FOREST ENTOMOLOGY A, T. Qillanders ^'*JLVT^A_^5aL^ — Forest Entomology SB • • • / Forest Entomology A. T. GILLANDERS, F.E.S. WOODS MAXAGER TO HIS GRACE THE DUKK OF NORTHUMKF.KLANO, K.G. WITH 354 ILLUSTRATIONS SECOND EDITION William Blackwood and Son: Edinburgh and London 1912 ALL RIGHTS RESERVED Jlli. 7 19/8 I, T>EDICATED BY PERMISSION TO HIS GRACE THE DUKE OF NORTHUMBERLAND, K.G. PEEFACE TO SECOND EDEFrON. As author of a work on a subject which does not appeal to a very wide circle of readers, I realise that it is a very high compliment that a second edition has been called for in a comparatively short time. Hence there is no need for drastic changes. I have, however, tried to benefit by the kindly advice given by entomological friends and others interested in the subject ; therefore the chapters have been re-arranged more in accordance with the sequence of entomological classi- fication. I have tried also, as far as I possibly can, to correct the recognition of photos and drawings of several friends. In this respect the first edition showed a slight error in acknow- ledging special photos as emanating from a friend rather than the firm of which he was a partner, and to whom I am very grateful for assistance. I am specially indebted to Professor Carpenter, of the Koyal College of Science, Dublin, for correcting several points in the Introductory chapter. I have every reason to be gratified by the many kind letters received from all parts, — from the landowner seeking further advice with regard to some special insect pest ; and from the young forester or nature-student trying to begin making a VIU PREFACE TO SECOND EDITION. collection of forest insects. It is therefore with very pleasant feelings that 1 submit this new edition, in the hope that it may command a wider circle of readers as associated with the growth and cultivation of forest trees, — a subject which is becoming more important every year. Park Cottage, Alnwick, December 1911. PREFACE TO FIRST EDITION, In submitting a book on a branch of Natural .Science which has required any degree of research during its preparation, or which otherwise is intended as a guide for research, perhaps no one realises its shortcomings more than the writer. At least this is certainly the feeling of the author on the present occasion. The book was begun in a very humble way. Having always had a strong inclination for Natural Science, and further for studying subjects associated with the scientific aspect of forest trees, I began as far back as 1887, while residing in Cheshire, to write papers on Forest Insects for the Warrington Field Club. To that district, with its Natural History clubs or societies, and the many excellent men I met in connection with them, I owe a very deep debt of gratitude. I cannot in the present circumstances enumerate all to whom I am indebted for assistance in the present book, but my first teacher in Entomology was the late Mr Berry Kendrick of Warrington, who gave me most ungrudgingly all the help any one could possibly give to a beginner. Of the Warrington Field Club itself it is impossible to speak too highly as a Natural History society. Composed of some fifty members. X PREFACE TO FIRST EDITION. original papers were given every fortnight during the winter months, often accruing from investigation during the summer montlis, and the members freely criticised the paper of the evening. This, though perhaps not always appreciated at the time, was in itself a most excellent school. In a similar way I am indebted to the Manchester ]\Iicro- scopical Society for a great deal of ready help from its members. This Society, unlike the former, published the papers of its members, and I have therefore to sincerely thank it for publishing my papers on " Forest Insects," and, further, for kindly giving me the Ijlocks prepared from my own neg- atives, some of which I have used for the present book. I have also to thank the Chester Society of Natural Science for many extended courtesies given from time to time. The material for this book has been collected in Cheshire and Northumljcrland, but more especially the former county. I am also indebted to the Country Gentlemen's Association, the Highland and Agricultural Society of Scotland, and the Royal English Arljoricultural Society for publishing papers on Forest Entomology. Turning from societies to individuals, it is almost impossible to acknowledge without being invidious those who have assisted me in the subject - matter of the volume. Two names may be mentioned. Mr R. Xewstead, author of ' A Monograph of British Coccidte,' gave me from time to time much assistance on Scale Insects. Mr A. Flatters, Man- chester, has given me great help in microscopical manipulation, and has, in addition to the figures acknowledged with his name, photographed several specimens from my own micro- scopic slides, as, for example, the tiny Cecidomyia liies. As regards the first works from which I derived consider- able assistance, I may specially mention 'A List of Insects on PREFACE TO FIRST EDITION. XI Forest Trees,' by Mr S. L. Mosley, Hudderstield, and the splendid ' Manual of Injurious Insects ' by the late IMiss Orinerod. Of the latter T cannot speak too highly, inure especially as Miss Ormerod gave me very great encourage- ment by correspondence. As the period during which I have studied the subject has extended over twenty years, it is obvious that much original work has been done by many workers during that time. I have tried, so far as means or leisure would afford, to keep in touch with all original work. In order, therefore, that the student may get the full advantage of that, I have, whenever I found the work of others more suitable than my own, quoted from those writers. In most cases this has been done by direct permission