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WILD BEES, WASPS AND ANTS

LONDON GEORGE ROUTLEDGE & SONS, LIMITED NEW YORK: E. P. DUTTON & CO.

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PREFACE

The object of this little book is to give in as simple a form as possible a short account of some of the British Wild Bees, Wasps, Ants, etc., scientifically known as the _Hymenoptera Aculeata_. Of these the non-scientific public rarely recognizes more than the Hive Bee, the Humble Bee, the Wasp, and the Hornet, whereas there are about 400 different kinds to be found in this country, and they can be recognized by any one who is disposed to make a special study of the group.

The author has not hesitated to make free use of the experiences of others in regard to the habits of the insects he describes, and he has not thought it necessary in each case to make separate acknowledgment of this. He takes this opportunity of thanking Mr. H. Donisthorpe and Mr. F. W. L. Sladen for assistance in the chapters on Ants and their Lodgers, and Humble Bees, respectively. {vi}

These pages are written only for the non-scientific, as the scientific entomologist will be already familiar with the elementary facts recorded; but it is hoped that they may be of interest to lovers of Nature who wish to know a little about the insects they see round them and how they spend their lives. Of this knowledge very little exists, as the scraps which have been here brought together evidence. There is an immense field open for research and observation, and the writer of this little book will be very glad if the following pages should encourage any one to take up the subject and add to our present scanty stock of information.

EDWARD SAUNDERS.

ST. ANN'S, WOKING.

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CONTENTS

PAGE

THE SUBJECT IN GENERAL, 1

THE SOLITARY GROUPS, 6

THE SOLITARY BEES, 9

THE CUCKOO BEES, 14

THE FOSSORS, OR DIGGERS, 18

THE SOLITARY WASPS, 24

THE SOCIAL GROUPS, 28

THE ANTS, 31

THE SOCIAL WASPS, 35

THE HUMBLE BEES, 39

THE BEES WITH BIFID TONGUES, 44

THE BEES WITH POINTED TONGUES, 48

LEAF-CUTTING BEES, 52

_Osmia_ AND ITS HABITS, 55

A COLONY OF _Anthophora_, 61

BEES AND POLLEN-COLLECTING, 65

ON BEES' TONGUES, AND HOW THEY SUCK HONEY, 72

A DREADFUL PARASITE, 77

{viii} AMONGST THE BEES AT WORK, 81

ANTS, THEIR GUESTS, AND THEIR LODGERS, 88

HOW CAN AN "ACULEATE" BE RECOGNIZED?, 92

MALES AND FEMALES, 95

THE VAGARIES OF COLOUR AND STRUCTURE IN THE SEXES, 100

THE DISTRIBUTION, RARITY, OR ABUNDANCE OF VARIOUS SPECIES, 105

ON BEES' WINGS, 110

ON BREEDING ACULEATES, ETC., 113

ON COLOUR, 119

THE DEVELOPMENT OF INSECTS FROM THE EGG, 124

ON STRUCTURE, 132

INDEX, 141

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LIST OF ILLUSTRATIONS IN THE TEXT

PAGE

Fig. 1. _Bombus_, larva and nymph: after Packard 11

" 2. _Ammophila_ 22

" 3-4. Spines on the tarsi of female _Ammophila_ 23

" 5. Tubular entrance to hole of wasp 25

" 6. Basal segments of ants 33

" 7. Rose-leaf partially eaten by bees 52

" 8. Tufted hairs of hind leg of _Andrena_ 67

" 9. Corbicula of humble bee 67

" 10-12. Cleaning apparatus of bees 69

" 13-18. Hairs of bees, magnified 71

" 19. Tongues of bees, magnified 73

" 20. Diagram of tongue of bee 75

" 21. _Stylops_ 77

" 22. _Stylops_ larva in abdominal cavity of bee 78

" 23. Antennae of "Keyhole" wasps 101

" 24. Legs of male "Keyhole" wasps 101

" 25. Tibia of male _Crabro cribrarius_ 103

" 26. Antennae of male _Crabro cribrarius_ 103

" 27. Head of male and female _Crabro clypeatus_ 103

" 28. Parts of the insect 133

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DESCRIPTION OF THE COLOURED PLATES

PLATE A

Figs. 1, 2, 3. _Formica sanguinea Latr._: male, female, and worker. The host of _Lomechusa_ (p. 89), also a slave-making species; makes irregular nests of dead leaves, etc., generally against a sloping bank.

Figs. 4, 5. _Mutilla europaea Linn._: male and female. One of the few British species of Aculeates where the female is wingless; found in sandy places running in the sun.

