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Westwood (J.0.), 1839-1840, An Introduction to the modern Classification of Insects, 2 volumes (in the separately-paged ‘Synopsis’ (pp. 158) appended to Volume 2, the species specified against the names of the genera there enumerated are to be treated as having there been selected to be the type species of those genera (Opinion 71)) (the dates to be accepted for the various parts of this work are as set out in the table below (Direction 63)) Direction 32
Date of Part No. ‘Introduction’ ‘Synopsis’ Publication (from cover) l | 1-48 > 1-16 [May 1838] 2 1: 49-112 [June 1838] 3 1: 113-160 el=32 [July 1838] A 1: 161-224 {Aug. 1838] 5 1 : 225-288 (Sept. 1838] 6 1 : 289-352 [Oct. 1838] 7 1 : 353-400 : 33-48 (Nov. 1838] 8 1 : 401-462 [Dec. 1838] 9 De l= 128 (Jan. 1839] 10 De 29-192 [Feb. 1839] 11 2: 193-224 [March 1839] 12 20225-=256 {April 1839] 13 2 : 257-288 : 49-80 [June 1839] 14 2)3289-352 [Nov. 1839] 15 2 : 353-400 : 81-96 [Jan. 1840] 16 2
: 401-587 : 97-154 (June 1840]
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THE MODERN CLASSIFICATION
OF
INSECTS;
FOUNDED ON THE NATURAL HABITS AND CORRESPONDING ORGANISATION OF
THE DIFFERENT FAMILIES.
By J.O. WESTWOOD, F.L.S.
HON. MEM. LIT. HIST. SOC. QUEBEC ; MEM. SOC. CHS. NAT. MOSCOW 3; PHYSIOGR. SOC. LUND 3 SOC. ROY. SCIENC. LILLE; SOC. HIST. NAT. MAURITIUS ; SOC. CUVIER. PARIS 35 PLIN. SOC. EDINBURGH ; LIT. PHIL. NAT. HIST. SOC. BELFAST, RICHMOND, SHEFFIELD 3 MEM. SOC. ENTOMOL. DE FRANCE 3 SECRETARY ENT. SOC. LONDON, ETC.
‘© Empirici, formice more, congerunt tantum et utuntur: rationales, aranearwm more, telas ex se conficiunt: apzs vero ratio media est, que materiam ex floribus horti et agri elicit ; sed tamen eam propria facultate vertit et digerit.””— Bacon, Nov. Org. lib. i. aph. 95.
IN TWO VOLUMES.
VOLE
LONDON:
LONGMAN, ORME, BROWN, GREEN, AND LONGMANS, PATERNOSTER-ROW.
1839,
Lonpow:
Printed by A. Srorriswoopr, New-Street-Square.
EXPLANATION OF PLATE.
Order CoLEorTERA. Fug. 1. Cicindela hybrida Zinn. (Linn. Cab.) Fig. 2. Anomala Donovani Steph. (Brit. Mus.) Order NEUROPTERA.
Fig. 3. Boreus hyemalis Latr.
Order HyMENOPTERA. Fig. 4. Dicladocerus ( W.) Westwoodii Steph. Fig. 5. Platymischus ( W.) dilatatus Steph. Order STREPSIPTERA. Fig. 6. Stylops Spencii Pek. Order HEmMiprerRA.
Fig. 7. Aphelocheirus zstivalis Westw.
Order Homoptera.
Fig. 8. Dorthesia characias, ¢ , Latr. Order Diprera.
Fig. 9. Phthiria fulva, 9, Laér.
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ERE A CE.
THE majority of Entomological Works which have appeared, during the last quarter of a century, and which have not only given so great an impulse to the science, but have also imparted to it a philosophical character, of which it was previously des- titute, may be described as exhibiting either generalised views of the subject; or, of elaborate technical details of the genera and species of insects.