of the author, but in all cases when giving an extract I have endeavoured to duly acknowledge the same. In this advanced age of ours, knowledge, more especially Entomological knowledge, is no longer the prop- erty of any one individual. Hence all quotations are freely given as epitomised knowledge suitable for the student. They are given to obviate at first the necessity of consulting other works, and yet at the same time be a guide as to what other works he may consult for fuller information. It will be apparent that the majority of quotations are of a systematic nature. Hence the great difficulty in preparing the book has been to amalgamate the systematic with the economic, and so maintain a due proportion. On the one hand, if too many details of a systematic nature were given, the work would not only have been very much overladen, but the practical man woidd be impatient in reading it : on the other hand, if too few systematic details were given, it would not act as a guide to fuller inqiuries, or form, as designed, a suitable text-book for students at agricultural and other colleges. The main Xll PREFACE TO FIRST EDITION. feature which is attempted is recognition of the insect from the damaffc, together loith systematic characters and life-history details. In studying the subject from the economic point of view, I have become convinced, from a prolonged study of the subject, tliat the economic student must of necessity study the system- atic side and associate himself with systematic men. Hence for example, I have added often, in cdenso, extracts from valuable works like Fowler's ' British Coleoptera,' &c. At the same time, the matter is so arranged that the beginner, or the practical man, may at first, if he chooses, skip them, and study the nature of the injuries, the life-history of the insect, and in some cases the remedies which may be adopted. If, how- ever, the subject is to be thoroughly studied, the systematic points must be ultimately mastered. With a view of making the work more reliable, I have submitted each chapter in proof to men who have specially studied particular sections or families of insects, and in all cases I am glad to count these specialists as personal friends. As the work embraces nearly all families of insects, this was considered all the more advisable, and though the changes made in the proof form by these specialists were compara- tively few, yet in every case they were most important from the systematic point of view. The following are the names of those who looked over the respective chapters in proof form — viz., Mr E. T. Connold, " Gall-Mites " and " Oak Galls " ; Mr R. S. Bagnall, " Cole- optera " ; Mr A. C. Forbes, " Scoly tid?e " ; Eev. F. D. Morice, " Saw - flies " ; Mr E. Newstead, " Scale - Insects " ; Mr J. Collins, " Lepidoptera " ; Mr F. V. Theobald, " Aphididre " and " Diptera." I have also to thank Mr J. F. Annand, Lecturer on Forestry, PREFACE TO FIRST EDITION. xiii Armstrong College, Newcastle, for kindly looking over the MS. previous to sending it to the publishers, and also for discussing many of the practical points with me. I am specially indebted to Commander J. J. Walker, K.N., Oxford, for kindly correcting the final proofs. With reference to the illustrations, for the loan of blocks or electros I am indebted to Messrs Sampson, Low, Marston, & Co., Ltd., for figs. 14, 16, 20, and 21; to Professor Miall for fig. 18 ; to Messrs L. Eeeve & Co. for figs. 39, 41, and 42 ; to the Eay Society for figs. 300 and 308, and also for granting permission to photograph figs. 251, 253, 255, 258, 282, 311, and 312; to Messrs Headley Brothers for figs. 175, 241, and 319 ; to the Proprietors of 'The Entomologist's Monthly Magazine' for figs. 157, 158, and 159. Mr W. E. Fisher has given me much encouragement from time to time, and I have to thank him for receiving from Messrs Bradbury, Agnew, & Co., Ltd., the large number of blocks — viz., figs. 47, 48, 49, 50, 60, 61, 78, 79, 92, 121, 126, 127, 163, 164, 209, 210, 211, and 212. I also owe thanks to Messrs Blackwood for figs. 45, 46, 72, 73, 79, 101, 102, 104, and 216, from 'The Forester,' by Dr J. Nisbet. With the exception of the figures taken from the valuable German works, and acknowledged under each figure respectively, all the others are original. Finally, I beg to say that I submit the knowledge con- tained in the book with a feeling that I have just about the necessary amount of knowledge to make a beginning rather than a finish ; and I trust that the student will take up the subject with the object of making a study of it on his own account, and verify each point by observation and rearing. In other words, the student must consider the work as an intro- duction only. Further, there is the most important point left XIV PREFACE TO FIRST EDITION. to the last — viz., the study of German literature. The Germans are our great teachers in this branch of knowledge, and I have added many points and illustrations from German sources with the distinct object of showing their superiority and inducing students to study their most valuable works. Park Cottage, Alnwick, May 1908. CONTENTS. PREFACE TO SECOND EDITION- PREFACE TO FIRST EDITION INTRODUCTION ... I. ERIOPHYID.K (PHYTOPTID/E) OR GALL-MITES II. COLEOPTERA (bEETLES) . iiL II — scolytid/E (bark-beetles) IV. HYMENOPTERA OAK GALLS V. n SAW-FLIES, ETC. VI. LEPIDOPTERA (mOTHs) . Vn. APHIDID^E (green-fly) . VIII. PART I. PSYLLID.E „ II. CIGADID.E IX. COCCID.E (SCALE-INSECTS) X. DIPTERA (two-winged FLIES) . XI. HINTS ON COLLECTING, PREPARATION, AND Xn. LIST OF TREES WITH IN.IURIOUS INSECTS XIIL BENEFICIAL INSECTS XIV. INSECTICIDES AND GENERAL REMEDIES APPENDIX ..... Vll ix 1 19 37 8.5 130 164 201 251 301 307 313 352 MOUNTING 376 388 403 . 407 418 423 \A^T (W ILLUSrUiVTIONS. INTRODUCTION. 