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Fig. 57.—Maxilla of Nemognatha, ♀, from Montana. A, base of maxilla enlarged to show the taste-papillæ (tp) and cups (tc), on the galea (ga). B, part of end of galea to show the imperfect segments and taste-organs: n, nerve; a ganglionated nerve supplies each taste-papilla or cup; l, lacinia; p, palpifer; s, subgalea.

Fig. 58.—Maxilla of Panorpa.

Fig. 59.—Maxilla of Limnephilus pudicus: mx, stipes; lac, galea.

The palpus is in general antenniform and is composed of from 1 to 6 joints, being usually 4– or 5–jointed, and is much longer than the galea. In the maxilla of the beetle Nemognatha (Fig. 57), the galea is greatly elongated, the two together forming an imperfect tube or proboscis and reminding one of the tongue of a moth, while the lacinia is reduced. In the Mecoptera the lacinia and galea are closely similar (Fig. 58); in the Trichoptera only one of the lobes is present (Fig. 59), while in the Lepidoptera the galea unites with its mate to form the so-called tongue (Fig. 60). The maxilla of the male of Tegeticula yuccasella is normal, though the galeæ are separate; but in the female, what Smith regards as the palpifer (the “tentacle” of Riley) is remarkably developed, being nearly as long as the galea (Fig. 61) and armed with stout setæ, the pair of processes being adapted for holding a large mass of pollen under the head.

Fig. 60.—Tongue of Aletia xylina, with the end magnified.—Pergande del., from Riley. A, much reduced maxilla (mx) of Paleacrita vernata; mx.p, palpus.

Fig. 61.—A, maxilla of Tegeticula yuccasella, ♂: g, galea. B, ♀: pl, enormously developed palpifer; mx.p, palpus; c, cardo; st, stipes; sty, stylus.

In coleopterous larvæ the maxillæ are 2–lobed (Fig. 62), the galea being undifferentiated, but in those of saw-flies the galea is present (Fig. 63, gal).

Fig. 62.—Larva of Rhagium lineatum: lat, lateral view of head and thoracic segments; mx, first maxilla; ml, undifferentiated lacinia and galea; v, under side of head and pro- and meso- thoracic segments; v.m.s., one of the middle ventral segments, magnified six times; mx′, 2d maxilla.

It now seems most probable that in the first maxillæ we have the primary form of buccal appendage of insects, the appendage being composed of three basal pieces with three variously modified distal lobes or divisions; and that the mandibles and second maxillæ are modifications of this type.

How wonderfully the maxillæ of the Lepidoptera are modified, and the peculiar shapes assumed in the Diptera, Hymenoptera, and other groups, will be stated in the accounts of those orders, but it is well to recall the fact that in the most primitive and generalized moth, Eriocephala, the lacinia is well developed (Fig. 64).

As Newport remarks, the office of the maxillæ in the mandibulate insects is of a twofold kind; since they are adapted not only for seizing and retaining the food in the mouth, but also as accessory jaws, since they aid the mandibles in comminuting it before it is passed on to the pharynx and swallowed. Hence, as the food varies so much in nature and situation, it will be readily seen that the maxillæ, especially their distal parts, vary correspondingly. Thus far no close observations on the exact use of the first and second maxillæ have been published.

The palpi also are not only organs of touch, but in some cases act as hands and also bear minute sense-organs, the function of which is unknown, but would appear to be usually that of smell.

Fig. 63.—Selandria larva, common on Carya porcina, with details of mouth-parts: leg, leg; mx, maxilla; gal, galea; lac, lacinia.

The second maxillæ.—The “under-lip” or labium of insects is formed by the fusion at the basal portion of what in the embryo are separate appendages, and which arise in the same manner as the first maxillæ. They are invariably solidly united, no cases of partial or incomplete fusion being known. The so-called labium is situated in front of the gula or gular region, and is bounded on each side by the gena, or cheek. As already observed, the second maxillæ appear to be the appendages of the last or occipital segment of the head.

Fig. 64.—Maxilla of Eriocephala calthella: l, lacinia; g, galea; mx.p, maxillary palpus; st, stipes; c, cardo.—After Walter.