Thus, whilst the delightful Introduction to Entomology of Messrs. Kirby and Spence, followed by Burmeister’s Manual, and, at more humble distances, by the Insect Architecture, Trans- formations and Miscellanies, the Grammar of Lintomology, by Newman, and my Lntomologist’s Text-Book, have made us ac- quainted with the general details of insect habits and structure; the Llustrations and Descriptions of the Genera of British Insects of Curtis, the Illustrations of British Entomology of Stephens, the Essay on the Kossorial Hymenoptera of Shuckard, the Lepi- doptera Britannica of Haworth, &c., have led us to the investi- gation of the minute details of generic and specific distinctions. The nature of these works necessarily rendered them essentially different in the information they conveyed; indeed, owing to the greater number of organs possessed by insects over the higher animals, and the consequently great modifications to which they are subjected in the different groups, in order to fit them for performing their various functions, it must be evident that the former class of works, unless extended to a great number of volumes, must necessarily exclude the description
A Y
1V PREFACE.
of genera and species; whilst the immense number of insect species in like manner prevented the latter class of works from entering into detailed accounts of habits and structure, or in- quiries into the relations of the different groups.
Thus the student was led at once from the general views he had gained of the subject, to the minute technical details of genera and species, there being no work which he could take up to serve as a guide to the developement of the principles of modern classification, in the distribution of the orders and families. For years this deficiency has strikingly manifested itself to me, and it is long since I announced my present undertaking, in which I had proposed to myself to show the application of the modern views which have been entertained relative to the na- tural relations of animals in the arrangement of the entire groups of winged insects; illustrating the subject by details of the natural habits, transformations, and structure of the different families.
Nothing can be more distinct than the views entertained by Linneeus and his immediate followers, and modern naturalists, as to the principles of classification. With the former, nothing further was requisite than the construction of an arrangement by which the name of a species might be arrived at in the most convenient, and, consequently, often in the most artificial, manner. ‘Totally regardless of the relations, more or less re- mote, existing amongst the different groups, their writings cannot be regarded otherwise, than as catalogues raisonnées. Modern entomologists, on the other hand, with Latreille at their head, have endeavoured to render the science more in accordance with nature by the establishment of ‘ Familles Naturelles” for the reception of the species most nearly according in habits and structure, the investigation of which, in all the various states of the insects’ existence, has been rendered requisite, in order to trace the limits, or to show the points of connexion existing between the different groups. Convinced that in our endeavours to perfect
PREFACE. Vv
this modern classification of insects, we should deserve the greatest assistance from an acquaintance with their preparatory states, I have, during sixteen years of attentive observation, con- stantly kept that object in view, carefully noting down with pen and pencil every fact which seemed to bear upon the subject. I have studied nature in the woods and fields, tending and observing insects in all their various transformations, well knowing, that the man who confines his researches to the mere collection and examination of museum specimens, can neither possess so intellectual an enjoyment, nor acquire so perfect a knowledge of the subject, as is to be derived from the exami- nation of living nature; and it is both with pleasure and with pride that I now submit the results of my numerous observa- tions to the reader.
Another object, which strongly impressed itself upon my attention, was the necessity for a careful examination and ar- rangement of the facts scattered throughout the voluminous transactions of foreign and native societies, and the various zoological and entomological magazines and other repertoria of science. ‘The time occupied in wading through the hundreds of volumes which this investigation has necessitated, has been im- mense; but the value of the observations otherwise left to slumber unnoticed, will speak for itself. Independent of the materials thus obtained and alluded to in the body of the work, the “ Bibliographical Notices” attached to each family, cannot but be of infinite service to the student.
It is not, however, to my own materials, and those derived from published works alone, that I have had recourse. I have industriously availed myself of every opportunity of studying the collections, as well in great Britain as in France and Ger- many; and I must take the present opportunity of returning my sincere thanks to all my friends who have assisted me in my undertaking, and whose communications I have punctually noticed in the body of my work; but more especially to M.
vi PREFACE.