1. Kggs oi Arctia mentha>^tri . •2. Eggs of Bombyx nemtria on birch .... .'}. Typical forms of larvie 4. Typical pupal form of an insect .... 5. Pupal case of Vanessa urtiaa G. First emergence of Tor- toiseshell Butterfly from pupal case 7. Emergence of Tortoiseshell liuttertiy more fully ad- vanced .... ■S, 9. Fully developed Tortoise- shell Butterfly 10. Typical mouth parts of in- sects .... 11. Compound eye of insect as seen under the micro- scope .... 12. Typical forms of antennae . 13. Typical form of leg . 14. Diagram of longitudinal section of an insect 15. Horizontal section througli the head of a blow-fly . 1 6. Diagram of the cliief trunks of tiie ti-acheal system of an insect 17. Trachea of butterfly . I 8. Food canal of cockroach . 19. Diagrams of the heart of an insect 20. Female genital organs of tile cockchafer 'A OK 4 21. Male genital organs of the cockchafer . . .17 ERIOPHYIDrE ((iALL-MlTES). 22. Eriophyes rudis . . 21 23. Abortive sv/oUen buds on hazel caused l)y Eriophyes np(dlan< liyniperda 201. Larva of Goat Moth 202. Pupa of Goat Moth 203. Zeuzera f&sr«7* 204. Orgyia antiqua 205. Pupa of Puss Moth on twig of birch 206. Dicranura n'liula 207. Pygm-a bucephala . 208. Larvaj of Buff-tip Moth on twig of oak . 209. Larva of Bordered White Moth .... 210. Pupa of Bordered White Moth .... 211. Fidonia piniaria {maXe) . 212. Fldonia piniaria (female) 213. Foliage of lime-tree eaten by larvse of Winter Moth .... 214. Leaf of wych elm eaten l)y larvie of Winter Moth . 215. Foliage of sycamore eaten by larvre of Winter Moth 210. Winter Moth : male, fe- male, and caterpillar . 217. Trachea piniperda . 218. Cone of silver fir parti- ally eaten by larva^ of Dioryctria ab/etella 219. Dioryctria abiefella . 220. Foliage of oak destroyed by larvaj of the Green Tortrix Moth 221. Foliage of hazel injured by larvse of Tortrix ribeana 222. Leading shoot of poplar injured by larva of Hedya oceUana . 223. Shoots of English yew injured by larvai of Batodes aiigustiorana . 224. Shoot of holly injured by Pd'discd ophthalmi- 206 206 207 207 208 208 209 210 212 212 213 215 215 215 215 216 216 217 217 219 219 220 226 225. "Leading" bud of Scots pine injured by larva of Rctinia turionana . 230 226. ' ' Leading shoot " of young Scots pine injured by larva of Betinia btioli- ana .... 230 227. Ectinia buoliana, male and female . . . 230 228. Gall of resin on Scots pine caused by Uetinia rednella . . .231 229. Betinia r: 279 286. Young form of Psylla cra- bursarius tivrp .... 305 264. Pupal stage of Pemphigus 287. Tip of hawthorn shoot in- 265. bursarius \Yingediormoi Painphigus bur sarins 280 280 288. jured by Psylla cratcgi Twig of alder injured by Psylla alni . 305 306 266. " Corkscrew " gall on leaf- stalk of poplar caused by Pemphi(/us spiro- CICADID.^. thecie . .' . . 281 267. Gall of Pemphigus pallidus 289. Typhlocyba 2dmi 311 on midrib of leaf 281 290. Typhlocyba ulmi 311 268. Galls of Tetraneura ulmi . 282 291. Young form of Typhlocybct 269. Gall of Chermes riridis 286 ulmi .... 312 270. Pkrly stage of gall of Chermes viridis . 286 COCCID/E (Scale-Insects) 271. Winged forms of Chermes (dnetis on spruce leaves 288 292. C( rat a) this lataniiv (fringed 272. ('hermcs larlcis on bark of aphis) on palm . 315 larch .... 290 293. Scale - like secretion of 273. Foliage of larch injured Alturodes on leaf of by attack of Chermes . 291 tomato 316 LIST OF ILLUSTRATIONS. XXlll 294. Female scale of As}ndiotus 295. Male scale of AspidioUiH . 296. Female scale of Diaspis . 297. Male scale of Bias/, is 298. Female scale of Chioua-yns and Mytilaspis 299. Male scale of Chionaspis . 300. Pygidium, or anal seg- ment, oiMytilasjjis pom- orum, showing dorsal and ventral parts, &c. .... 301. Male scales of Chionaspis salicis .... 302. Female scales of Chion- aspis salicis 303. Life-history of Chionaspis salicis .... 304. Scale of Aspidiotus on leaf of Lapageria 305. Scale of Aspidiotus on leaf of Stephanotis 306. Scale of Aulacaspis 7'os(v on dog-rose 307. Female scales of Lecanium capreo', " brown scale," on sycamore 308. Adult female Lecanium after treatment with potash X 20 . 309. Typical antennae of female Lecanmm . 310. Male and female scales of Pidvinaria . 311. Male of Pxdvinaria, species found on currant . 312. Male of Lecanium, species found on sycamore 313. Scales of Physohennes ahietis 314. Depressions on twigs of oak caused by Astero- lecanium variolosum. 3 1 5. Psetidococcus cicensonbark of hornbeam 316. Antenna of Pseudococcus aceris .... 317. Leg of Pseudococcus aceris 318. Rostrum of Pseudococcus aceris .... 319. Cryptococcus fagi on beech .... 320. Apterococcus fraxini 318 318 DIPTERA (Two-wiNOKD Fli ES). 319 319 321. Leaves of Salix viminalis rolled by Cecidomyia 319 mn r(ji)ie.mtorquens 356 319 322. Cecidomyia marr/inem - torqueius (male) . 357 323. Cecidomyia ynart/iiiem- torquens (female) 357 324. Shoot of hawthorn in- 320 jured by Cecidomyia arUa'fji 358 321 325. Twig oif yew injured by Cecidomyia taxi . 358 321 326. Shoots of Salix caprea in- jured by Cecidomyia 322 rosaria 359 327. Cecidomyia rosaria (male) 359 329 328. Section of willow stem showing injuries caused 329 by Cecidomyia salici- perda .... 360 329 329. Portion of bark and wood cut off to show the injuries of Cecidomyia 331 saliciperda . 360 330. Leading shoots of a species of willow injured by 332 Cecidomyia heterobia . 362 331. Twig of Salix caprea in- 332 jured by Cecidomyia salicis .... 363 336 332. Cecidomyia salicis (male) . 363 333. Leaf of lime-tree injured 336 by Cecidomyia tiliam rolens .... 364 339 334. Shoots of lime-tree injured by Cecidomyia tiliam 340 volens .... 364 335. Foliage of ash injured by Diplosis hotulariic 364 343 336. Edges of oak leaves folded by Diplosis dryohia 365 344 337. Galls on leaf -stalks of aspen poplar caused by 345 Diplosis trem^da> . 