LIBRARY OF THE UNIVERSITY OF ILLINOIS AT URBANA-CHAMPAIGN 595.79 Sa8h § WJ Biology The person charging this material is re- sponsible for its return to the library from which ,t was withdrawn on or before the Latest Date stamped below. Theft, mutilotion, ond underlining of books ore reasons for disciplinory oction and may result in dismissal from the University UNIVERSITY OF lUINOIS UBRARY AT URBANA-CHAMPAIGN L16I — O-1096 HYMENOPTERA ACULEATA. p' IIYMENOPTERA ACULEATA BRITISH ISLANDS. A DESCKIPTIYE ACCOUNT OF THE FAMILIES, GENERA, AND SPECIES INDIGENOUS TO GREAT BRITAIN AND IRELAND, WITH NOTES AS TO HABITS, LOCALITIES, H.U3ITATS, ETC. EDWARD SAUNDERS, F.L.S. LONDON : L. REEVE & CO., Puhlishers to the Home, Colonial, and Iii'Uan Governmenii, 6, Hexbiktta Street, Cove.nt Garden. 1896. ^ "^^^ PEEFACE. There are few people who are not more or less interested in ants, bees, and wasps, and their habits, and yet there are very few who systematically take up their study. Probably this is, to a certain extent, due to the somewhat uncompromisingly scientific look of the literature offered to them, and it is hoped that the coloured figures in the larger edition of this work will remove, at any rate to a certain extent, the apparent difficulties of the subject. The study of the aculeates is full of interest. On account of their nest- making habits they lend themselves peculiarly well to an investigation of their life histories, which abound in interesting details. The diversity of their structure also, and the wonderful specialization of the various organs of each species to its necessary habits, give a field for examination and study possessed, the writer thinks, by no other group of insects. The fertilization of plants depends greatly on their visits, and there seems to be in many cases a distinct correlation of structure between insect and flower, the short-tongued bees visiting only flowers whose honey is near the surface, as in such orders as the Rosaca', Componlfx, &c., the flowers with long tubes, such as those of the Lnhiatr tribe, being dependent, so far as the Ilymenoptera are concerned, on the visits of the long-tongued bees. Hermann Miiiler, in his celebrated work, " Die Befruchten der Blinnen, &c.," has gone very carefully into this subject, and at the present time Messrs. Willis & Burkill are publishing, in iLl- ij>'-inSrs vi PREFACE. tbe "Annals of Botany," observations on "Flowers aad Insects in Great Britain," giving the names of the insect visitants of the various flowers in the localities where they have been able to make observations. The principal works on the Hymenoptera Aculeata of this country are the following: — Kirby, " Monographia Apum Anglise (1802)/' the classical work on the British Anthophila; Shuckard, "Essay on the Indigenous Fossorial Hymenoptera," 1837; F. Smith, " Catalogue of British Hymenoptera, in the Collection of the British Museum, Part I., Apidse," 1855, and 2nd edition of the same, 1876; " Catalogue of British Fossorial Hymenoptera Forraicidse and Vespidce," in the same col- lection, 1868; E. Saunders, "Synopsis of the British Hymenoptera" Heterogyna, and Fossores, Trans. Ent. Soc. Lon. 1880, pp. 20L-300; Biploptcra and Andrenidse, ibid., 1882, pp. 165-290; Apida% ibid., 1884, pp. 159-250. The above works give descriptions of all the British species known at the dates of their publication. Besides these strictly technical treatises, the Rev. W. Farren White has published, without date, a volume entitled " Ants, and their Ways," which gives a popular account of the habits of the ants of the British Isles, and, in an appendix, short descriptions of the species ; and Shuckard, in 1866, pub- lished his " British Bees," a semi-popular work, giving descriptions of the various genera of the Anthophila, with copious notes on Labits, &c., illustrated with coloured plates. Very many interesting observations and experiments on instincts and faculties are recorded by Sir John Lubbock, in tbe Journal of the Linnean Society 1874-81, and in his well-known popular work, " Ants, Bees, and Wasps," published in 1882. Of Continental works, other than monographs on special groups, the only one likely to be of use in the determina- tion of the British aculeates is C. G. Thomson's " Hymen- PREFACE. vii optera Scandinavia?," but this does not include the ants. Two exhaustive works ou European Hymenoptera Aculeata have been commenced: Schmiedcknocht, " Apida3 Europa;a3," and Andre, " Species des llymenopteres d'Europe et d'Al- gerie." Of the former, two volumes only have appeared, treating of the genera NumaJa, Psithyrus, Bomhus, An- drena, and Osmia ; while the latter, so far, has dealt only with the Heterogijna, Yespidx, and part of the Sphegidse. The present work owes much, as will be seen in its pages, to the important papers on special groups of