Victor Audouin, professor of entomology at the Jardin des Plantes, who has kindly permitted me to make unlimited use of his unique collection and voluminous manuscripts relative to the economy of insects in completing my present work.
I have commenced the work with general observations on insects, and then proceeded to divide them into orders: I have afterwards taken up each order separately, dividing it into families, and giving an account of the characters, habits, trans- formations, and general distribution of the insects comprised within each family, with an illustration of their characteristic anatomical details and preparatory states.
It is thus that I have endeavoured to make my work a fitting “Sequel” to the Introduction to Entomology of Messrs. Kirby and Spence, who, upon being made acquainted with its nature, kindly sanctioned my thus styling it. That it may be deemed worthy of such a title, and of a place by the side of their volumes, is the highest object of my ambition.
At the same time, in order that this work may serve as a precursor to the works of Curtis, Stephens, &c., I have added a synopsis of the British genera, brought down to the present time. The idea of the addition of this synopsis was derived from Latreille’s Considérations Générales, in which the genera are shortly characterised, and the names of the typical species given in an Appendix. The additions of generic synonymes, references to generic figures and indications of the number of British species, will render the synopsis more complete, although it must be evident that it can serve but as a guide to more ex- tended research. +
The numerous figures with which the work is illustrated are, in almost every instance, original, and drawn by myself.
I cannot conclude this preface without alluding to the endless
.
PREFACE. vil
gratification to be derived from the study of this branch of natural history, of which the present work bears such ample testimony. For a long series of years, the collection of materials for its completion has been a ceaseless labour of love. Indeed, had it not been thus, it would have been impossible for me to lave proceeded in an undertaking “ of which the profit, if by great chance there should be any, could not be expected to repay, even the cost of books required in it, and from which any fame must necessarily be confined to a very limited circle*,” and in which the time absorbed in the preparation of the text, and in microscopically dissecting so many minute objects re- presented in the wood-cuts, has been so great. I have, however, persevered, and, if I shall have succeeded in inducing any of my readers to pursue the science with a higher aim than that of collecting specimens, by investigating the habits and the corre- sponding organisation of these animals with a view to the dis- covery of their natural relations and classification, my labours will not have been in vain.
* Kirby and Spence, Introd, pref. vol. i, p. 10.
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CONTENTS
OF
THE FIRST VOLUME.
Page Page
Observations upon Insects in Hydradephagous Adephaga - 93 general - - ae * Dyticiwx - =o General Structure of Insects - 5 GyYRINIDE - 105 Head, &c. - = 16 Philhydridous Rypophaga - 111 Mouth - - Sh OS) HETEROCERIDA = le Thorax, &c. - =5 LO ParNiID& - = iss Abdomen - - - I4 HELOPHORIDE - 120 Internal Systems - - 15 HypDROPHILID® - 122 Transformations =e = 06 SPH ERIDIIDE - 128 Distribution of Insects into AGATHIDIDE - 150 Orders = - - 18 Necrophagous Rypophaga - 132 Metamorphotic System - 19 SCAPHIDIIDE - 134 Alary System - - 20 SILPHIDE - Br lists Cibarian System - = 21 NITIDULIDE - - 140 Eclectic System - - 23 ENGID& - - 143 Representative System - 26 Trogositides - 145 COLEOPTERA. Cucujides - - 148 Bibliographical References 30 Paussip& - - 150 Characters of Order - 33 MYCETOPHAGIDE - 152 Arrangement of Order - 37 DERMESTID& - 155 Pentamerous Coleoptera - 44 Brachelytrous Rypophaga - 161 Geodephagous Adephaga 45 Sv APHYLINIDE - 162 CICINDELID& 47 Pselaphides - 174 CaRABIDE - 57 | Clavicornes - - 178
Larvee of Carabidee - 65 Byrruipx% . - 178 Arrangement of Cara- HisTEeRID& - 181
bidze - - 71 Lamellicornes - - 184
Brachinides ara LUCANIDE- - - 185
Scaritides = - 81 Petalocera - - 191 Harpalides - 83 GEOTRUPIDE - - 201 Carabides - - 89 SCARABEIDE- - - 203 Bembidiides = Ml APHODIIDE - 207
VOL. I. a
TROGIDE - DyYNASTIDE RUTELID - ANOPLOGNATHIDE MELOLONTHIDE GLAPHYRIDE CETONUDE - Serricornes - Buprestip& - EUCcNEMIDE - ELATERIDE - CEBRIONIDE - CYPHONIDE - LAaMPyYRID& - TELEPHORIDE MELYRIDE - CLERIDE - Prinip& - LYMEXYLONIDE BostrRICHIDE ScyDMENIDE Heteromerous Coleoptera Trachelia - Noroxipz - PYROCHROID#£ LaGriip® - Horupe - MorDELLIDE- CaNTHARIDE SALPINGIDE - CGEDEMERIDE. - MELANDRYID£ Atrachelia . CISTELID.E -
CONTENTS.