366 345 338. Galls on upper surface of beech leaves caused by 345 Horniomyia piliyer 367 339. Leaves of goat willow 347 galled by Hormomyia 350 capreie 367 XXIV LIST OF ILLUSTRATIONS. 340. Galls on broom caused by A sjihonthilia sarolhamni 341. A-^/>li(iiiiii//i(i farol/ianvii . 342. Pupa of A yp/iondylia mro- t ham III 343. Gall on stem of bramble caused by Lasiojitera rubi .... 344. Galls ou stems of willow caused by Agromyza i^chineri 345. Leaf of snowberry mined by Chromatomyia oh- scurella 346. Leaf of holly blotched by Chromatomyia ilicis 347. Bibio marci (males and females) 348. Three-year-old spruce in- jured by larvffi of Bibio marci .... 368 369 369 369 370 372 372 374 374 COLLECTING, PREPARATION, AND MOUNTING. 349. Simple arrangement for dissecting specimens under a magnifying- 3o0. Proboscis of honey-bee . 351. Tongue of house-dy 352. Life - liistory stages of " Swallow-tail" Butter- fly ... . 380 385 385 386 APPENDIX. 353. Boring apparatus of the female of Nemahis Erichsonii . . .418 354. Six-year-old larch injured by larvte of Nematufi laricis . . . . 421 FOREST ENTOMOLOGY. INTRODUCTION. It may appear somewliat paradoxical to say that '^' Forest Entomology" differs from the " Entomology of the Forest." Under the latter head- ing the major portion of the science of entomology may be included, inasmuch as the student of general entomology, or the specialist of any particular group, will often find the forest one of his happiest hunting-grounds. This is due not only to the varieties of arboreal food and the varied forest flora, but to the game-preserving laws or customs continually furnishing carrion food, and demanding a varied cover for game-birds and feathered songsters, which doubtless act as sources of dissemination for many species of insects with comparatively poor powers of locomotion. With regard to "Forest Entomology," even in its widest sense we can only embrace directly all insects pre- ferring an arboreal diet, and indirectly all other insects parasitic on arboreal-feeding insects. As the field of natural history becomes broader and broader, natural science tends more and more towards specialisation ; and while it is essential that the young student should gain a compre- hensive view of kindred subjects, the adult who aspires to pose as a public instructor cannot make progress in any branch without limited concentration. Perhaps this narrowing is more essential in entomology than in many other branches of natural history. The science of entomology may be studied from two different aspects — viz., from that of the collector, and of the economic ento- mologist or investigator. The collector, who generally limits his work to one or two groups, A 2 FOREST ENTOMOLOGY. tries by observation and comparison to classify his captures according to the difference in colour, size, and other variations. He also strives to enrich the local fauna by fresh captures. On the other hand, the economic entomologist seizes these data of the collector or systematic biologist, goes a step farther, and considers insects in relation to man, his person and property, either in so far as they affect his industrial products, his cultivated plants, or the wild plants of nature. From a husbandry point of view, the science of economic ento- mology may be divided into three principal parts — viz., garden pests, farm pests, and forest pests. The last is obviously the most difficult to deal with. Before any pest can be satisfactorily checked, it is indispensable to study the life-history of the insect — viz., the egg, the larva, the pupa, and the perfect insect, — and then to deal with that stage in which the most practical results can be obtained. Hitherto it has been customary to study only those insects which, by their numerical strength, have been injurious in the forest, but I venture to think it would be far more interesting and educational, either as an economic or a biological subject, to study all insects which prefer an arboreal diet. To carry this out, two methods of arrangement suggest themselves — viz., a botanical and an ento- mological. With regard to the former, it would be necessary to classify the respective trees, and then arrange those insects Avhich feed on them. This method has been adopted by some American writers. It has, however, this drawback, that while several insects are fastidious as to their diet, others (except for the distinction between hardwoods and conifers) are general feeders. Several German writers follow an entomological method, and this course appears to be far more educational to the student of forest entomology. As the forest is a field literally teeming with varieties of animal life, it may just be as well in passing to remind the practical husband- man and general naturalist that there are numerous creatures in the forest, under rotten bark and so forth, which are not in scientific language insects. Such creatures are snails, centipedes, spiders, and worms of various genera. Hence two questions arise — viz. : (1) What relative position do insects occupy in the animal kingdom? and (2) What is an insect? As regards the position of insects in the animal kingdom, natur- INTRODUCTION. 3 alists differ slightly in their arrangement ; but it may be said that insects constitute a siibdivision of the animal kingdom known as Arthropoda, which are characterised by having jointed limbs, and, as a rule, a distinctly divided body. They rise in an ascending series in the following order — ^viz. : ARTHROPODA (4 classes). 1. Crustacea . . Cmhs, Lobsters, ^-c. 2. Arachnida . . Mites, Scorpions, Spiders. 3. ]\Iyriapoda . . Centipedes and Millepedes. 