The second maxillæ are very much differentiated and vary greatly in the different orders, being especially modified in the haustellate or suctorial orders, notably the Hymenoptera and Diptera. In the mandibulate orders, particularly the Orthoptera, where they are most generalized and primitive in shape and structure, they consist of the following parts: the gula (a postgula is present in Dermaptera), submentum (lora of Cheshire, i, p. 91), mentum, palpifer, the latter bearing the palpi; the lingua (ligula) and paraglossæ, while the hypopharynx or lingua is situated on the upper side. The labial palpi are of the same general shape as those of the first maxillæ, but shorter, with very rarely more than three joints, though in Pteronarcys there are four. Leon has detected vestigial labial palpi in several Hemiptera (Fig. 73). As to the exact nature and limits of the gula, we are not certain; it is not always present, and may be only a differentiation of the submentum, or the latter piece may be regarded as a part of the gula.

We are disposed to consider the second maxillæ as morphologically nearly the exact equivalents of the first pair of maxillæ, and if we adopt this view it will greatly simplify our conception of the real nature of this complicated organ. The object of the fusion of the basal portion appears to be to form an under-lip, in order both to prevent the food from falling backwards out of the mouth, and, with the aid of the first pair of maxillæ, to pass it forward to be crushed between the mandibles, the two sets of appendages acting somewhat as the tongue of vertebrates to carry and arrange or press the morsels of food between the teeth or cutting edges of the mandibles.

The spines often present on the free inner edges of the first and second maxillæ (Figs. 54, 62) form rude combs which seem to clean the antennæ, etc., often aiding the tibial combs in this operation.

The submentum and mentum, or the mentum when no submentum is differentiated (with the gula, when present), appear to be collectively homologous with the cardines of the first pair of maxillæ, together with the palpifers and the stipites.^[15] These pieces are more or less square, and have a slightly marked median suture in Termitidæ, the sign of primitive fusion or coalescence.

The most primitive form of the second maxillæ occurs in the Orthoptera and in the Termitidæ. The palpifer is either single (Periplaneta, Diapheromera, Gryllidæ) or double (Blatta orientalis, Locustidæ). In Prisopus the single piece in front of the palpifer is in other forms divided, each half (Blatta, Locustidæ, Acrydidæ) bearing the two “paraglossæ,” which appendages in reality are the homologues of the lacinia and galea of the first maxillæ.^[16] In the Termitidæ (Fig. 65) the lingua is not differentiated from the palpifer, and the two paraglossæ (or the lamina externa and interna of some authors) with the palpus are easily seen to be the homologues of the three lobes of the first maxillæ. In the Perlidæ (Pteronarcys, Fig. 66) the palpifer is divided, while the four paraglossæ arise, as in Prisopus and Anisomorpha, from an undivided piece, the lingua not being visible from without. In the Neuroptera the lingua or ligula is a large, broad, single lobe, without “paraglossæ,” and the palpifer is either single (Myrmeleon, Fig. 67), or divided (Mantispa, Fig. 68). In Corydalus (Fig. 29) the palpifer forms a single piece, and the lingua is undivided, though lobed on the free edge.

Fig. 65.—Second maxillæ of Termopsis angusticollis: li, the homologue of the lacinia; le, galea.

In the metabolic orders above the Neuroptera the lingua is variously modified, or specialized, with no vestiges of the lacinia or galea, except in that very primitive moth, Eriocephala, in which Walter found a minute free galea, me, and an inner lobe (Figs. 76, 77), the lacinia.

Fig. 66.—Second maxillæ of Pteronarcys californica.

Fig. 67.—Second maxillæ of Myrmeleon diversum.

Fig. 68.—Second maxillæ of Mantispa brunnea.

The hypopharynx.—While in its most generalized condition, as in Synaptera, Dermaptera, Orthoptera, and Neuroptera, this anterior median fold or outgrowth of the labium forming the floor of the mouth may retain the designation of “tongue,” lingua, or ligula; in its more specialized form, particularly when used as a piercing or lapping organ, the use of the name hypopharynx seems most desirable. And this is especially the case since, like the epipharynx, it is morphologically a median structure, and while the epipharynx forms the soft, sensitive roof of the mouth, or pharynx; its opposite, the hypopharynx, rises as a fold from the floor of the mouth, forming in its most generalized condition a specialized fold of the buccal integument. In certain cases, as in the honey-bee, the very long slender “tongue” or hypopharynx is evidently, as in the case of the epipharynx, a highly sensitive armature of the mouth.