Page 208 210 213
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Rhyncophora BrucHIp& ATTELABIDE
CURCULIONID&
ScOLYTIDE Longicornes - PRIONIDE CERAMBYCID& LEPTURID Phytophaga - CRIOCERID CassIDIDE
GALERUCIDE -
CHRYSOMELID®
Pseudotrimerous Coleoptera
EROTYLIDE
ENDOMYCHIDE
CoccINELLID
EUPLEXOPTERA (For-
FICULIDE) ORTHOPTERA BuatTrip#&
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ERRATA AND ADDENDA TO VOL. I.
2
Page 1. line 8. for ‘and antenne ” read “and two antenne.” line 11. after “ Aristotle” read “and the class of insects of Lamarck.” 3. line 5. for “mites” read “lice.”
In reference to the observations in sect. 1. relative to the importance of metamorphosis as the leading character of the Ptilota, Mr. MacLeay, in consequence of the observations of Mr. J. V. Thompson, and the confirmation assumed to have been afforded to them by the recent researches of Captain Ducane (Annals Nat. Hist. No. 9. Nov. 1838.) has suggested the theory, that “the Ptilota may be characterised by their change of form taking place during their last two or three stages of ecdysis ; while the metamorphosis of all other annulosa only oceurs during the first or second moult after leaving the egg.” ( Illustr. Zool. So. Africa, No. 3. p. 53.) In opposition to this theory, I will only here reply that the most elaborate examinations hitherto made of the eggs of spiders and crustacea by Heroldt and Rathke, together with
my own of the egg of one of West Indian Land Crabs, have clearly demonstrated that, at least in the species under examination, not the slightest change worthy of the name of metamorphosis takes place. 11. line 16. for “ininsects” read ‘in those insects.” line 18. for “being” read “are.” line 20. dele.
line 30. for “are as destitute” read “ which are as destitute.’
6. line 3. for “oxyginating ” read “ oxygenating.” 7. line 10. 29. for “ Homomorphous” read “ Monomorphous.”
line 19. for “larve” read “ larva.”
] 1 21. line 21. dele “‘and” before Dermaptera. 32. line 44. for “ Harris in ditto?” read “ Harris in Nat. Hist. Soc. Hartford, vol. i.” 37. line 37. Mr. Haliday states in the Zoological Journal, that Clambus pos- sesses five joints in the tarsi. 42. line 40. for “ Philyhdrida” read “ Philhydrida.” 83. line 14. Mr. Newman assures me that Eutoma tinctilatus possess a max- illary unguis, which escaped my microscopical examination of it, If this be the case, it cannot enter into the genus Carenum, in which Latreille noticed the want of a maxillary hook. (R. An. second edition, tom. iv. p. 381.) 89. line 30. for “ Cbhychrus” read “ Cychrus.” 93: The elaborate volume of descriptions of Dyticide and Gyrinide recently published by M. Aube, must be particularly noticed. 118. line 9. for “to be the” read ‘to be those of the.”’ 123. note *, line 2. for “ species” read “ spaces.” 126. note +, M. Audouin has published a note upon this subject in the Hist. Nat. Ins. tom. vy. p. 268.
xil ERRATA AND ADDENDA.