4. Insecta . . . Insects. In answer to the question, What is an insect % the general definition, with some common exceptions, is that an insect is a creature whose body is divided into three parts, has two antennae, six legs, and, as a rule, passes through four stages of metamorphosis. Accepting this general definition, it may be said that the creatures which, in point of anatomical structure and economic injuries, approach nearest to insects, are mites and spiders ; and as a rough-and-ready classification, it may be said that a mite is a creature whose body is of one piece, a spider two pieces, and an insect three. The mites most injurious to vegetation are the Eriophyidse (Phytoptidae), or four-footed mites. Spiders do not affect trees directly, and therefore need not be considered. We find a certain species of red-spider ^ injurious to the foliage of ivy and other plants, sucking the juices from the leaves, and causing them to assume a sere appearance. The whole class of Insects is divided into seven general or principal orders — viz., Coleoptera (beetles) ; Orthoptera (earwigs, crickets, grasshoppers, and locusts) ; Neuroptera (dragon -flies) ; Hymenoptera (ants, bees, and saw-flies) ; Lepidoptera (butterflies and moths) ; the Hemiptera, including the various species of bugs ; and DiPTERA (two-winged flies). The Hemiptera is subdivided into two principal divisions — viz., Hemiptera-Heteroptera, including the bugs ; and Hemiptera-Homop- tera, including Aphidse or plant-lice, Cicadas, Psyllidse, and Coccidse or scale-insects. 1 The so-called "red-spider" — the dreaded enemy of the gardener — is only a species of mite. FOREST ENTOMOLOGY With the exception of Orthoptera and Neuroptera, all the other orders are represented in forest entomology. Hence the necessity of versatility in entomological knowledge will be apparent. Insects are further arranged into Families, Sub-families or Tribes, Genera, Species, and Varieties. There is, of / course, no sharp distinction between a species "■^j^, , and a variety. In order to understand our subject a little more fully, let us, by way of introduction, briefly consider the life -history and structure T^ii^ of a typical insect. It has just been remarked that, as a rule, insects pass through four stages of metamor- phosis— viz., Bgg, larva, pupa, and perfect insect. When an insect passes through all these stages, the metamorphosis is said to be complete. When, however, it does not, — as, for instance, in the case of Aphidae or plant -lice, — -the metamor- phosis is said to be incomplete. Let us follow a typical ex- ample of a complete metamor- phosis, commencing with the egg stage. The eggs of insects are always deposited in the immediate vicin- ity of the food for the larvas, as Pig. 2 -Eggs of {j^ fi„g_ J ^^^^ 2. In cases of the Lackey Moth ° (Bombyx neu- niany arboreal insects the eggs stria) on hirch "^ °P tirig. Found in are often very difficult to dis- Cheshire. . cover. Sometimes they are con- cealed within the buds, in the midribs of leaves, the roots, bark, or other convenient places, according to the food of the larvse. Insects adopt many devices to protect their eggs against weather and natural enemies, such as covering them over by a gummy secretion, the shedding of hairs, the formation of cocoons, and so forth. The eggs of insects possess very strong powers of vitality, and frost gener- Fig. \.—Egg^ of mtltc Ermine Moth (Arctia menthastri) on grass. (Photo by Flatter.s & Garnett, Ltd., Manchester.) INTRODUCTION. 5 ally has very little effect on them. They vary very much in size, form, and markings. Some are smooth and spherical, others cor- rugated, sculptured, elongated, and tailed. The structure and development, as seen under the microscope, form not only a very interesting study, but present many difficult biological problems. Amongst the higher egg-producing animals no egg will develop without male fertilisation, but amongst insects there are many exceptions to this rule. In numerous cases eggs laid by virgin moths have been known to develop. Amongst saw-flies, certain generations of gall-wasps, plant-lice, and others, we have reproduction by virgin females. This law is known as parthenogenesis, or virgin reproduc- tion without the intervention of a male, and a few special peculiarities will be considered under the respective families. The morphological structure of the egg is somewhat complicated, more especially when we remember the philosophical writings of Weismann and others. The term larva, caterpillar, maggot, or grub is generally applied to insects as they hatch out from the egg, and at this stage they gener- ally differ in form and structure from the perfect insect. On the other hand, the term "nymph" is applied to the creature when it '^ Fig. 3. — Typical forms of la: (From drawings by Mr G. A. Diinlop.) bears a very strong resemblance to the perfect insect, as, for example, in the case of mites and Hemipterous insects. The larva of a moth, on hatching from the egg, is usually a seg- mented body possessing six true legs. The caterpillar of a saw-fly has six true legs, like the larva of a moth, but has also several additional " prolegs," and consequently is easily distinguished. The maggot of a fly is footless. Fig. 3 represents typical larvse. FOREST ENTOMOLOGY. In the case of the Aphididse or greenfly, scale-insects, and plant-bugs, the young in the first stage, whether hatched from eggs or produced alive by the parent insect, resemble the perfect insect very much more than those of beetles, moths, and flies. But whatever shape and form they may take, the young insect in the first stage is always scientific- ally known as a larva. We have in forest insects all forms and classes of larvse, and this variety is of great interest as a biological question. For instance, we find, as arboreal-feeding insects, active six-legged larvae of a greenish colour, and harmonising with the foliage or lichens on the bark ; while