v. Hagens, Haudlirsch, Kohl, Perez, and Schletterer, to the Council of the Linnean Society for permission to reproduce the figures of tongues on Plates 1a and 1b, and to that of the Entomological Society of London for similar permis- sion as regards the figures of details scattered throughout the plates of this work, as well as to the assistance in various ways of the following entomologists ; the late Frederic Smith, whose kindness to the author, in deter- mining doubtful specimens and in helping whenever applied to, will always be in his memory ; Messrs. A. Beaumont ; W. H. Bennett; G. C. Bignell ; T. R. Billups; E. A. Butler; Rev. E. N. Bloomfield ; J. B. Bridgman; G. H. Carpenter; H. S. Cuthbert ; C. W. Dale; F. Euock ; P. Freke; G. E. Frisby ; "Willoughby Gardner; W. H. Harwood; T. McGregor; R. McLachlau ; E. G. Marquaud ; Rev. T. A. Marshall ; P. B. Mason ; Rev. F. D. Morice ; R. Newstead ; R. C. L. Perkins; Vincent R. Perkins; A. Piffard; H. Ramsden ; G. A. J. Rothuey; F. W. L. Sladen ; Dr. H. Swale; W. H. Tuck; and to the following local lists : Devon, (Parjitt) ; Gloucestershire, ( V. IL Perliliis) ; Hastings and St. Leonards and vicinity ; Lan- cashire and Cheshire, {Gardner); Land's End, {Mtinjiiand) ; Norfolk, {Bridijmaii) ; R;igby, [Morice) ; Yorkshire, (7iVe- huch). It has been thought by some that notes on the distii- viii PREFACE. bution of the species outside the British Isles could be usefully added to the localities given, but there is so much doubt about the identity of many of the British and Con- tinental species which bear the same names, and different countries have been so unequally worked by collectors, that to define areas at present would, if the author may adapt an expression from a critic of his former work on llfiinptera Hdeioptcra, throw more light on the distribution of Hymenopterists than on that of Hymenoptera. The species of Hymenoptera Aculeata now known to be indigenous to this country number 374. Of these 20 belong to the Heterogyna, or Ants; 127 to the Fossores, or Sand Wasps; 23 to the Dqdop (era, ov true Wasps; and 204 to the Anfhophila , or pollen-collecting Bees. There are, no doubtj others yet to be found, and the author trusts that all who have doubtful specimens will send them to him for examination, as he is always ready to be of use if possible, and it is amongst the doubtful ones that additions to our fauna are likely to be discovered. Edward Saunders. St. Ann's, Woking Eehruarj/, 181)6. HYMENOPTEEA ACULEATA. INTRODUCTION. The Hymenoptera, whicli include the Bees, Wasps, Ants, Ichneumons and Savvflies, as they are familiarly called, may be known by their mandibulate mouths, complete metamor- phoses, their four membranous wings with branching nerves, enclosing a few comparatively large sized cells, or rarely almost nerveless, and by the form of the thorax, all the segments of which are exhibited dorsally. Some forms are wingless. The Aculeata may be considered as the highest section of the order, both on account of the ingenuity displayed in their domestic economy and also of their more highly specialized organs. The antenna) as a rule are thirteen jointed in the male and twelve jointed in the female; there are three ocelli in the vertex of the head, and two large lateral compound eyes ; these, however, are sometimes reduced to a single facet, or entirely absent. The abdomen is pedicellate, and the ? is armed with a sting, whoso wound owes its paiufulness to the poison ejected from its poison bag, but even this prominent character is not always present, the tvaQFurmicidw being destitute of a sting. Tiie sting is always retractile, and almost always hidden when withdrawn. In some few genera the J' is winged and the ? apterous, and in one or two exceptional instances the J is apterous, but as a rule both sexes have fully-developed 2 HYMENOrTERA ACULEATA. ■wings. In the social groups a second form of the ? exists, in which the ovaries are not fully developed. Such females are genei'ally called " workers," and in the Heterogyna or ants are always wingless. By social species is understood those of which many individuals combine in making a common nest. Of these there are three groups, the Heterogyna or ants, the Vespidee or true wasps, and the higher Apidse, which include the humble bees and the hive bee. All the rest are solitary, one ? making and provisioning its own nest. Many of the solitary species occur in colonies, and this habit of colonization is considered by some to be a tendency towards socialism. In general foi'm the Aculeata may be said to vary comparatively very little, especially among the Anthophila. The specific characters are often most obscure, and frequently the females of two species are so alike that in the absence of the c? their real names cannot be assigned to them with certainty. The genera