Fig. 6. _Cerceris arenaria L._: female; burrows in the sand, and provisions its nest with beetles (p. 20).

Fig. 7. _Ammophila sabulosa L._: female; burrows in the sand, provisions its nest with caterpillars, peculiar for its very elongated waist (p. 22).

Fig. 8. _Crabro cribrarius L._: male; peculiar for its paddle-like tibiae and flattened antennae (p. 103).

Fig. 9. _Odynerus spinipes L._: male; peculiar for the form of its middle femora, which are cut out almost in two semicircles (p. 101); female makes a tubular entrance to her nest (p. 25).

PLATE B

Fig. 10.--_Colletes succinctus L._: female; lines its cells with a gluey material (p. 44); colonizes in sandy banks; host of _Epeolus rufipes_ (fig. 19).

{xii} Fig. 11. _Sphecodes subquadratus Smith_: female; cuckoo of a species of _Halictus_; female hibernates like its host (p. 17).

Fig. 12. _Halictus lencozonius Schr._: burrows in the ground; the host of _Sphecodes pilifrons Thoms_ (p. 17).

Fig. 13. _Vespa crabro L._: female (the Hornet), nests in hollow trees; host of the rare beetle _Velleius dilatatus_ (p. 38).

Fig. 14. _Vespa vulgaris L._: female: one of our commonest wasps; nests usually in the ground (p. 35); host of a peculiar beetle (_Metoecus paradoxus_) (p. 38)

Figs. 15, 16. _Andrena fulva Schr._: male and female; the bee which burrows in lawns, etc. (p. 9); host of _Nomada ruficornis var. signata_ (p. 15).

Fig. 17. _Panurgus ursinus Gmel._: Female; legs loaded with pollen, burrows in hard sandy paths, etc. (p. 49). Males sleep curled up amongst the rays of yellow composite flowers.

Fig. 18. _Nomada ruficornis L. var. signata_: cuckoo of _Andrena fulva_ (figs. 15 and 16).

Fig. 19. _Epeolus rufipes Thoms_: female; cuckoo of _Colletes succinctus_ (fig. 10).

PLATE C

Fig. 20.--_Megachile maritima Kirby_: female; burrows in the ground, makes its cells of pieces of leaves, which it cuts out with its mandibles; host of _Coelioxys conoidea_.

Figs. 21, 22. _Coelioxys conoidea Illig_: male and female; cuckoo of _Megachile maritima_.

Fig. 23. Burrows of _Megachile Willughbiella Kirby_, in a piece of rotten willow; each burrow originally contained six cells, but two of the left-hand series have been lost.

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PLATE D

Figs. 24 and 25. _Anthophora pilipes F._: male and female. A spring bee, the male of which may often be seen in gardens, darting from flower to flower (p. 81); while the female collects pollen; it forms large colonies (p. 62).

Fig. 26. _Melecta armata Pz._: cuckoo of _Anthophora pilipes_.

Fig. 27. _Anthidium manicatum L._: invests its cells with the down off the stems of labiate plants, which it strips off with its mandibles (p. 50).

Fig. 28. _Osmia bicolor Schr._: female; nests in snail-shells, which it sometimes covers up with small pieces of grass-stems till a little mound is formed, resembling a diminutive ants' nest (p. 59).

Fig. 29. _Bombus terrestris L._: female. One of the commonest of our Humble Bees; it nests in the ground. It is the host of _Psithyrus vestalis_, which resembles it very closely in colour; it is this species that was exhibited by Mr. Sladen at the Maidstone Agricultural Hall (p. 41).

Fig. 30. _Bombus lapidarius L._: another common Humble Bee, also an underground builder; it is the host of _Psithyrus rupestris_.

Fig. 31. _Psithyrus rupestris F._: female; the cuckoo of _Bombus lapidarius_, which it closely resembles except for the nearly black colour of the wings.

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THE SUBJECT IN GENERAL

I think I ought here to say why I propose to limit myself to an account of a certain portion only of the Hymenoptera. The reason for this, in the first place, is that the section which I have selected is the only one of which I have any special knowledge; it consists of the bees, wasps, ants and sandwasps, four groups which make up the stinging section of the order--or perhaps more accurately, which have poison bags connected with their egg-laying apparatus or _ovipositor_. Another reason for their selection lies in their nesting habits; these enable one to get a further insight into their economy and ways than can be obtained from those of almost any other group or order--at any rate they make them comparatively easy to study; one can, so to say, find these little creatures at home, whereas in most orders there seems to be no definite home to which the {2} individuals may be traced; a great advantage also in selecting the stinging groups for study is that they are creatures of the spring and summer, and of the sunshine, so that the weather which tempts them out to their duties is of the kind most agreeable to those who wish to investigate their habits.