on the other hand we find the footless whitish grub feeding within the wood. Thus we have two important factors in the theory of evolution brought before us — viz., the absence and apparent absence of legs through disuse, and the harmony of colour to surroundings as a protection against natural enemies. In the larval stage the insect feeds voraciously, and as a rule grows very fast. The skin of the creature, however, does not grow beyond certain limits, and when this stage is arrived at, the creature halts, as it were, throws ofli" its skin, assumes a fresh coat, often of a different colour, and again commences feeding. This process is known as moulting, and occurs several times during the larval stage until full growth is reached. The duration of larval life in arboreal insects is very varied indeed. In some cases the period occupies only a few days, in others several years. The pupa, or third stage of the life of an insect, is a period of rest or quiescence, when the insect takes no food. It is the period of quiet transformation, when the ugly crawling caterpillar is being transformed into the beauti- ful moth, or the carrion maggot into the swift airy fly. The pupal forms and habits of forest insects are very varied. Thus in beetles all the parts of the future insect are visible, saw-flies and moths form cocoons, and the two-winged flies form cases. Some descend into the ground to pupate, while others form hard cases on the food-plant. Fig. 4 may be considered, for general purposes, as a typical pupa. The stages thus referred to — viz., egg, larva, and pupa — are the Fig. i.— Typical pupal form of a II insect. (From draw- ing by Mr G. A. Dunlop.) INTRODUCTION. periods of growth and development. As soon as bursting from the pupal case takes place, the creature is then literally the perfect insect, inasmuch as there is no further change or development. It is Avell to bear this in mind, as it is a popular notion amongst certain people not conversant with natural history I that small moths and flies are simply "young ones." Such, however, is not the case, for while there is often a difi'erence in size amongst individuals of the same species, it should be remembered that no growth takes place after full pupal emergence. In demonstration of this, perhaps a typical case may be taken — viz., that of the Small Tortoiseshell Butterfly, Vanessa urticce. Fig. 5 represents the pupal case. In fig. 6 we have a representation of the first emergence from the pupal case ; in fig. 7, which is later. i Fig. 5. — Pupal case of Small Tortoiseshell BtMerfly (Vanessa urtica?). (Flatters & Garnett.) Fig. G.— First emergence of Tortoiseshell Butterfy from, pupal case. (Flatters & Garnett.) the wings are slightly more expanded ; and in figs. 8 and 9 we see a representation of the fully developed butterfly. The whole process Fig. 7.— Emergence of Tortoiseshell Butterfly more fully advanced. (Flatters & Garnett.) of development, which is chiefly a question of wing expansion, takes at most only a few hours. FOREST ENTOMOLOGY. In many cases where we have incomplete metamorphosis, the insect is injurious in all its respective stages, as, for instance, in Aphidse and Cicadse. On the other hand, when the metamorphosis is com- plete, we find the insect injurious in larval and perfect stages, or Fully developed Tortolscshell Butterfly. (Flatters & Garnett.) simply injurious in the larval stage alone. The perfect insect (or "imago," as it is termed in entomology) often takes no food what- ever, except sipping water or juices, and the duration of life is often so short as to have become proverbial. As a branch of economic entomology, the student of forest insects should strive to ascertain where and when egg-deposition takes place, how long the insect remains in the egg stage, how long before the first moult and between the other larval moults, when and how long it remains in the pupal stage, and the time and appear- ance of the perfect insect — in short, a full life-history. With regard to the structure of insects, it may be repeated, and must always be borne in mind, that an insect is a creature whose body is divided into three parts — head, thorax, and abdomen. As a biological subject, insect anatomy and physiology is a very broad and interesting Fig. 9. — Fully developed Tortoisesliell Butterfly. (Flatters & Garnett.) INTKODUCTION. 9 one, inasmuch as serial section -cutting and the highest powers of the microscope must be employed. This, though highly inter- esting in itself, is certainly not of vital importance to the young student of forest entomology. It is, however, indispensable that the salient features of head, thorax, and abdomen should in all cases be well studied, and in some special instances be committed to memory. It is upon these points that generic and specific characters depend. The advanced student with a fair amount of leisure would do well to study the internal anatomy of insects. The head is in reality composed of several segments fused together, but looks superficially as if made of a single piece. On the under side vrji.ri. ^^'^4J 'i.v.,iii. (v.,j' n'orh Fig. 10.