are more easily defined, but the difficulty of discovering characters by which to sort the genera into families or groups is very great. The males and females are often very dissimilar, and in some groups rarely occur together. The inquiline (or cuckoo) genera, of which there are many among the Anthophila, call for special notice. Their species inhabit the ?ame nest or lay their eggs in the same burrows as the bees to whom the nests belong. They apparently do nothing towards the economy of the family into which they intrude, but lay their eggs so that their offspring shall have a better chance of nourishment when hatched than that of their hosts. Yet, notwithstanding this, they appear to live on friendly terms with the owners of the nests, and generally closely resemble them in appearance, although in a few cases the dis- similarity between them is very strongly marked. The inquilines, however, are always destitute of the specialized organs fur pollen collecting which characterize their hosts. INTRODUCTION. 3 The habits of the Aculesites are most diverse. All are believed to be vegetable feeders in the imago state, but iu the larval state some feed on animal and some on vegetable substances. Some nest in the ground, some in bramble stems, some in gate posts ; key holes and locks prove attractive to some ; old disused snail shells to others ; holes in walls are utilized by certain species, and so plastered up as to leave no suspicion of a hidden nest ; whilst others make mud nests, and suspend them from a twig, or attach them to a stem of grass. The Carder bees belonging to the genus Bomhus make a covering to their nests, which are placed on the ground, of moss, grass, etc., but some have been kcown to utilize a forsaken bird's nest. The habits of the various genera and species will be treated of under their respective heads. The metamorphoses of the Aculeata are complete ; the pupa3, which are either naked or enclosed in a cocoon, are almost exactly of the form of the perfect insect, except that the wings are enclosed in short pad-like cases. The larvte are pale, fleshy, and grub-like. A remarkable peculiarity exists in the transformations of these creatures. The first and part of the second segment of the larva unite to form the head of the imago, the fifth segment of the larva, which in other insects forms the first abdominal segment in the imago, is transferred during the insect's passage through its pupal conditions to the thorax, so that what looks like the metathorax in the imago is really formed partly of the first abdominal segment, aud the constriction which follows it is really between the first and second segments of the abdomen, instead of between the abdomen and thorax. This transferred segment has been called the " propodeum," by Newman, the median seg- ment by some authors ; but, for convenience sake, I shall treat it here as part of the metathorax, and call the first segment of the abdomen that which follows the second regional constriction. In position the Hymenoptera should probably be placed 4 HYMENOPTERA ACULEATA. at the head of the lusecta. Their economic and often social instincts tend to place them in this position, and their right to it is supported by their wonderfully specialized structure, the very strongly defined regional constrictions and the highly cephalized imago. In the arrangement of the families I have followed the old plan, as it is quite impossible to get a satisfactory linear arrangement, and therefore I have not thought it worth while to disturb the existing one of our lists. Fossil Hijmenoptera — The oldest known representatives of this order have been obtained from the Solenhofen State of Bavaria and the Purbecks of this country, both belonging to the Upper Oolite (Secondary Period), but the Hijmenoptera do not appear to have been common and generally distributed until towards the middle of the Tertiary Period. The remains of Hymenoptera are not uncommon in the middle and upper Eocene and the lower, middle and upper Miocene. In certain strata of middle Miocene age Professor Heer found the Hymenoptera more numerously represented than any other order of insects. Internal Anatomy. Digestive System. — The food passes through a groove at the back of the labium into an enlarged chamber which opens into the oral cavity, thence through the pharynx into the oesophagus, which runs as a narrow tube through the thorax, opening out at the base of the abdomen into an enlargement called the first stomach, honey sac, or crop. At the posterior end of this stomach is a constriction, or kind of mouth corresponding to the gizzard, which can be opened to let the food pass into the chyle stomach, or can be kept clo; ed, eg as to allow the contents of the crop to be regurgitated. Leon Dufour says that in the Crabronidie there exists a lateral stomach or pouch leading out of the cesophagus, sometimes on the left hand side and sometimes IXTRODUCTION. 