CHAPTER VIII. A NEW ARRANGEMENT OF BRITISH BEES, WITH ITS RATIONALE, AND AN INTRODUCTION TO THE FAMILY, SUBFAMILIES, SECTIONS, AND SUBSECTIONS.

If perfection of instinct, and an organization exquisitely moulded to a complete adaptation to the many delicate and varied functions of that instinct, as well as to the exercise of every faculty incidental to the class, be certainly a proof of pre-eminence, we may justly claim this position for the Order Hymenoptera. There is no characteristic in which they are deficient, nor any in which some of the members of the Order do not transcend in aptitude the insects of all the others.

If they have not been placed at the head of the class Insecta, it has been because systematic convenience did not permit the transposition, on account of the interruption it would have caused to the convenient linking of the rest in a consecutive arrangement. Yet are they the most volatile fliers, the most agile runners, the most skilful burrowers, and consummate architects.

The beauty resulting from the combinations of symmetry of form, elegance of motion, brilliancy of colour, and vivacity of expression, is to be found exclusively amongst them. Either in the velocity of their flight, or in its playful evolutions and graceful undulations, they are unsurpassed, and they hover in the execution of their designs with pertinacious perseverance. No insect structure can more thoroughly exemplify the most appropriate adaptation to its uses, and the most admirable elegance in the formation of the means of execution.

I thus claim for them, and which I think I may without infraction of dispute, the distinctive rank amongst insects.

Having fixed the station of the Hymenoptera generally, we have next to seek the relative rank of the natural divisions into which they readily separate.

Taking structure and instinct conjunctively, there can be no doubt that the first position will be conceded to that division of the Order which comprises the aculeated tribes—those armed with stings,—some of whose members, in each of the three large divisions into which they fall, being social, that is, living in communities, organized by a peculiar polity or administration.

These aculeates divide into, first, the fossorial Hymenoptera, or burrowers; and the equivalent branch the Diploptera, or wasps, distinguished and named from their folding the superior wings longitudinally in repose; secondly, the heterogeneous Hymenoptera, or ants, named from the dissimilarity either in size or structure of their females, a peculiarity incidental to all the social Hymenoptera, but living in community is more peculiarly characteristic of this division, it being in the other divisions restricted to a few genera only, whereas here the solitary habit is the exceptional. In all cases of socialism there are three classes of individuals,—males, females, and abortive females. In the other social kinds of Hymenoptera, these abortive females, called neuters, perform the labours of the community, and they are always winged; whereas amongst the ants they are never winged, and they constitute civil and military departments, the former attending to domestic matters, and the latter making predatory excursions to enslave the inhabitants of other communities, to aid their civilians in their many duties.

The third and last division of the aculeate Hymenoptera contains the MellicolligerÆ, the bees, or honey-gatherers.

Thus each division of the aculeated Hymenoptera is closely linked to the others by the strong affinity of the social habits of some of the genera of their several families.

The food of these three divisions of the aculeated Hymenoptera differs considerably, the Fossores being raptorial flesh-feeders, which hunt down and destroy their prey, and supply it as food to their young; the HeterogynÆ are omnivorous,—grain, fruits, or carrion being equally welcome to them; but in these climates I am not aware that they destroy life, although their wide migrations within the tropics are undertaken in the very spirit of the Huns and Vandals, for they devastate everything they come across; but the whole family of bees are exclusively honey-feeders without any carnivorous propensities, and use their stings merely as weapons of defence.

Although all the social aculeates are edifiers, and although the wasp in its papier mÂchÉ domicile may vie with the honey-bee in capacity and skill in the structure of the hexagons of the habitation it erects or suspends, which are as perfect, and almost as delicate, although fabricated of a coarser material than those within the hive, and wherein also the several compartments form a more homogeneous unity, and the uniformity of the several layers or floors is more in accordance with architectural symmetry,—yet must the palm of precedence be accorded to the bee, from the more elaborate and perfect development of the social instinctive faculty.

We may be the more excused for this preference when we weigh the interest of the genus Apis to man. The wasp boots us nothing, but is the pilferer of our fruits, and a marauder upon the hive, whose inhabitants it destroys and consumes their produce, it being indifferent to them which they obtain—the bee or the honey,—either furnishing them with sustenance. The ant is obtrusive and incommodious, making incursions upon the pantry, the store-room, the green-house, and the hothouse; disfiguring our flower-beds, and often disgusting us with our aliment by the impertinent intrusion of its appearance. But the bee stores up for us honey, whose cruses are as inexhaustible as the oil cruse of the good widow of Zarephath, and whose waxen shards furnish us with a beautifully soft light, which in Catholic worship adds solemnity to the rites of religion. In doing this the bee fulfils a sovereign function in the economy of nature, by the fertilization of the flowering plants, with which she reciprocates benefits; the preponderance, however, is importantly in favour of the flower.

