BRITISH ANTS THEIR LIFE-HI~TORY AND CLASSIFICATION BY ~ \ H. ST. J. K. DONISTHORPE, F.Z ..~ ~~~~~\:~ With 18 Plates and 93 Diagrams LONDON GEORGE ROUTLEDGE AND SONS, LIMITED BR.OADWAY HOUSE: 68-74 CARTER LANE, KC. 1927 BRITISH ANTS - ~ ~~ " \\ u_3. F'1' (b~3 • Printed in Great Britain at The Mayflower Press, Plymouth, William Brendon & Son, Ltd, DEDICATED AS A TOKEN OF ESTEEM AND FRIENDSHIP TO WILLIAM MORTON WHEELER PH.D., PROFESSOR OF EdoNolllIC ENTOMOLOGY HARVARD UNlVEltSITY, U.S.A. • PREFACE TO SECOND EDITION WHEN the first edition of this book was written I did not~tiCiPate that in ten yea.rs' time a second edition would be requ . R~lizing, however, that the copies were nearly e " 81 sted I decided to accept MessrS. Routledge's offer to prepare a econd edition -bringing the work up to date, and also to write a ew book devot;ed exclusively to the British Myrmecophilous fauna.\ T~ former is now, complete; the latter I hope to acco~lish in the dot distant future. \ The study of ants has made considerable progress duriJ;tg the last ten years, and many important 'memoirs and books hav\ been published-some of them to be briefly referred to below. . \ The increase in our knowledge of the distribution of the B~.is h Fotmicidae has been disappoij_ting. Entomologists often do ot appear to realize that for the purposes of a local list the common, at species are quite as necessary as the rarest. If the reader will kindly note from the information supplied in this book, the counties aIill. vice-counties from which our commonest ants have never been, recorded, and as opportunity offers will make a point of searching and of ma.king his observations available, our knowledge of the distribution of British a.nts would soon be greatly increased. In the meantime I desire to thank Messrs. Britten,.Butterfield, Hallett, Harwood, Hodson, Joy, Keys, C. Nicholson, Perkins, Phillips, Saunt, Stelfox, S. O. Ta.ylor, and Waterston for kindly sending me ante} to determine, and information concerning their occurrence in Britain. Since the first edition of "British Ants" was published two new species have been discovered-Myrmica se'hencki Emery, by Mr. 'lI. M. Hallett, and AcanthomyopB (Donisthorpea) brunneus Latr., by Dr. N. H. Joy; also two new varieties are added-Ponera Ill'll,nctatissima var. exada Santa., due to Dr. F. Santschi; and r:anthomyops (Ohthonolasiua), umhratus Nyl. var. aflino-umhratus br. nov., by myself. Among recent works which the student of Myrmecology will find viii PREFACE TO SECOND EDITION essential I place first, Chapterij IV and V in W. M. Wheeler's "Social Life among the Insects" (1923) .• It is truly remlU"kable that the author should have been able to compress into this limited space such a store of information on nearly every subject connected with ants. Also' his great work on " The Ants of the :Relgian Congo" (1921-22), consisting of over 1000 pages and containing a .key to all the genera. of the ants of the world, should be consulted by all myrmecologists. The same author's" Ants of the Baltic Ambm-" (1914). is also a most useful and instructive work; and to those who are interested in the geological history of ants, attention ma.y be directed to papers by Prof. Cockerell and myself on the British fossil ants, of which the titles will be found in the Bibliography at the end of this volume. • Emery, whose death on May 11th, 1925, leaves