.) 401

Aldrovandus, U. De animalibus insectis *Schwarz, C. Neuer raupenkalender, libri septem cum singulorum iconibus ad Ntirnberg, 1791. Abth. I, p. 59" vivum expressis Denub impress Bonon. Fitch, A. 8th rept. on the noxious and apud Clementem Ferronium, i638 p. 273. other insects of... New York. (Trans. N. Y. [The first edition was in 16o2.] state agric, soc., I862, v. 22, p. 657-684.) 5Goedart, J. Metamorphosis naturalis sire p. 677 [Separate, p. I95 ]. insectorum historia... Amstelodami, 17oo. 6SColeman, N. Notes on orgyia leucostig- Pars 2, p. 136. [French ed. of I7OO, t. 2, p. ma. (Papilio, Nov.-Dec. 1882 [Jan. I883], 162; Lister's Latin ed., Lond., 1685, p. 60.] v. 2, p. 164-166. p. 165 '2Rdaumur, R. A. F. Mdmoires pour servir 69Coquillett, D. W On the early stages of k l'histoire des insectes Paris, 1736, t. 2, some moths. (Can. entom., March I88O, v. p. 266-269, pl. 21-22. t. 2, partie 2, p. 21-23 x2, p. 43-46.) 45. of the Amsterdatn ed. of P. I737-z748. 7Dufour, L. Recherches anatomiques sur aMeckel, H. Mikrographie einiger drii- les carabiques et sur plusieurs autres coleop- senapparate der niederen thiere. (Arch. tdres. (Ann. d. sci. nat., 1826, t. 8, p. 5-54.) anat. phys. "u. wiss. reed., I846 p. 1-73 pl. F. 1-3.) p. 46. Miiller, The habits of various insects. (Nature, ti June 1874, v. IO, p. lO2-1o3, 4I Schiiffer, J. C. Neuentdeckte theile an cn. und raupen zweyfaltern Regensburg, :'2Miiller, F. Die stinkk61bchen derweibli- 1754. then maracuj',tfalter. (Zeitschr. f. wiss. Sulzer, J. H. Die kennzeichen der insek- zool., 1877, bd. 3o, p. 67-17o, pl. 9.) ten... Zfirich, fig. 176 p. 65-67 pl. 5, 34. :aMfiller, F. Beobachtungen an brasilia- Kirby and Spence. Introduction to ento- nischen schmetterlingen. . Die duftschup- mology... 2d ed., Lond., 818, v. 2, p. 238- pen der miinnlichen maracujMalter. (Kos- 239. mos, Aug. 1877, bd. t, p. 391-39S fig. S-6, 174 Laboulbgne, A. Note sur les caroncules cm.) thoraciques ou cocardes rouges du raalachs :Miiller, F. Der anhang am hinterleibe @ustlas. (Ann. soc. entom. Fr., 1858, der acraea-weibchen. (Zool. anzeiger, 6 s. 3, t. 6, p. 521-528. Aug. 1883, jahrg. 6, p. 4IS-416.) SLiegel, H. Ueber den aussttilpungs- :Doubleday, E: Genera of diurnal lepi- apparat von malac/ius und verwandten fbr- doptera, Lond., 1846-185o, p. 139. men. Inaugural-dissertation... Hannover date_. :Rogenhofer, A. Drei schmetterlings- metamorphosen. (Verh. k.-k. zool.-bot, ge- 5Petzhold, C. P. Lepidopterologische sells. Wien, I862, bd. 12; Abh., p. I24-i3o. beytrRge. (L. G. Scriba's Beitrge zu er p. 1227. insekten-geschichte, Frankfurt am Main, heft nGoossens, T. [Sur un organe entre la 1793, 3, P. 23o-251.) tte et la premiere paire de pattes de quelques Guende, A. D'un organe particulier que chenilles. (Ann. soc. entom. Ft., 1869, s. prdsente une chenille de lycaena. (Ann. soc. 4, t. 9; Bull., p. 6o-61.) entom. s. v. Fr., 1867, 4, 7, P. 665-668, pl. 13, T. Introduction fig. 9-12.) SLacordaire, J. l'ento- mologie. 838, t. 2, p. 45. mEdwards, W H Notes on lycaena 3seud- T. De la corne chez les argiolus and its larval history. en- 7OGoossens, (Can. (Ann. soc. entom. Fr., s. t. tom., 1878, v. IO, p. 1-14, fig.) idae. t873, 5, Jan. 3, P. 123-128") nEdwards, W: H: On the larvae of lyc. Mgmoire and attendant ants. SBonnet, C: sur une nouvelle 2seudargiolus (Can. pattie commune k plusieurs espces de che- entom., July v. lO, p. fig. 1878 131-136 8.) nilles. (M6m. math. d. savants trang. naScudder, S H. Butterflies their struc- Paris, 1755, t. 2, p. 44-52.) (Collection com- ture, changes, and life-histories... N. Y., plete des ceuvres de C Bonnet, t. 2, 1779, p. 1881 p. 26-27. 3-16.) n4Gerstaecker, C. E. A. Ueber das vor- mMtiller, F. Ueber haarpinsel, filzflecke kommen von aus.ttilpbaren hautanhiingen und iihnliche gebilde auf den flfigeln miinnli- am hinterleibe an schaben. (Archiv f. natur- chef schmetterlinge. (Jena. zeitschr, f. gesch., 1861, bd. 2I, th. I, p. IO7-i5. naturw., 1877 bd. II, p. 99-114.) nSBrunner yon Wattenwyl, K: [Ueber ein SBertkau, P. Duftapparat an schmetter- neues organ bei den acridiodeen.] (Ver- lingsbeinen. (Entom. nachrichten, Sept. handl. k.-k. zool.-bot, gesells. Wien, 1879, bd. 1879 jahrg..5, P. 223-224.) 9; Sitzungsber., p. 26-27. International Journal of Peptides

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