5 on the right, aud varying considerably in form. Beyond this the oesophagus contiuues for a short distance as a narrow tube, and then dilates into the second or chyle stomach. The chyle stomach also receives the urinary vessels near its apex. Beyond the chyle stomach extends the smaller intestine which widens out posteriorly into the colon, which again narrows to the apical opening of the abdomen. The sides of the colon internally are usually strengthened by longitudinal plates varying in form and number, but in the genus Bomhxi^, Dufour says there is no trace to be found of this character. Salivary Si/stem. — The salivary glands in the Hymen- optera play a very important part in their economy, and in the hive bee are very largely developed. Some of these lie in the head, some in the thorax. The pharynx beneath is chitinized into a slightly concave plate called the hypo- pharangeal plate, near the front margin of which are two nipple- like openings, which lead into the salivary ducts, these diverge laterally into two systems of glands that lie near the centre of the head. Behind these are two other lateral systems in the head, and two more lying in the thorax; these latter four unite into one common duct, which terminates in a valvular opening near the base of the tongue on its upper surface. This description is taken from the ? of the houey bee; pi-obably modifications of these systems would occur in many of the families. Leon Uufour fails to find any salivary glands in some of the Fossores. At the base of each mandible there is a small gland, but there seems to be doubts as to its functional use. The Nervous Sytiieia, as in other insects, consists of a system of ganglia united along the centre of the insect by two longitudinal cords, from which ganglia nerves are distributed to the various organs. The cephalic ganglia are largely developed ; there are five or six abdominal ganglia, the rest being thoracic. The Circninforii Si/stnn consists of a dorsal vessel, \vhich 6 HYMENOPTERA ACULEATA. lies along the back of the abdomen, from wliiob tbe blood is circulated through the vessels by the contractions and dilatations of the organ. The Besinratury System is carried on through the agency of spiracles, which open into the air tubes or tracheee, which distribute the air through the system. These tracheae con- sist of two main branches dilating towards the base of the abdomeu into two very large vessels, which help the insect to sustain itself in the air. The spiracles in the Aculeata are not easy to see, as those of the abdomen are mostly at the base of the segments liidden by the overlapping of the preceding one. That on the first abdominal segment is however exposed, and those of the metathorax and propodeum may be easily examined. l\p,2ii'oductive System. — In the J' the organs of this system are, as in other insects, the testes, the spermatic ducts which lead from the testes into the seminal vessels, and the ejaculatory duct terminating in the intromittent organ. In some cases the testes are enclosed in a common sac ; in others distinct. In some the speimatic ducts lead directly from the testes into the seminal vessels ; in others the seminal vessels are at the ends of tubes which lead into the spermatic duct. In the ? the organs are the ovaries, each of which consists of a series of tubes united together at the ends, varying considerably in the different genera, the oviduct, the spermatheca and ejaculatory ducts. Amongst the social genera, where a worker form exists, its ovaries are less developed than in the ? . The ? is supposed to be able to voluntarily control the passage of spermatozoa into the eggs, so that (J or ? eggs can be laid at will, those unimpregnated always producing S offspring. Before closing these short remarks on internal anatomy, I must mention two other important sets of organs in the ? and 5, viz. the poison and wax-secreting organs. The former of these consist of a sac into which the secreting tubes enter, and which can empty its contents through the lyTRODUcrw.y. 7 canal of the sting. The latter, which only exist in the worker bees, consist of glands which secrete the wax from the blood, and from which the wax in a liquid state soaks through the abdominal wall, and is collected and cools on the ventral segments of the abdomen. ExTKRNAL Anatomy. The skeleton is hard and usually more or less shining and black, but yellow and red markings are frequent, and some species are entirely pale and testaceous. Many of the species of the Anthophila are very hairy, and in this section the hairs are plumose — at any rate those on the thorax — whereas in the other sections they are simple, or in some cases twisted. Head large, but