In all insects this organ—whether forming a soft, tongue-like, anterior portion or fold of the labium, and “continuous with the lower wall of the pharynx,” or a hard, piercing, awl-like appendage (fleas and flies), or a long, slender, hairy or setose, trough-like structure like the “tongue” of the honey-bee—has a definite location at the end and on the upper side of the labium, and serves to receive at its base the external opening of the salivary duct.

The hypopharynx, as well shown in its lingua condition in Orthoptera, is continuous with and forms the anterior part or fold of the base of the coalesced second maxillæ. It does not seem to be paired, or to represent a pair of appendages.

Opinion regarding the homology of this unpaired piercing organ is by no means settled, and while there is a general agreement as to the nature of the paired mouth-parts, recent observers differ very much as to the morphology of the organ in question.

It is the langue or lingua of Savigny (1816), the ligula of Kirby and Spence (1828), the langue ou languette (lancette médiane du suçoir) of Dugès (1832), the lingua of Westwood (Class, ins., ii, p. 489, 1840), “the unpaired median piercing organ” (“the analogon of the epipharynx of Diptera”) of Karsten (1864), the “tongue” of Taschenberg (1880).

The name hypopharynx was first proposed by Savigny in 1816, who, after naming the membranous plate which has for its base the upper side of the pharynx, the epipharynx, remarks: “Dans quelques genres, notamment dans les Eucères, le bord inférieur de ce même pharynx donne naissance à un autre appendice plus solide que le précédent, et qui s’emboîte avec lui. Je donnerai à ce dernier le nom de langue ou d’hypopharynx. Voilà donc la bouche des Hyménoptères composée de quatre organes impaires, sans y comprendre la ganache ou le menton; savoir, la lèvre supérieure, l’épipharynx, l’hypopharynx, et la lèvre inférieure, et de deux organes paires, les mandibules et les mâchoires.”

As stated by Dimmock: “The hypopharynx is usually present in Diptera (according to Menzbier absent in Sargus), and contains a tube, opening by a channel on its upper surface; this channel extends back, more or less, from the tip, and is the outlet for the salivary secretion. The tip of the hypopharynx may be naked and used as a lance (Hæmatopota, according to Menzbier), or may be hairy (Musca). The upper side of the base of the hypopharynx is continuous with the lower wall of the pharynx; its under surface may entirely coalesce with the labium (Culex, male), may join the labium more or less, anterior to the month (Musca), or, if either mandibles or maxillæ are present, its base may join them (Culex, female).” (p. 43.)

Fig. 69.—Section of head of Machilis maritima: hyp, hypopharynx; lbr, labrum; t, tentorium; ph, room in which the mandibles move on each other; p, paraglossa; mx, labium; sd, salivary duct; s.gl, salivary gland. oe, œsophagus.—After Oudemans.

We will now briefly describe the lingua, first of the mandibulate or biting insects, and then its specialized form, the hypopharynx of the haustellate and lapping insects.

The lingua (hypopharynx) exists in perhaps its most generalized condition in the Thysanura (Fig. 69), where it forms a soft projection, having the same relations as in Anabrus and other Orthoptera.^[17]

In the cockroach (Fig. 70), as stated by Miall and Denny, the lingua is a chitinous fold of the oral integument situated in front of the labium, and lying in the cavity of the mouth. The common duct of the salivary glands enters the lingua, and opens on its hinder surface. The lingua is supported by a chitinous skeleton (Figs. 70, B; 82, shp). “The thin chitinous surface of the lingua is hairy, like other parts of the mouth, and stiffened by special chitinous rods or bands.” (Miall and Denny.)

Fig. 70.—Hypopharynx of Periplaneta orientalis; the arrow points out of the opening of the salivary duct: A, origin of salivary duct. B, side view. C, front view.—After Miall and Denny.

In the Acrydiidæ (Melanoplus femur-rubrum) the tongue is a large, membranous, partly hollow expansion of the base of the labium. It may be exposed by depressing the end of the labium, when the opening of the salivary duct may be seen at the bottom or end of the space or gap between the hinder base of the tongue, and the inner anterior base of the labium, as shown by the arrows in Fig. 70. It is somewhat pyriform, slightly keeled above, and bearing fine stiff bristles, which, as they point more or less inwards, probably aid in retaining the food within the mouth. The base of the tongue is narrow, and extends back to near the pharynx, there being on the floor of the mouth, behind the tongue, two oblique, slight ridges, covered with stiff, golden-yellow hairs, like those on the tongue. The opening of the salivary duct is situated on the under or hinder side of the hypopharynx, between it and the base of the labium, the base of the former being cleft; the hollow thus formed is situated over the opening, and forms the salivary receptacle.