The habits of the hive bee have not been touched on, as so many excellent treatises have been written on them that any observations here would be superfluous.

Although these groups are distinguished by their stinging habits, it is only the female that possesses a sting--the male is a most harmless creature and quite incapable of injuring any one. A male wasp or even a male hornet may be handled with absolute impunity, only it is wise to be certain as to the sex of the individual before presuming to play with it too much! A word here may perhaps be said about stinging. People often talk about a gnat stinging or a stinging fly; it may be difficult to define exactly what "to sting" means, but the writer has always considered that a sting is inflicted by the tail end of the creature or a {3} bite by the mouth. A fly or gnat no doubt inserts its proboscis into one's flesh just as a wasp does its sting; but the actions of such opposite parts of the body surely demand distinct names. As we have been alluding to flies it may not be inappropriate to say here that all the creatures we are going to consider have four membranous wings except the worker ants and a very few forms which are comparatively seldom met with. By this character they may at once be known from flies, which have only two membranous wings. The large brown "drone flies", so often seen on the windows of our rooms, especially in autumn, and which most people mistake for hive bees, to which they certainly bear a considerable general resemblance, may be detected at once by wanting the two hind wings of the bee.

The "aculeate", or stinging, Hymenoptera, are divided into sections and families according to their structure; but the groups which stand out most clearly in regard to their habits are the solitary and social species, the predaceous and non-predaceous and the inquilines or cuckoos. {4}

The vast majority of the aculeate Hymenoptera are what are called "solitary", i.e. one male and one female alone are interested in the production of the nest; but there are also three "social" groups--the ants, the true wasps, and the humble and hive bees.

These are called social because they form communities and all work together towards the maintenance of the nest. In the social species there are two forms of the females--the queens and the workers; these latter have the ovaries imperfectly developed, and in the humble bees and wasps they only differ outwardly from the fully developed females or queens by being smaller. In the ants, however, the workers are wingless, and of a very different form from that of the queen. The role of these workers seems to be to do the general work of the nest; they have been known to lay fertile eggs, but the resulting offspring has always been male.

Between these conditions of solitary and social we know of no actually intermediate stages. We do not seem to see any attempts on the part of solitary bees to become social or vice versa. The only condition known which {5} could possibly be considered as intermediate is shown in certain species where a number of individuals make their nests close to each other in some particular bank, forming a colony. These colonies are sometimes very extensive, and the burrows of the individual bees very close together; it has also been shown that the burrows sometimes unite--at the same time there seems to be no positive evidence that there is any work done in the colony which could be considered as done for the common good.

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THE SOLITARY GROUPS

All the solitary kinds appear to feed themselves on vegetable juices, honey, etc., but there is a well-marked division between those who provision the cells of their offspring with insects, either fully developed or in the larval stages, and those who provision them with the pollen of flowers, honey, etc. The theory is that originally all fed their cells with insects, but that by degrees the more progressive found that the food which suited themselves would equally nourish their offspring, and accordingly provided them with vegetable nourishment. We find no intermediate stages. A certain class still goes on feeding on the old principle. The members of this class are known as "_fossors_" or diggers, while those which feed on the new principle are called "_Anthophila_" or flower-lovers. These are not very happy names, as many of the _Anthophila_ dig out holes for their nests just {7} in the same way as the _fossors_ do, and many of the _fossors_ are found in flowers, apparently enjoying them just as much as a truly anthophilous species would, although no doubt often with the ulterior object of capturing some insect for their young! Still these names are known as representing these two sections all over the world, and therefore it is better to keep to them even if they are not as descriptive as one would like them to be.

The _fossors_, or "diggers", have all comparatively short and bifid tongues, and have, as a rule, little in the way of hairy covering, and what hairs they have are simple and only in very rare instances branched or feather-like. The hind legs of the females are not modified in any way so as to enable them to collect pollen, their legs are usually long and slender, and they are admirably adapted to their life habits of hunting spiders, insects, etc., for their young.