If captious objectors should dispute the position we thus claim for the bees, we will willingly leave them the wasp with its sting, whilst we sedulously cultivate the active and industrious bee, whose associations range through all the fields of poetry, but nowhere more lusciously than in the beautiful compositions of the Sanskrit poets Kalidasa and Yayadeva.

The position of the family, whose English constituents I shall subsequently treat of, being thus fixed, I have next to explain the several subdivisions into which it is divided in the following arrangement.

I am prompted to propose this new distribution of the British bees, by the manifest imperfection of the several arrangements of them already extant. The defects of these systems I shall have occasion to exhibit in reference to the course I have been induced to take.

Mr. Kirby’s keenness of observation led him to surmise, from the absence of polliniferous brushes upon the posterior legs, or other parts of the body of some, that there might be a class of bees analogous to the cuckoo, amongst the birds, who did not rear their own young, or undertake any of the cares of maternity; but that led by a peculiar instinct they deposited their eggs in the nests of more laborious kinds, for their young to be nurtured upon the provision laid up in store by the latter for the supply of their own progeny. This being merely a supposition, Mr. Kirby made no use of it in the distribution of his families.

Observation has since confirmed the conjecture, and the fact lends material aid to the combination of the bees into detached groups, and which has been partially applied since by all systematizers.

Conjunctively with the assistance derived from this circumstance, the various modes whereby pollen is collected and conveyed, either on the legs or on the belly, further facilitates the grouping of the family. Other structural or economical peculiarities lend their aid, and although the arrangement primarily emanates from the differences in the formation of the tongue, these are corroborated by differences in other organs, and the general distribution, as well as the special combinations, all result from natural characteristics.

The simplicity of the arrangement thus effected is very striking; and we thus find all the bees having similar habits, and with a similar structure united together by it in distinct groups.

I will here insert my scheme, and exhibit why and in what it differs from those of my predecessors; and, where necessary, I shall append such observations upon the several methods extant, as will sufficiently show the necessity, and vindicate the introduction of a new one.

Family MELLICOLLIGERÆ (Honey collectors).

Subfamily 1. AndrenidÆ (Subnormal Bees).

Section 1. With lacerate paraglossÆ.

Subsection a. With Emarginate Tongues.

Genus 1. Colletes.

2. Prosopis.

Subsection b. With Lanceolate Tongues.

Genus 3. Sphecodes.

4. Andrena.

5. Cilissa.

Section 2. With entire paraglossÆ.

Subsection c. With Acute Tongues.

Genus 6. Halictus.

7. Macropis.

8. Dasypoda.

Subfamily 2. ApidÆ (Normal Bees).

Section 1. Solitary.

Subsection 1. Scopulipedes (brush-legged).

a. FemoriferÆ (collectors on the entire leg).

† With two submarginal cells.

Genus 9. Panurgus.

b. CruriferÆ (collectors on the shank only).

† With two submarginal cells.

Genus 10. Eucera.

†† With three submarginal cells.

Genus 11. Anthophora.

12. Saropoda.

13. Ceratina.

Subsection 2. Nudipedes (naked-legged).

a. With three submarginal cells.

Genus 14. Nomada.

15. Melecta.

16. Epeolus.

b. With two submarginal cells.

Genus 17. Stelis

18. Coelioxys.

Subsection 3. Dasygasters (hairy-bellied).

All with two submarginal cells.

Genus 19. Megachile.

20. Anthidium.

21. Chelostoma.

22. Heriades.

23. Anthocopa.

24. Osmia.

Section 2. Cenobites (Dwellers in Community).

Subsection 1. Spurred.

† Parasitical.

Genus 25. Apathus.

†† Collectors.

Temporarily social.

Genus 26. Bombus.

Subsection 2. Unspurred.

Permanently social.

Genus 27. Apis.