Fig. 71.—Section through the anterior part of the head of Anabrus (the mandibles removed), showing the relations of the hypopharynx (hyp) to the opening of the salivary duct (sd): g, galea; l, lacinia; mt, mentum; oe, œsophagus; lbr, labrum; cl, clypeus.

In the Locustidæ (Anabrus, Fig. 71) the tongue (hypopharynx) is a broad, somewhat flattened lobe arising from the upper part of the base of the mentum and behind the palpifer. This lobe is cavernous underneath, the hollow being the salivary receptacle (sr); the latter is situated over the opening of the salivary duct, which is placed between the base of both the hypopharynx and the labium. The salivary fluid apparently has to pass up and around on each side of the hypopharynx in order to mix with the food.

These relations in the Orthoptera are also the same in the Perlidæ, where the hypopharynx is well developed, forming an unusually large tongue-like mass, nearly filling the buccal cavity.

Fig. 72.—Lingua of a May-fly, Heptagenia longicauda, ×16: m, central; l, lateral pieces.—After Vayssière from Sharp.

In the Odonata the lingua is a small, rounded lobe, as also in the Ephemeridæ; in the nymph, however, of Heptagenia (Fig. 72) it is highly developed, according to Vayssière, who seems inclined to regard it as representing a pair of appendages. The tongue in Hemiptera is said by Léon to be present in Benacus griseus (Say) and to correspond to the subgalea of Brullé or hypodactyle of Audouin (Fig. 73), but this appears to correspond to the labium proper, rather than a true lingua, the latter not being differentiated in this order. In the Coleoptera the lingua is rather small. In beetles, as Anopthalmus (Fig. 74), it forms a setose lobe; and a well-developed nerve, the lingual nerve, passes to it, dividing at the end into several branches (n-l). In Sialis the lingua is short, much less developed than usual, being rounded, and bears on the edge what appear to be numerous taste-hairs, like those on the ends of the maxillary and labial palpi.

Fig. 73.—A, labium of Zaitha anura. B, of Z. margineguttata. C, of Gerris najas: mt, mentum; lp, labial palpi; sg, subgalea; l, lacinia (= intermaxillare and præmaxillare of Brullé); g, galea.—After Léon.

In the adult Panorpidæ the lingua is a minute, simple lobe.

Fig. 74.—Section through head of a carabid, Anopthalmus telkampfii: br, brain; f. g, frontal ganglion; soe, subœsophageal ganglion; co, commissure; n. l, nerve sending branches to the lingua (l); mn, maxillary nerve; mx, 1st maxilla; mm, maxillary muscle; mx′, 2d maxilla; mt, muscle of mentum; le, elevator muscle of the œsophagus; l of the clypeus, and a third beyond raising the labrum (lbr); eph, epipharynx; g, g, salivary glands above; g², lingual gland below the œsophagus (oe); m, mouth; pv, proventriculus; md, mandible. A, section passing through lingual gland (g²).

In the larval Trichoptera the spinneret is well developed, and in structure substantially like that of caterpillars, and it is plainly the homologue of the hypopharynx, receiving as it does the end of the silk-duct.

In the adult Trichoptera the hypopharynx is a very large, tongue-like, fleshy outgrowth, and is, both in situation and structure, since it contains the opening of the silk-duct, exactly homologous with the hypopharynx of insects of other orders, being somewhat intermediate between the fleshy tongue or lingua of the mandibulate insects, especially the Neuroptera, and the hypopharynx of the bees (Fig. 86). Lucas describes and figures it under the name of “haustellum,” but does not homologize it with the hypopharynx. The caddis-flies have been observed to drink water and take in both fluid and fine particles of solid food, and to use the haustellum for this purpose, the end being provided with minute sense-organs like those on the first maxillary lacinia, and possibly of a gustatory nature.

Fig. 75.—Head of Anabolia furcata: A, front view, showing the labrum removed. B, side view; ant, antenna; oc, ocellus; ol, labrum; gh, articulatory process; cmx₁, cardo; stmx₁, stipes; lemx₁, outer lobe (galea); ptmx₁, palpus of 1st maxilla; pl, palpus of 2d maxilla; ha, haustellum; so, gustatory pits; spr, opening of salivary duct; chsp, chitinous hook of the clasp; spr, furrow or gutter of the haustellum.—After Lucas.