a void which can never be filled in the ranks of myrmecologists, had, fortunately for science, completed his great work in the « Genera Insectorum," 1901-25. Among his recent papers must be mentioned also a valuable memoir on the ants of Italy published in 1916. During his life he wrote some 295 publications on ants; not only was he a myrmecologist of the highest eminence, but also a just and generous man, who always gave full credit to others for their work. Forel in 1915 republished the taxonomic part of his "Fourmis de la Suisse," bringing the synonymy up to date; and in 1923 he completed the five volumes of "Le Monde Social des Fourmis " of which an English translation will shortly appear. This, although in many ways a fine work, is somewhat disappointing in that it is not up to date, and that the opportunity has been made for airing the author's socialistic views. I should wish in particular to protest against the ~ts being employed as a supposed weapon in political controversy. In my opinion an entomological work is not the appropriate means for the introduction of political theories of any kind, still less for their glaring a.dvertisement. Let those, however, who are so ready to set forth'the social life of ants as a lesson to human beings, and an argument favouring a socialistic community, reflect on the following fa.cts :-To all intents and purposes the working classes of ants are sterile! They have no trade union rules; each worker does as much work as she can from . early morn to dewy eve, and often during the hours of the night as well, All are willing to sacrifice their lives at a moment's notice • for the good of the state, and are ever industrious and contented. PREFACE TO SECOND EDITION ix leveloped heads in order to contain the powerful muscles of the aws with which they ,crush "the hard seeds reqUired for food; ,ut when these workers are no longer needed by the colony, the ither ants cut off their heads and throw them on the refuse heap. rhis is a very' drastic, but effective, method of getting rid of a uperfluous working class . . Wasmann,. whose interesting papers on ants and myrmecophiles lumber up to the present date some 250, published in 1925 a 'aluable work on ant mimicry, which is 'indispensable for those Ilterested in that subject. Schmitz, in 1916, ~ote an interesting .ccou1J.t of the Ants and Myrmecophiles of Holla ij; and Bondroit, 11 1918, a book on the Ants of France and Bel ~. The value of he latt~r is lessened, however, by the unnecess rJl multiplication If species, and the creation of many on the poore of characters. The attention of those interested in the M~'eCOIOgy of other ontinents is directed to the following public tions--Arnold's ompletion of his monograph on ·the ants of South . rica, 1915-25 ; he papers on Australian ants, by Clark, Crawley, :.and Wheeler; un ants of Africa, South America, etc., by Santschi \ and on ants and myrmecopbiles in the Argenthle, etc., by Bruch. \ At the end of, the novel "The Ants of Timothy T~iimmel," by A. Ferenczy (1924), w:41 be found some sixty pages of ~tes on the social life of ants, compiled with the assistance of Mis Cheesman and myself. . Finally, mention may be made of the chapter written\ by me in "The Life-Work of Lord Avebury (Sir John Lubbock)," 1924, edited by his daughter, The Hon. Mrs. Adrian Grant Duff. I wish to express my best thanks to Mr. C. K. Ogden to whose good offices the publication of this edition is due; to Miss\F. J. Kirk for compiling the Bibliography and the Index, and for hltIp in other ways; to Mr. Engel Terzi for translations from the Ita~n ; and to Professor E. B. Poulton for kind help and encouragement with the revision, etc. HORACE DONISTHORPE. J 'Une 30th, 1926. CONTENTS \\ . PAGE ZFAOE vii , l'BODUOT,ORY \: 3 ~RNAL STRUCTURE 3 hE HlIlAD . 