varying little in general form. The face is vertical or nearly so ; the vertex usually convex and transverse, sometimes nearly quadrate. The compound eyi^s generally occupy a considerable portion of each side of the head, and are composed of many facets, although amongst the anis they are sometimes wanting, or are reduced to a single facet. Besides the compound eyes there are usually three ocelli on the vertex, arranged more or less in a tiiangle. The side of the face below and behind the eye is called the clipek. This varies much in length, and in some genera the eyes meet, or nearly meet, the mandibles. From each side of the face between the eyes spring the antennce. These are as a rule thirteen jointed in the S , twelve jointed ia the ? and 5. The first joint, which is very long in the ants, is called the scape, and the remaining joints the Jlagel- lum. The antennce in the Aculeates are mostly filiform, or very slightly thickened to the apex, but the basal joint is often much thicker than the others, the second almost always very short; in the ants the apical joints form a distinct club ; the lower central portion of the face is nearly always divided from the upper, and often from the side portions by distinct impressed lines. The area thus limited S HYAfENOPTERA ACULEATA. is called the clypeii^. Just above tbis in the ants is a small triangular space called the frontal area. On to the front margin of the clypens is hinged another piece or flap called the lahrum ; the inandihles articulate on to the cheeks at their apex, and vary very much in form and development, they fold across, below, or sometimes over, the lahrum,- under the labrum is the epipharynx. On the underside of the head is a deep wide groove into which the cibarial apparatus fits when at rest. The ceiling of this groove is the posterior side of the wall of the face ; the sides of the groove are sub-parallel and nearly perpendicular, and unite posteriorly in a semi-circular curve, anteriorly the groove is free except when closed by the mandibles folding across it. Into the sides of this groove articulate the cardiues, two narrow joints each slightly widened at the apex. On the widened apices of these swing the lora, which are two narrow joints united so as to form a /\ shaped body. These vary a good deal in length and width, and in some genera are wanting. From the centre of this f^ depends the submenfum, a more or less triangular, sometimes hyaline body. This is followed by the menhun, which is semi-tubular, and forms a sheath in which lie some of the softer parts of the apparatus. From each side of the apex of the meidinn spring the lahialjmJpi, whose joints may be either cylindrical or sheath-like, and from between them extends the llgida or lingua. The ligula itself is grooved posteriorly, and its anterior surface is traversed by very fine ridges set with bristly hairs. In the higher J/j/f?*' there isa ladle-like organ, or " houton " at the apex. The liijula may be short and bifid at the apex, as in the Fossures, ants and wasps, and the early genera of the Antliofhila called the ohtusilinyues, or short, wide, and pointed, as in the genera Ilalictus, Andrena, etc., or many times longer than wide, as in the higher Apidse. The upper side of the base of the ligula is ensheathed by the paraglossa\ which vary much in shape and size. All the central or labial organs are laterally ensheathed by the vrn.vilhe, which articulate to the widened apices of the I.XTRODUCTIOA'. 9 cardines just outside tlie articulation of the arms of the lorit. 1'be maxilla? are sbeath-like organs, each emitting near its centre laterally a palptif: called the ma.rillary palpuK, the number of whose joints varies greatly in the different genera ; two narrow sclerites, called the scleritcs of fhe Injpopharynx, extend from the oral groove, close to the articulation of the cardines, down to the base of the weritum, passing between the rftnxiUie, to whose basal membrane they appear to be attached. In many genera from its point of attachment springs a scale-like organ of variable shape frequently fringed with bristly hairs. Between all the sclerites and organs at the base of the cibarial apparatus extends a membrame limited anteriorly by the fclerites of the hijpoph(ir?/nx, the investing membrane forming a complete bag. The whole apparatus can be folded up and p'lcked away in the oral groove thus : — The blades of the maxillx in the higher Apidie fold back on to their stems, the lora fold back between the cardines, and the cardines are drawn up into the groove, so that their dista,l extremities point back- wards. In the short-tongued bees there is no folding of the maxillfe on themselves, so that the point of the tongue when folded up is anterior. THORAX.^This, as in other insects, is composed of three segments — the prothorax, me.