Fig. 76.—Hypopharynx of Eriocephala calthella: lig, ligula, its membranous hinder edge; lig′, anterior horny edge of the ligula-tube opening outwards; hp, contour of the hypopharynx; mi, mala interior (lacinia); me, mala exterior (galea), of second maxilla; mx′ p, labial palpus.—After Walter.

The spinneret of the larvæ of Lepidoptera is evidently the homologue of the hypopharynx of insects of other orders. It will be seen that the homology of the different parts is identical, the common duct of the silk-glands opening at the end of the hypopharynx, which here forms a complete tube or proboscis extending beyond the end of the labium, in adaptation to its use as a spinning organ.

Walter refers to Burgess’s discovery of a hypopharynx in Danais archippus, remarking that this organ in the adult Eriocephalidæ (Fig. 76) exhibits a great similarity to the relations observable in the lower insects, adding:—

Fig. 77.— Labium of Micropteryx anderschella seen from within (the labial palpi (mx.′ p) removed to their basal joint). Lettering as in Fig. 76.—After Walter.

“The furrow is here within coalesced with the inner side of the labium, and though I see in the entire structure of the head the inner edge of the ligula tube extended under the epipharynx as far as the mandible, I must also accept the fact that here also the hypopharynx extends to the mouth-opening as in all other sucking insects with a well-developed under-lip, viz. the Diptera and Hymenoptera.”

He has also discovered in Micropteryx a paired structure which he regards as the hypopharynx (Fig. 77). As he states:

Fig. 78.—Hypopharynx (hph) of Danais: cl, clypeus; sd, salivary duct; m, labial palp muscles; fm, frontal muscle; ph, pharynx; cor, cornea.—After Burgess.

“A portion of the inner surface of the tube-like ligula is covered by a furrow-like band which, close to the inner side, is coalesced with it, and in position, shape, as well as its appendages or teeth on the edge, may be regarded as nothing else than the hypopharynx.”

A hypopharynx is also present in the highest Lepidoptera, Burgess having detected it in Danais archippus. He states that the hypopharynx forms the floor of the pharyngeal cavity; “it is convex on each side of a median furrow (Fig. 78, hph) and somewhat resembles in shape the human breast. The convex areas are dotted over with little papillæ, which possibly may be taste-organs.”

As a piercing organ the hypopharynx reaches its greatest development in the Siphonaptera and Diptera, where the chitinous parts are greatly hypertrophied, the fleshy tongue-like portion so developed in the mandibulate orders being greatly reduced. The chitinous parts are alike on each side of the median organ, being bilaterally symmetrical.

Fig. 79.—A, hypopharynx of Pulex canis: x, basal portion situated within the head; s. d, common duct of the four bladder-shaped salivary glands; s. d′, opening of the tubular salivary glands into the throat. B, end of the hypopharynx, showing the gutter-like structure and teeth at the end.—After Landois.

Fig. 80.—Beak of Vermipsylla: hyp, hypopharynx.—After Wagner.

In the fleas the hypopharynx is a large, slender, unpaired, long, chitinous trough, as long as the mandibles, and toothed at the end. Figures 79 and 80 show its relations to the other parts of the mouth; in Fig. 79, x, is seen where the salivary duct opens into the pharynx. Although this organ is not unanimously referred to the hypopharynx, yet from the description of Landois and others, it is evident that this structure does not correspond to the labrum or epipharynx, but belongs to or arises from the floor of the mouth, and, being in close relation to the labium, and also receiving the salivary duct, must be a true hypopharynx.

In the Diptera the hypopharynx reaches its highest development as a large, stout, awl-like structure.

Meinert, in his detailed and elaborately illustrated work, Trophi Dipterorum (1881), has made an advance on our knowledge of the hypopharynx and its homologies, both by his evidently faithful descriptions and dissections, and by his admirably clear figures.

Fig. 81.—Culex pipiens, section of head: oe œsophagus; sm, upper muscle, lm, lower muscle of the œsophagus; ph, pharynx; rm, retractor muscle of the receptacle (r) of the salivary duct (s.d); lbr, labrum; ep, left style of the epipharynx; f, part of front of head.—After Meinert.

Fig. 82.—Pharynx and hypopharynx of Simulium fuscipes: lph, lower lamina of the pharynx; p, the salivary duct (s.d) perforating the pharynx; o, orifice of the duct; shp, styles of the hypopharynx; mph, membranous edge of the hypopharynx; m, protractor muscle of the pharynx; gp, gustatory papillæ.—After Meinert.