3 hE THORAX r 7 \ THE WINGS. \. 9 .\ THE ABDOMEN , , 9 . THE GENITALIA I, \, 13 17 !l'Ji:&l\TAL STRFCTV'RE '\ bE GLANDS 19 \ GLANDS IN THE HEAD 19 , THORACIC GLANDS 19 ABDOMINAL GLANDS. 20 B»PRODUCTIVE ORGANS 22 THE CIRCULATORY SYSTEM .\ 25 THE RESPIRATORY SYSTEM .~5 THE,MusOULAR SY;STEM' . 5 THE NERVOUS SYSTEM. 26" [FE HISTORY • 26 PRoPAGATION 26 CoLONY FOUNDING 27 METAMORPHOSIS 29 PARTHENOGENESIS 30 LARVAE 30 PuPAE' 33 . POLYMORPHISM 34 5YOHOLOGY • 44 HoW' ,ANTS KNow EAOH OTHER 48 • CONTENTS PAGK How ANTS COMMUNICATE WITH EACH OTHER 48 • How ANTS FIND THEIR WAY 49 GEOGRAPHICAL DISTRmUTION 52 GEOLOGICAL RECORD • 53 55 COLLECTING MOUNTING AND PRESERVINU • 57 OBSERVATION -NESTS 58 :t INDIGENOUS GENERA AND SPECIES PONEIUNAE 67 PONEBA 77 MYBMIOINAE. 79 MYBMEOINA • FOBMICOXENUS 88 95 ANEBGATES 102 MONOMORzUM 109 80LENOPSIS MYRMIOA 115 152 8TENAMMA 161 LEPTOTHOBAX 189 TETRAMORIUM 199 DOLIOHODERIN AE 199 TAPINOMA 205 FORMIOINAE • 206 ACANTHOMYOPS 281 FORMICA COSMOPOLITAN AND INTRODUCED SPECIES • 384 385 PONEIUNAE 387 MYRMICINAE. 394 DOLIOHODEIUNA E 396 FOBMICINAE • 407 BmLIOGRAPHY • 429 SYSTEMATIC INDEX: ANTS AND MYRMECOPHTLES LI~T OF IL~~STRATIONS PLATES SEPAR~~~LY PRINTED PJ!IA.B , \ FACING PAGE I. Eggs, larvae, and pupae of Po~ra coarriata, Myrmecina gramini- ,-_- cola, Solenop8is fugax, and M~mica ruginodis • . . 30 II. . Eggs, larVae: and pupae of nergate8 atratulu8, Monomorium pharaoni8, Leptothorax inteTTu tus, and Tetramorium caespitum 32 (II. Eggs, larvae, and pupae of TaPlnoma erraticum, Acanthomyop8 • niger, and Formica rufa . ~ . . . . . • 34 IV. Ergatan~ron1orph. of Formica ru arbis, and Gynandromorph of Formtca 8angUtnea . - ',' . . _ . • 42 V., Male, female, and worker of Ponei'f! coarctata, P. punctatissima, I and Myrmecinagraminicola. ' 82 VI. Male and female of Anergate8 atratiljlu8, and male, female, and worker of Formicoxenu8 nitidul1l8 ~d Monomorium pharaoni8. 98 rn. Male, female, and worker of SolenopBi,s fugax and Myrmica Bul- cinodi8 ' \ ,- 114 [II. Male, female, and worker of Leptothorax interruptu8, L. acervorum, and Stenamma westwoodi' 160 tx. Male, female, and worker of Tetramorium caeapitum and Tapinoma eTTaticum \ 198 X. Male, female, and worker of Acanthomyop8fuJ.tgino8Us and A. alienus 206 XI. Carton nest of Acanthomyop8fuliginosu8 and ~lUndB of A. flat"l!8 . 216 :;11. Nest of Acanthomyop$ umbratu8 266 [11_ Nests of Formica rufa . " 290 IV. Nests of Formica TUfa v. alpina and F. praten8i8 . 308 (V. ·Male, female, and worker of Formica pratensis 310 VI. Nests of Formica pratensis and F. ex8ecto 318 rn. Male, female, and worker of Formica rufibarbis 370 [II. Nests of Formica exsecta and F. picea . 382 FIGURES PRINTED IN THE TEXT -WOrker of FQl'miea praIe1ui8 • PAGE2 UnderVde_ of bead of Formiea pratenBi8 ljj 4 Mandible of hrmiea ~ !.! 5 Maxilla. of Formiea praUnBi8 ~ 5 Labium of Formiea pratenBi8 _~ 6 ~hora.x of Strt1Jlogna,tiU8 adMopiCUB S 7
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