The primary division of the bees into two large branches, viz. into the AndrenidÆ, or abnormal bees, and the ApidÆ, or normal bees, is effected by the mode in which they fold the cibarial apparatus in repose. In the description of the structure of the imago, I have enlarged upon these organs, and for their explanation I must refer to that chapter where diagrams exhibit the structure of the different kinds of trophi of the bees, as well as their mode of folding. Here it is only necessary to notice that in the AndrenidÆ, the joint at the base draws back the basal portion when protruded, and this basal portion is further jointed at the point of the insertion of the paraglossÆ and labial palpi, and parallel with which joint the maxillÆ are likewise jointed close to the sinus where the maxillary palpi are inserted laterally upon it. The basal portion thus throws the anterior part forward or retracts it, at the will of the insect, and in the latter case, being then in repose, it lies in contiguous parallelism to the basal half, but beneath it. When thus withdrawn, the short tongue itself, with its paraglossÆ and labial palpi are sheltered beneath the coping of the labrum and the lateral protection of the mandibles, whilst the horny sheathing of the maxillÆ protect the softer parts folding underneath.

In the ApidÆ, or normal bees, the basal joint has the same action in withdrawing the entire organ into its place of rest; but the joint which gives it this power is not in an analogous situation to that in the AndrenidÆ, for it is seated short of the joint which lies at the base of the several organs of the cibarial apparatus. By bending these downwards, it carries their apex backwards towards the basal fulcrum through the action of these two joints, and, when there, the more delicate ones are protected from abrasion or injury, by the lateral overlapping of the horny skin of the maxillÆ. All being thus withdrawn within this covering, upon the joint which folds them back, seated at the base of the tongue, the labrum falls, and further to strengthen this protection, the mandibles close over it like forceps.

That this difference in the arrangement of the cibarial apparatus points to any distinctive peculiarities of economy has not been ascertained, for the habits of the Scopulipedes greatly resemble those of the AndrenidÆ; although the habits of one of them, Anthophora furcata, are remarkably like those of the foreign genus Xylocopa, in its mode of drilling wood. But the ApidÆ have cross affinities amongst themselves, thus Ceratina resembles Heriades, and some of the OsmiÆ, in the way in which it nidificates.

The tongues of the AndrenidÆ are always shorter, broader, and flatter than those of the ApidÆ, in which they are always long, cylindrical, and tapering. In the first section of the AndrenidÆ, the paraglossÆ are obtusely terminated at the apex, thence called lacerated, and where they are fringed with brief bristles. The peculiar form of the tongue in this section suggests its being separated into two subsections, that organ being in the first subsection very broad and bilobated, which gives those insects their position in the series by approximating them to the preceding family of the Diploptera, or wasps, whose tongues have the same bilobate form, but each lobe in them is furnished with a gland. These tongues, in both cases of the wasps and these bees, may conduce to the building or plastering habits of the insects. The form may aid the wasp and the Colletes, the first in the moulding of its hexagonal papier-mÂchÉ cells, as it may the second in shaping and embroidering the silk-lined abode of its embryonic progeny. Why Prosopis should have this organization is difficult to conceive, unless it be from an analogy of structure incidentally previously referred to, beyond which any special object has hitherto escaped detection.

In the second section of the AndrenidÆ, which have the paraglossÆ entire and terminating in a point, the tongues all also terminate acutely with a lateral inclination inwards. In the lanceolate-tongued tribe they bulge outwards laterally, although pointed at the apex.

All this subfamily of AndrenidÆ, excepting only the two genera reputed parasites, viz. Prosopis and Sphecodes, are essentially Scopulipedes, densely brush-legged, for the conveyance of pollen which they vigorously collect; but from the brevity of their tongues they are restricted to flowers with shallow petals and apparent nectaria, their favourite plants being the abounding CompositÆ and UmbelliferÆ, as well as the RosaceÆ, whence they derive the agreeable odours which many of them emit upon being captured.

Their peculiar mode of collecting is a further reason for bringing the brush-legged ApidÆ collectively to the top of the normal bees, in juxtaposition to the AndrenidÆ, where the transition is made very naturally from Dasypoda to Panurgus.

The whole of the cibarial apparatus, or trophi, is always complete in all its constituent parts throughout the AndrenidÆ; and it is only with Ceratina, in the group of scopuliped ApidÆ, that it begins to show the tendency it has to abnormal deficiencies, by the paraglossÆ, in that genus, being obsolete. This characteristic, then, exhibits itself in the Nudipedes with two submarginal cells who are parasitical upon the Dasygasters, in whom also the maxillary palpi participate in a deficiency in the authentic number of their joints, whilst in Apis both maxillary palpi and paraglossÆ are unapparent. This shows that the numerical completion of the organs of the mouth have nothing to do with the qualifications of the creature, the best endowed in other respects being thus curtailed, the final cause of which is not yet understood.