“The hypopharynx, a continuation of the lower edge (lamina) of the pharynx, most generally free, more or less produced, acute anteriorly, forms with the labrum the tube of the pump (antliæ). (The hypopharynx when obsolete, or coalesced with the canal of the proboscis, is the theca; in such a case the siphon or tube is formed by the theca and labrum.) Meanwhile the hypopharynx, the largest of all the trophi (omnium trophorum maximus), constitutes the chief piercing organ (telum) of Diptera. The hypopharynx is moved by protractor, most generally quite or very powerful, and by retractor muscles.

“The efferent duct of the thoracic salivary glands (ductus salivalis) perforates the hypopharynx, more or less near the base, that the saliva may be ejected through the canal into the wound, or that it may be conducted along the labella. Very rarely the salivary duct, perforating the hypopharynx, is continued in the shape of a free, very slender tube.

“The salivary duct behind the base of the hypopharynx forms the receptacle or receptaculum, provided with retractor and levator muscles.”

Fig. 83.—Labrum-epipharynx (lbr and eph) and hypopharynx (hyp) of Tabanus brominus: oe, posterior cylindrical portion of the œsophagus; a, anterior swollen portion of the same; ph, pharynx; ph.m, pharyngeal muscle; p.ph, protractor muscle of the pharynx; r.oe, retractor muscle of the œsophagus; r.ph, retractor muscle of the pharynx; f.oe, flexor muscle of the pharynx; t.oe, twisting muscle of the œsophagus; s.r, receptacle of the salivary duct; l, its elevator muscle; s, its retractor muscle; cl, clypeus.—After Meinert.

Fig. 84.—Œsophagus (oe), pharynx (ph) with epipharynx and labrum (lbr) of Asilus atricapillus: m, ph, pharyngeal muscle; sr, salivary receptacle; t, twisting; r, l′r, retractor muscles; other lettering as in Fig. 83.—After Meinert.

It has been carefully studied by Meinert in a species of Culex (Fig. 81), Simulium (Fig. 82), Tabanus (Fig. 83), and in Asilus (Fig. 84), where it is seen to attain enormous proportions. In the Hymenoptera, this organ in its most specialized condition is a trough-like rod, adapted for lapping nectar (Fig. 85, 86, hyp). The tongue or hypopharynx of the honey-bee has been elaborately described by Cheshire in his Bees and Bee Keeping.^[18] He calls it the tongue or ligula. It is situated in a tube formed by the maxillæ and labial palpi, and can be partially retracted into the mentum. He states that it can move up and down in the tube thus formed, and then describes it as covered by a hairy sheath, its great elasticity being due to a rod running through its centre enabling it to be used as a lapping tongue. The sheath

“passes round the tongue to the back, where its edges do not meet, but are continuous with a very thin plaited membrane (G, pm) covered with minute hairs. This membrane, after passing towards the sides of the tongue, returns to the angle of the nucleus, or rod, over the under surface of which it is probably continued. The rod passes through the tongue from end to end, gradually tapering towards its extremity, and is best studied in the queen, where I trace many nerve threads and cells. It is undoubtedly endowed with voluntary movement, and must be partly muscular, although I have failed completely in getting any evidence of striation. The rod on the underside has a gutter, or trough-like hollow (cd, the central duct) which is formed into a pseudotube (false tube) by intercrossing of black hairs. It will also be seen that, by the posterior meeting of the sheath, the space between the folded membrane (G, sd) becomes two pseudotubes of larger size, which I shall call the side ducts.

Fig. 85.—Head of honey bee, worker: a, antenna; g, epipharynx; m, mandible; mx, maxilla; mxp, maxillary palpus; pg, paraglossa; lp, labial palpus; l, hypopharynx; b, its spoon.—After Cheshire; from Bull. Div. Ent. U. S. Dept. Agr.

“These central and side ducts run down to that part of the tongue where the spoon, or bouton (K, Fig. 86) is placed. This is provided with very delicate split hairs (b, Fig. 86) capable of brushing up the most minute quantity of nectar, which by capillarity is at once transferred by the gathering hairs (which are here numerous, long, and thin) to two side groove-like forms at the back of the bouton, and which are really the opened-out extremity of the centre and side ducts, assuming, immediately above the bouton, the form seen in F, Fig. 86. The central duct, which is only from 1 600 inch to 1 1000 inch in diameter, because of its smaller size, and so greater capillary attraction, receives the nectar, if insufficient in quantity to fill the side ducts. But good honey-yielding plants would bring both centre and side ducts into requisition. The nectar is sucked up until it reaches the paraglossæ (pa, B, Fig. 86), which are plate-like in front, but membranous extensions, like small aprons, behind; and by these the nectar reaches the front of the tongue, to be swallowed as before described.”