— Typical mouth parts of insects. (From drawings liy Mr G. A. Dunlop.) it bears the mouth, which is adapted either for biting or sucking. The mouth arrangement separates the whole of the class Insecta into two principal divisions — viz., Mandilndata, or biting insects; and Ha usteJ- lata, or sucking insects. The biting mouth of the beetle and the trunk-like proboscis of the moth are cases in point. Fig. 10 repre- sents typical mouth parts of insects. The eyes in many cases form a prominent part of the head, and are of two kinds — compound and simple. The latter are termed ocelli. In some insects — as, for example, in the house-fly and hive-bee — the compound eyes cover nearly the whole of the head. The orbit of the eye is covered by a transparent skin termed the cornea. An examina- tion under the microscope shows that the surface of the cornea is 10 FOREST ENTOMOLOGY. made up of a large number of six-sided areas, each of which is termed a corneal facet (fig. 11). The ocelli or simple eyes are placed between the compound eyes, or just in the middle of the forehead. The antennjB, or horns, rise from the head. They are composed of several joints, and form good points for the discrimination of species. Any text-book of general entomology, or monograph of some particular group, will show that specific points very largely depend on the antennae. This, for instance, is the case with the Aphididae or plant- lice. The joints of the antennae have to be counted and compared. By a joint is meant not so much the division line between the pieces- Fig. 11. — Compound eye of insect as seen under the inwroscope. (Flatters & Garnett.) as the actual portions themselves if separated at these lines. In counting joints we must begin from the head outwards, making sure the horn is properly and clearly separated' from the head. As a rule, the antenna is stouter at the junction Avith the head, and gradually tapers towards the other extremity : thus we say in a general way, from base to apex. The antennae receive various designations, accord- ing to the structure, as, for example, in fig. 12. The physiological functions of the antennae are not fully understood, but they are sup- posed to be, to some extent, sense-organs. The thorax or middle region bears the legs and wings. It is INTRODUCTION. 11 composed of three distinct segments — viz., prothorax, mesothorax, and metathorax, each of which may be built up of several pieces. These segments vary in proportion in different in- sects, and they also vary in the respective stages of the life of an individual insect. The thorax and its compon- ent parts are of considerable importance in the discrimin- ation of species, as, for in- stance, in the Hymenoptera. The third region is known as the abdomen, and is very variable in form, according to order and genus. It is generally composed of nine segments, and bears the organs of reproduction. The wings, as appendages, are very important points in the discrimination of genera and species In fact, some writers use the wings as a basis of classification. So far as forest entomology is concerned, it is highly essential to make a detailed study of the structure of the wings of saw-flies, gall-flies, Aphididae, and Diptera. The structure of the leg is also of importance in the classification of species. If we separate a leg from the body of the insect and examine it, from the connection with the body to the tip, we shall find it is composed of several parts. The leg is attached to the body by a joint called the coxa ; next to this is a very small portion known as the trochanter, which serves as a sort of joint or hinge connecting the Fig. 12. Tiipical forms of antenna;. (From drawings by Mr G. A. Dunlop.) Coxa. 'frocbaofer. Fcnjur. "Tibia. "tarsus. —Typical form of leg. (From draw- ing by Mr G. A. Dunlop.) 12 FOREST ENTOMOLOGY. femur ; then we have a long joint called the tibia ; and finally, the joints (five or fewer) which make up the tarsus ov foot. Fig. 13 is a representation of the different parts of the leg. The foregoing characters represent the more important points as regards external structure, but it is somewhat more difficult to deal with internal structure and function. To deal with too many details would simply embarrass the ordinary student, and at the same time the subject cannot possibly be intelligible to the student if too few points are presented. The accompanying diagram (fig. 14), representing a longitudinal Fig. 14. — Diagram of longitudinal section of an insect, shoiring principal anatomical points of an insect. (From ' Text-Book of Zoology,' by Dr J. B. V. Boas.) 1-3, first and third pairs of legs cut away; o, anus; c, cerebral ganglion; ch, mesenteron ; e, proctodseum ; g, genital aperture; h, heart; k, crop; m, mouth; «, ventral ganglion; sp, salivary gland; «, uialpighian tubule; a', ovary. section of an insect, shows the relative position of the more important anatomical points from a side view. The nervous system of an insect resembles that of most other animals, inasmuch as the terminal seat of the nerve-centres is the brain, whence proceed other nerve-centres and nerves all over the body. The nerves receive impres- sions from the outside world, which react on the organism, and thus stim- ulate or restrain muscular action. Fig. 15 shows a photograph of a horizontal section through the head of a blow-fly. The various convolutions of the brain should be noted ; and as the development of these convolutions is indicative of intelligence, it is no wonder that certain insects, in this respect, are considered as approaching nearer to man than many other animals. The respiratory organs of an insect are composed of a system of Fig. 15. — Horizontal section through the head of a Mow-fly, showing eyes and brain. (Section and photo by A. Flatters.) INTRODUCTION. 