The shape of the tongue itself thus separates the AndrenidÆ into three well-defined divisions readily perceptible. These, as I have just observed with respect to the differences in the mode of closing the oral apparatus in both cases, yield no clue to economy and habits, for which observation must supervene to illustrate it. This, patiently carried out, is very desirable, as it is still in discussion whether, notwithstanding the elucidation structure affords, Prosopis and Sphecodes are or are not parasitical. Structure says they are, for, like the cuckoo-bees forming the group Nudipedes in the ApidÆ, they are destitute of the requisite apparatus for collecting pollen. Mr. Kirby, however, gives direct testimony in favour of Sphecodes being a burrower, in the case of which bee it ought not to be a matter of much difficulty to determine, for on sandy plateaus I have occasionally found it very abundant, especially where there was ragwort (Senecio) in flower in the vicinity, to which the males resorted; but being at the time more intent on other matters, I neglected the opportunity. Other observers concur with Mr. Kirby as regards Sphecodes, and also say as much for Prosopis (better known as HylÆus). I strongly incline to the opinion enunciated by Latreille and Le Pelletier de St. Fargeau, that they are parasites. My opinion is based upon peculiarities in them other than, although strengthened by, the negative characteristic of absence of polliniferous organs. A negative cannot be proved, it is true, yet what has been positively asserted may as certainly result either from defective observation, or from too strong a desire to find no parasites among the AndrenidÆ. My reasons occur elsewhere in this work, and I need not repeat them. It is still an open question, and the young entomologist, if entering the arena unprepossessed, might win his spurs in determining it. It would be well worth the trouble of attending to for those who have leisure, and if decided in favour of the independency of these genera, which must be corroborated by a plurality of observations, and not confined to one locality, they would form strong and remarkable instances of a defective analogy in nature’s workmanship, and suggest looking further for the causes of so extraordinary an anomaly, and urge us to endeavour to trace the equivalent which supersedes it.

The main subdivision of the ApidÆ results from the habits of the insects, which divides them into SOCIAL and SOLITARY. The only tangible characters the social tribes present to distinguish them from the solitary is the glabrous surface of the posterior tibiÆ, with their lateral edges fringed with bristles slightly curved inwards, and which form, with the slightly indented surface of the limb, a sort of natural basket for the conveyance of pollen or other stores to the nest. This, however, has not been made use of as a main feature for scientific distribution, although they might follow the Dasygasters, as corbiculated bees, or little basket bearers, in which case they would form as pertinent a group as any of the rest, and the whole distribution of the bees, ApidÆ, would then rest upon the absence of, or the mode in which the polliniferous organs were present. But the wonderful attribute of their extraordinary instinct prohibits their being treated with the rest in a consecutive line, and renders it rationally imperative that all the Cenobites should group together in a section by themselves, and separate from the rest. Therefore in my arrangement I have not availed myself of this very natural character, and here indicate it, to show that I have not passed it from not noticing it.

Although the division into social and solitary yields in itself no tangible character whereby the insects may be separated, it being wholly empirical, yet is it so natural and necessary that it is impossible to gainsay it. We find the solitary section readily resolve itself into groups or subsections, determined by positive structural characters, indicative of certain habits, and having a conforming economy, besides which they are equivalents.

Thus the first subsection presents us with the brush-legged ApidÆ (Scopulipedes), which collect pollen upon their posterior legs. These are further subdivided into those which collect it upon the whole limb, viz. the coxa, the femur, the tibia, and first joint of the tarsus, (the femoriferÆ), and those which gather it merely upon the shank and basal joint of the foot (the cruriferÆ). These collectively form a well-defined group, and why Panurgus should be separated from the brush-legged bees, when it is a most conspicuous instance of the faculty, even more so than any other of the Scopulipedes, I have yet to learn. It is true its mode of collecting closely resembles that practised by the AndrenidÆ, as does also the furniture for the purpose of its posterior legs, but being essentially collocated with the ApidÆ or normal bees by its tongue, it fittingly links itself to the other brush-legged ApidÆ (which have hitherto been placed between the Dasygasters and the Social Bees), by means of the genus Eucera, by reason of its two submarginal cells, the structure of its maxillary palpi, its mode of burrowing, and by each being infested by a similar parasite—a Nomada, which in accommodation to the size of the sitos is the largest of the genus. Nomada does not occur as a parasite upon any other of the brush-legged bees, or indeed upon any other of the true bees at all, which peculiarity brings these two genera into close contiguity to all non-parasitical AndrenidÆ, all of which have their legs furnished with polliniferous brushes, and upon which subfamily, exclusively of these two instances of Panurgus and Eucera, Nomada is solely parasitical.