Fig. 86.—Tongue or ligula of the honey bee: A, under side of the tongue; lp, labial palpi; r, r, rod; p, pouch; sh, sheath; gh, gathering hairs; b, bouton or spoon. B, under lip or labium, with appendages, partly dissected; l, lora or submentum; a, a, retractor linguæ longus; sd, salivary duct; rb and b, retractor linguæ biceps; mx, maxillæ; lp, labial palpi; pa, paraglossa; gr, feeding groove; sh, sheath of ligula. C, D, E, sections of ligula; hp, hyaline plate of maxilla; h, hairs acting as stops; mx, maxilla; lp, labial palpi; sd, side duct. F, cross-section of extremity of tongue near the “spoon”; th, tactile hairs; r, rod; n, nucleus; gh, gathering hairs. G, cross-section of tongue without gathering hairs, × 400 times; sh, sheath; b, blood space; t, trachea: ng, gustatory nerve; cd, central duct; sd, lateral duct; pm, plaited membrane. H, same as G, but magnified two hundred times, and with pm, plaited membrane, turned outwards; h, closing hairs; lp, labial palpi; b, blood; n, nucleus; r, rod; h, closing hairs. I, small portion of the sheath; lettering as before. K, extremity of the tongue, with spoon; b, branching hairs for gathering.—After Cheshire.

Cheshire then settles the question which has been in dispute since the time of Swammerdam, whether the bee’s tongue is solid or tubular. He agrees with Wolff that the duct is a trough and not a tube, and proves it by a satisfactory experiment. He remarks:

Fig. 87.—Longitudinal section through the head of the honey bee, ♀, just outside of right antenna: ant, antenna with three muscles attached to mes, mesocephalic pillar; cl, clypeus; lbr, labrum; 1, chyle-gland (system no. 1, of Siebold); o, opening of the same; oc, ocellus; br, brain; n, neck; th, thorax; oe, œsophagus; s.d², s.d³, common salivary ducts of systems 2 and 3; v, salivary valve; c, cardo; ph, pharynx; mx′, labium; mx.′p, labial palpi; mt, mentum; mx, maxilla; hyp, hypopharynx; s, bouton.—After Cheshire.

“Bees have the power, by driving blood into the tongue, of forcing the rod out from the sheath, and distending the wrinkled membrane so that in section it appears as at H, Fig. 86, the membrane assuming the form of a pouch, given in full length at A. It will be seen at once that this disposition of parts abolishes the side ducts, but brings the central duct to the external surface. The object of this curious capability on the part of the bee is, in my opinion, to permit of cleaning away any pollen grains, or other impediment that may collect in the side ducts. The membrane is greasy in nature, and substances or fluids can be removed from it as easily as water from polished metal. If, now, the sides of a needle, previously dipped into clove oil in which rosanilin (magenta) has been dissolved, so as to stain it strongly red, be touched on the centre of the rod, the oil immediately enters, and passes rapidly upwards and downwards, filling the trough.”

Does the hypopharynx represent a distinct segment?—The facts which suggest that the hypopharynx may possibly represent a highly modified pair of appendages, arising from a distinct intermaxillary segment, are these: Heymons plainly shows that, in the embryo of Lepisma, the hypopharynx originates as a transverse segment-like fold in front of the 2d maxillary segment, and larger than it, and though he does not mention it in his text, it appears like the rudiment of a distinct segment; the hypopharynx of Ephemeridæ; arises and remains separate in the nymph from the labium (see Heymons’ Fig. 29, and there are two lateral projections; see also Fig. 72, and Vayssiere’s view that it may represent a pair of appendages; Kolbe also regards it as representing a third pair of maxillæ, his endolabium, p. 213). Though what is called an unpaired organ, it is composed of, or supported by, two bilaterally symmetrical styles, both in Myriopods (Fig. 6, labiella, stil) and in insects (Fig. 77, etc.). On the other hand, in the embryo of pterygote insects, an intermaxillary segment has not been yet detected.