On the other hand, the _Anthophila_ or "flower-lovers", are specially adapted for pollen collecting. Their tongues vary from a short form like that of some _fossors_ to the long tongues of the humble bees. Their hairs are always plumose {8} or branched on some part of the body and the hind legs of the females in most species are provided on the tibia or shin with a special brush on which pollen may be collected. In some of the long-tongued bees, however, this brush occurs on the underside of the body instead of on the tibia. The pollen-collecting arrangements of the different genera of the _Anthophila_ and the corresponding organs for cleaning off the pollen again are amongst the most interesting instances of modification and adaptation: some of the more striking of these will be mentioned later on. (See pp. 65 _sqq._)

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THE SOLITARY BEES

The life-history of an ordinary pair of solitary bees is, roughly, as follows: I will take for an example one of the spring species of _Andrena_. Many people know the little red bee, which for some apparently unaccountable reason suddenly appears in myriads on their lawn or gravel path, throwing up little mounds of finely powdered earth--in this respect being quite different from worm casts, which are formed of wet mould and the particles of which cling together--sometimes causing considerable alarm as to the possible effect on the lawn. These have hatched out from burrows made by their parents in the previous year, the mouths of which have been filled up with earth and therefore are quite invisible till the newly fledged bees gnaw their way out. They, in their turn, are now making fresh burrows for their own broods; possibly they infested some one else's lawn the year before or were only in comparatively small {10} numbers on the lawn under notice and so passed unrecognized. They may safely be left alone, as they never seem to breed many consecutive years in one such locality: probably the treatment of a lawn does not suit them, mowing and rolling upsetting their arrangements. We will now consider these arrangements. The female bee, so soon as she realizes that she is charged with the duty of providing for her future offspring, makes a burrow in the ground, and the earth thrown up from the tunnel forms the little heap which is so observable; this burrow varies in depth from 6 to 12 inches and has short lateral branches; each of these she shapes, more or less, into the form of a cell, provisions it with a small mass of pollen mixed with honey for the maintenance of the larva when hatched, and lays her egg; she then seals up that cell and proceeds to the next, and in this way fills the burrow up until pretty near the surface. The bee caterpillar when hatched is a white grub-like creature which, after devouring the food provided for it, becomes more or less torpid; it then makes its final change of skin, after how long a period is probably uncertain, and appears in the nymph stage. {11} [Illustration: FIG. 1. Bombus, larva and nymph: after Packard.] This stage corresponds to the chrysalis of a moth or butterfly, the creature being shortened up and rather more like the perfect insect compacted into the smallest form possible. People are often misled into the idea that the caterpillar forms the chrysalis over its former self, whereas the chrysalis has been all the time forming inside the caterpillar and only shows itself when the final skin is shed; of course some caterpillars spin a cocoon over themselves before they change their skin, but then the true chrysalis is found inside the cocoon. A curious fact connected with the change from the nymph to the perfect insect is that this takes place sometimes as early as August in the year preceding their appearance; so that cells dug up in August may contain fully fledged insects which are not due to appear till April or May of the following year. It is wonderful also how long life can be {12} sustained by these creatures in the "full-fed larva" condition. Some years ago I collected a number of pierced bramble stems in order to breed out some of the small "sandwasps" which nest in them. On opening them in May, when the perfect insects are generally ready to appear, I found that several of the larvae had rather shrunk up and had not changed into nymphs. These I left in the stems, covering them up again, and they appeared as perfect insects in the May of the following year.

The account given of the nesting habits of the above _Andrena_ of our lawns, etc., is more or less true of nearly all the solitary bees. Their methods vary, some burrow in the ground, some in old wood, some in snail shells, some in bramble stems or straws or the hollow stems of various plants, some in holes or crevices in walls, etc., and their methods of building their cells vary exceedingly: all of these are of great interest and some display an ingenuity which is quite surprising. Of these special nesting habits some of the most striking will be mentioned later on.

Before leaving these general remarks on the {13} solitary bees the habits of two genera must be specially noticed, as they differ in an essential point from those of the others. These are known to entomologists under the names of _Halictus_ and _Sphecodes_.

In most species of these the males and females of the new brood are not hatched out till after midsummer, and no work is done for the provisioning of new burrows that autumn; but the female, after having undertaken the duties of maternity, hibernates, i.e. goes back into a burrow and lives there till the next spring, the males dying off before the winter. In the spring the [female] wakes up and does the necessary work for the future brood just as any ordinary spring bee would--but there are no attendant males--the duties of that sex having been performed in the autumn. The larvae contained in these burrows hatch out after midsummer and therefore never spend a winter in the ground. In this respect they resemble the social bees and wasps, about which more hereafter; in the meanwhile a few words must be said about the cuckoos or inquilines, which are perhaps the most interesting creatures of all.

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{14}

THE CUCKOO BEES

These cuckoos live at the expense of their hosts. The mother of the industrial brood makes her cell and provisions it, and lays her egg. The cuckoo bee manages to enter also and lay her egg in the same cell, the usual result being that the cuckoo devours most of the food instead of the rightful offspring, which gradually gets starved and dies, the cuckoo appearing in its place; but there have been cases, how frequent they are is difficult to say, in which both offsprings have emerged.