With respect to the two submarginal cells to the wings, nature must have some reason for the limitation, for we find it prevalent also throughout the Dasygasters, or hairy-bellied bees.

The next very natural group is consistently central. It comprises the cuckoo-bees, which are naked-legged (Nudipedes), by reason of their parasitism, they not requiring organs to collect what they have no occasion to use. Their parasitism extends both upwards and downwards, those with three submarginal cells being parasitical upon all the brush-legged bees, whether subnormal AndrenidÆ or the Scopulipedes, those with two submarginal cells being restricted in their parasitism to the Dasygasters.

These Dasygasters, or hairy-bellied bees, form the next very natural group. Their general peculiarity of structure I have had occasion to advert to, in treating, in a former section of the work, upon the structure of the imago, and to which I now refer to avoid repetition. This group contains the majority of the artisan bees, whose habits I shall particularize when I speak of the genera specially; but we find carpenters amongst the Scopulipedes, and essentially builders amongst the Cenobites, which form a further and the last of our natural groups. A true cuckoo-bee (Apathus) consorts amongst these Cenobites, and properly so, from many causes. The anomaly would have been too great to have removed it to a place amongst the Nudipedes, for although in obsolete paraglossÆ, and in a deficiency in the normal number of the joints of the maxillary palpi, it resembles some of these, its general habit and general structure, bating that controlled by its parasitical habits, are so like Bombus, that it cannot well be separated far from the latter,—especially as we know too little of its habits to say that it does not regularly dwell in the nest of its sitos, which may well mistake it for one of its own community, it resembling the species it infests so closely; it therefore consistently associates systematically with the temporarily social societies.

Having thus cursorily skimmed the surface of the method I suggest, I have next to give my reasons for proposing it in lieu of adopting any yet extant.

My exhibition of Kirby’s grouping, in the preceding section, where I treat of the scientific cultivation of British bees, will fully explain why I could not adopt that arrangement.

Why I cannot follow Latreille’s, is, that in his last elaboration, in his ‘Families Naturelles,’ published in 1825, which must be considered as his final view, he does not satisfactorily divide the AndrenidÆ, of the genera of which he has made a complete jumble. With the ApidÆ in his group of Dasygasters, he intermixes Ceratina, separating it from the group of Scopulipedes, where it truly belongs by every characteristic, and he mingles also with them the two cuckoo genera Stelis and Coelioxys, which are merely parasites upon these Dasygasters, and can only be associated by the structural conformity of the two submarginal cells to the superior wings, and the length of the labrum, the latter being a character of very secondary importance; and further, he dissevers the Scopulipedes in placing Panurgus at the commencement of the ApidÆ, and the rest proximate to the social bees.

Westwood, in his modification of Latreille’s system, certainly divides the AndrenidÆ better than his master had done, but he does not go far enough. Besides, he interposes Halictus and Lasioglossum, (the latter admitted as a genus merely out of courtesy to Curtis, who had elevated it to that rank in his ‘British Entomology,’ although it is nothing more than a male Halictus), between Sphecodes and Andrena with Cilissa, these having lanceolate tongues with lacerate paraglossÆ, whereas Halictus has a very acute tongue, and its paraglossÆ are entire, as is also the case with Dasypoda, from which Halictus is thus divided. In the ApidÆ, he does not separate the cuckoo-bees, but with Latreille intermixes Coelioxys and Stelis with the artisan bees, although without retaining Latreille’s convenient and suitable name of Dasygasters, for this group of mechanics. The same objection I take to his Scopulipedes as that expressed above, relative to Latreille’s.

Precisely the same fault I find with the AndrenidÆ of Smith, as that urged above with respect to Westwood’s. He is more careful with his ApidÆ, his CuculinÆ being all genuine parasites, but he includes Ceratina with the Dasygasters, with which it has no affinity of structure, and only a slight analogy in the form merely of its abdomen without its hairiness beneath, to that of Osmia, from whose proximity he takes it to place it near Heriades, when it is certainly intimately allied in every respect with the Scopulipedes, and by reason of its subclavate antennÆ might suitably be brought into juxtaposition with Panurgus, did not its obsolete paraglossÆ and three submarginal cells interfere with its occupying this position. To his Scopulipedes the same objection is valid as that taken to Latreille’s and Westwood’s disposition of them. Amongst the social bees he separates Bombus from Apis, by the intervention of Apathus, which is scarcely consistent.

It is in no spirit of captiousness that these objections are made; they are deduced from collocations whose conspicuous incoherence is patent to the most superficial observation. The distribution I have here introduced has been made merely to ameliorate, and make more cogent, what was so palpably defective and feeble.