13 tubes, which are well represented in fig. 16. These tubes, or tracheae, ramify throughout the whole of the body ; and the air passes into them by means of special open- ings known as spiracles, Avhich are situated alongside the body of the creature. These spiracles are somewhat complicated open- ings, inasmuch as they open to receive air, shut to maintain a supply, and again open for ex- pulsion. The act of breathing, therefore, is somewhat after the natvire of a bellows, inasmuch as air is received at a special open- ing and driven throughout the tubes. In addition to the func- tion of breathing, it is obvious that the inflating of the tubes is associated with the machinery of flight. Fig. 17 is a photo- graphic representation of a trachea from a butterfly. The organs concerned with feeding and digestion occupy a considerable space in the body- cavity of many insects. From actual specimens this can only be understood by careful dis- section and microscopical man- ipulation. The accompanying figure, however (fig. 18), taken from Miall and Denny, will con- vey a very good idea of these internal organs. In the centre of the figure the food - canal is represented. At s we have the salivary glands and reservoir which contain the saliva, which is not only beneficial in digesting the food within the canal, but in some sucking insects is mixed with the crude Fig. 16. — Diagram of the chief trunks of the tracheal system of an insect; the central nervous system is also shown. a, antennce ; o, eye ; st', anterior stigTiia ; I, longi- tudinal trunk. — After Kolbe. (From ' Text- Book of Zoology,' liy Dr J. E. V. Boas.) 14 FOREST ENTOMOLOGY. sap of the plant before the juice is taken up by the proboscis. At c we have the long crop, and below this we have the gizzard, which organism in some species is a favourite object for the micro- scope. Then we have a stellate arrangement of the csecal tubes or blind tubes ; and proceeding from this there is the stomach, which is simply a tube. At k we have the kidney tubes, and then the small and large intestine, ending in the rectum and anus. In fig. 19 Fig 17.— Trachea of butfcvfly. (Flatters & Garnett.) we have a representation of tlie heart, which is a long muscular tube separated by valves. Having now considered the various structural details of an insect, it may be well to turn briefly to the reproductive organs, the func- tions of which serve to perpetuate the race. Insect reproduction may be either sexual or asexual. In the former, the union of two dissimilar cells is required — viz., a small active (male) sperm cell (spermatozoon) with a large (passive) female cell (ovum) for the production of new individuals. In the latter class, female cells or INTRODUCTION. 15 eggs give rise to new individuals without the intervention of male (parthenogenesis). The reproductive organs of insects may, according to instructions given in certain works on microscopy, be carefully dissected from the abdomen, but the accom- panying figures, together with the explanation, convey a very good idea of the repro- ductive organs of insects. The accompanying figures and explanation have been selected because they are not only large insects, but arboreal in their habits. It may also be noted that, as regards insects, the hive bee has been carefully worked out in this respect, so that reference to any of the more important works on the bee would be found very helpful. Genital organs. — ■ ** The female, as in other Arthro- poda, possesses a pair of ovaries. Each consists of a varying number of tubules (ovarioles), which usually ex- tend like fingers from the anterior end of the oviduct. Each ovariole is surrounded by a thin membrane, and is immature anteriorly, consist- ing of small homogeneous cells ; farther back there are larger cells, young ova, lying in the middle of the tube, and surrounded by smaller cells, which provide them with nutriment, and also secrete the shell (chorion) for the fully developed egg. The mature ova occupy the posterior ends of the ovarioles, and pass thence into the oviduct. "When an kroach. , salivary glands and reservoir ; c, crop (the gizzard below it); ca;, cecal tubes (below them the stomach) ; k, kidney tubes ; i, intestine ; r, rec- tum. Twice natural size. (From 'The Cockroach,' by Miall and Denny.) 16 FOREST ENTOMOLOGY. egg passes into the latter, the corresponding portion of the ovarian tubule shrinks, and thus the egg next in front is brought nearer to the duct. The two oviducts unite to form an unpaired portion, the vagina, which opens ventral to the anus, either freely on the surface or into a cloaca, an invagination occurring at the hinder end of the body. There is usually an evagination of the vagina which serves as a receptaculum seminis, and one or a pair of accessory glands, which secrete either a sticky fluid to attach the ova to foreign bodies, or the mucus sur- rounding them {e.(j., in insects Sometimes there is also an evag- ination of tlie vagina to form the bursa copulatrix, into which the Fig. 19. — Diagnrms cftlie heart of an which lay their eggs in water). ^0.— Female genital organs ofiJie cockchafer. On the right, the ovarioles are lying together in the natural position ; on the left they are separated, and two are cut away, g, vagina ; A-, glands which open into the receptacula ; I, oviduct; o, segments of the ovarioles, containing almost ripe ova; o', regions of the same, containing immature ova ; j), bursa copulatrix, ; r, anterior ; ?•', r", posterior buds of the ovarian tubules ; s, glands ; sg, receptacula ovorum. (After Boas.) penis of the male is inserted in copulation. Not infrequently there is at the female aperture an ovipositor (Locusts), consisting of complicated knife -like or dagger - shaped laminae, or a sting INTRODUCTION. 17 (Hyinenoptera) ; or tlie last abdominal segments, which then are thin and elongate, and may be telescoped, serve in this capacity (Diptera and others). The chorion is often very hard, frequently covered with a delicate and regular sculpturing, and always provided with one or more openings, the micropyles, through which the spermatozoa may enter. The outer form of the eggs varies : it may be spherical, oval, discoid, rough, stalked, &c. " The male genitalia are for the most part a repetition of those of the female. There is a pair of testes, each consisting of several long seminal tubes or shorter seminal pouches, situated at the end of the Fig. 21. — Male genital or