Fig. 6—A, Worker of the honey-bee (Apis mellifica), with pollen plates laden; B, basal portions of a middle-leg (trochanter with part of coxa and of femur) with plumose hairs and grains of pollen; C, one hair bearing pollen-grains.
The hind-legs of bees are very largely used in the industrial occupations of these indefatigable creatures; one of their chief functions in the female being to act as receptacles for carrying pollen to the nest: they exhibit, however, considerable diversity. The parts most modified are the tibia and the first joint of the hind-foot. Pollen is carried by other parts of the body in many bees, and even the hind-leg itself is used in different ways for the purpose: sometimes the outer face of the tibia is highly polished and its margins surrounded by hair, in which case pollen plates are said to exist (Fig. 6, A); sometimes the first joint of the tarsus is analogous to the tibia both in structure and function; in other cases the hind-legs are thick and densely covered with hair that retains the pollen between the separate hairs. In this case the pollen is carried home in a dry state, while, in the species with pollen plates, the pollen is made into a mass of a clay-like consistence.[[11]] The legs also assist in arranging the pollen on the other parts of the body. The males do not carry pollen, and though their hind-legs are also highly modified, yet the modifications do not agree with those of the female, and their functions are in all probability sexual. The parasitic bees also do not carry pollen, and exhibit another series of structures. The most interesting case in this series of modifications is that found in the genus Apis, where the hind-leg of male, female, and worker are all different (Fig. 25); the limb in the worker being highly modified for industrial purposes. This case has been frequently referred to, in consequence of the difficulty that exists in connection with its heredity, for the structure exists in neither of the parents. It is, in fact, a case of a very special adaptation appearing in the majority of the individuals of each generation, though nothing of the sort occurs in either parent.
The proboscis of the bee[[12]] is a very complex organ, and in its extremely developed forms exhibits a complication of details and a delicacy of structure that elicit the admiration of all who study it. In the lower bees, however, especially in Prosopis, it exists in a comparatively simple form (Fig. 9, B, C), that differs but little from what is seen in some Vespidae or Fossores. The upper lip and the mandibles do not take any part in the formation of the bee's proboscis, which is consequently entirely made up from the lower lip and the maxillae, the former of these two organs exhibiting the greatest modifications. The proboscis is situate on the lower part of the head, and in repose is not visible; a portion, and that by no means an inconsiderable one, of its modifications being for the purpose of its withdrawal and protection when not in use. For this object the under side of the head is provided with a very deep groove, in which the whole organ is, in bees with a short proboscis, withdrawn; in the Apidae with a long proboscis this groove also exists, and the basal part of the proboscis is buried in it during repose, while the other parts of the elongate organ are doubled on the basal part, so that they extend backwards under the body, and the front end or tip of the tongue is, when in repose, its most posterior part.
For the extrusion of the proboscis there exists a special apparatus that comes into play after the mandibles are unlocked and the labrum lifted. This extensive apparatus cannot be satisfactorily illustrated by a drawing, as the parts composing it are placed in different planes; but it may be described by saying that the cardo, or basal hinge of the maxilla, changes from an oblique to a vertical position, and thrusts the base of the proboscis out of the groove. The maxillae form the outer sheath of the proboscis, the lower lip its medial part (see Figs. 7 and 9); the base of the lower lip is attached to the submentum, which rises with the cardo so that labium and maxillae are lifted together; the co-operation of these two parts is effected by an angular piece called the lorum, in which the base of the submentum rests; the submentum is articulated with the mentum in such a manner that the two can either be placed in planes at a right angle to one another, or can be brought into one continuous plane, and by this change of plane the basal part of the tongue can also be thrust forwards.
Fig. 7.—Side view of basal portions of proboscis of Bombus. a, Epipharyngeal sclerites; b, arrow indicating the position of the entrance to pharynx, which is concealed by the epipharynx, c; d, hypopharyngeal sclerites; e, vacant space between the scales of the maxillae through which the nectar comes: f, lobe; f′, stipes; g, cardo of maxilla: h, encephalic pillar on which the cardo swings; i, angle of junction of lores and submentum lorum; k, mentum; l, base of labial palp; m, maxillary palp.
There is considerable variety in the lengths of these parts in different genera, and the lorum varies in shape in accordance with the length of the submentum. The lorum is a peculiar piece, and its mechanical adaptations are very remarkable; usually the base of the submentum rests in the angle formed by the junction of the two sides of the lorum, but in Xylocopa, where the submentum is unusually short, this part reposes in a groove on the back of the lorum, this latter having a very broad truncated apex instead of an angular one; in the condition of repose the apex of the lorum rests in a notch on the middle of the back of the oral groove, and in some of the forms with elongate submentum, this depression is transformed into a deep hole, or even a sort of tunnel, so as to permit the complete stowing away of the base of the tongue, which would otherwise be prevented by the long submentum; another function of the lorum appears to be that, as it extends, its arms have an outward thrust, and so separate the maxillae from the labium. In addition to these parts there are also four elongate, slender sclerites that are only brought into view on dissection, and that no doubt assist in correlating the movements of the parts of the mouth and hypopharynx; one pair of these strap-like pieces extends backwards from the two sides of the base of the epipharynx; Huxley called them sclerites of the oesophagus; a better name would be epipharyngeal sclerites (Fig. 7, a): the other pair pass from the terminations of the epipharyngeal sclerites, along the front face of the hypopharynx, down to the mentum, their lower parts being concealed by the stipites of the maxillae; these are the hypopharyngeal sclerites, and we believe it will prove that they play a highly important part in deglutition. When the labrum of a bee is raised and the proboscis depressed, the epipharynx is seen hanging like a curtain from the roof of the head; this structure plays an important part in the act of deglutition. The entrance to the pharynx, or commencement of the alimentary canal, is placed below the base of the epipharynx. As we are not aware of any good delineations of the basal parts of the proboscis we give a figure thereof (Fig. 7). The maxillae in the higher bees are extremely modified so as to form a sheath, and their palpi are minute; in the lower bees the palpi have the structure usual in mandibulate Insects.
Returning to the consideration of the lower lip, we find that there is attached to the mentum a pair of elongate organs that extend forwards and form a tube or sheath, enclosed by the maxillary sheath we have previously mentioned; these are the greatly modified labial palpi, their distal parts still retaining the palpar form; and in the lower bees the labial palpi are, like the maxillary, of the form usual in mandibulate Insects. Between the labial palps and the central organ of the lip there is attached a pair of delicate organs, the paraglossae.
There remains for consideration the most remarkable part of the proboscis, the long, delicate, hairy organ which the bee thrusts out from the tip of the shining tube formed by the labial palps and the maxillae, described above, and which looks like a prolongation of the mentum. This organ is variously called ligula, lingua, or tongue.[[13]] We prefer the first of these names.
According to Breithaupt and Cheshire the structure of the ligula is highly remarkable; it is a tube (filled with fluid from the body cavity), and with a groove underneath caused by a large part of the circumference of the tube being invaginated; the invaginated part can be thrust out by increase of the pressure of the fluid in the tube. A portion of the wall of the invaginate part is thickened so as to form a chitinous rod.