Page 151. Mr. MacLeay states that his brother has observed that an Austra- lian species of Cerapterus resides in ants’ nests, and likewise possesses the power of crepitating. ‘ 152. note *, for “ Mycelophagide ” read ‘“‘ Mycetophagide.” 176. note *, Within a month from the time of the publication of this note (namely, on the 30th of August, 1838), I had the pleasure to capture the Claviger foveolatus in an ant’s nest in Whychwood Forest, Oxfordshire. 183. note t+, read “ Faun. Boreal. Americana.” 290. line 30. and $1. dele the statement of this cocoon being that of a Lepi- dopterous insect. The observation of Mr. MacLeay having applied to the supposed gall of Cecidoses figured by Mr. Curtis on the same plate, and which Mr. MacLeay informs me is the cocoon of a Lepidopterous insect, and nota gall. 221. The family Cetoniide has formed the subject of an Essay by Mr. MacLeay in the Jilustrations of the Zoology of Southern Africa.
231. line 34. This passage was written (but not published) previous to the pub- lication of the second edition of Stephens’s Nomenclature, in which the genera of Buprestide were introduced.
281. line 23. for “ palpi to appear to belong, this” read “ palpi, appear to belong
to this.”
304. Fig. 35.1. The antenne ought to have been inserted at the base of the
rostrum.
,
365. line 21. for “naturally” read ‘ materially.”
MODERN CLASSIFICATION
OF
INSECTS.
I. OBSERVATIONS UPON INSECTS IN GENERAL.
Insects, as proposed to be treated of in the following pages, may be defined to be, Annulose Animals breathing by trachez, having the head distinct, and provided in the adult state with six articulated legs and - _antenne, subject also to a series of moultings previously to attaining perfection, whereby wings are ordinarily developed.
This definition, which comprises the characters _. of Be al or winged insects, of Aristotle, does not, indeed: accord with that required by the group of insects as extended by many recent authors ; but the group thus defined is regar ded Mr. Macleay (Linn. Trans. vol. xiv. p. 67.) as pre-eminently natural, and as constituting the typical division of Annulose Animals; and, indeed, when we call to mind the general characters of the classes of the Annulosa and other invertebrated animals, it will appear evident, that the extraordinary metamorphoses to which the Ptilota are subject, attended as they are by the ulterior development of organs of flight, which exist in no other group of invertebrates, ought to be regarded as especially entitling these insects to be treated not only as a distinct, but as a most natural, group*, and consequently as giving a superior degree
* It will, perhaps, be objected by some persons, that the existence of metamor- phoses, as an exclusive character, is denied by the researches of Mr. Thompson, who has asserted that the Crustacea undergo equally striking transformations. I have, however, and I trust satisfactorily, disproved the statements of this author, in a Me- moir published in the Philosophical Transactions for 1835. It may also be said, that the character employed by Mr. Newman to isolate the winged insects (that of their incapability to reproduce their limbs) ought to have been employed. But this character has also been long since refuted by Dr. Heineken in the Zoological Journal, and by Dr. Burmeister in his Handbuch der Entomologie
B
g MODERN CLASSIFICATION OF INSECTS.
of weight to that system in which they are regarded as distinct, over the numerous other methods which have hitherto been proposed from time to time, of some of the more celebrated of which it will be convenient to give a very short notice. :
Linneus and his disciple Fabricius introduced into the great division of Insects, not only those subject to metamorphoses and the acquisition of organs of flight, but also all other articulated animals possessing articulated legs, which formed in the Linnzan system the order Aptera, but which Fabricius separated into several other classes. In the former of these systems we perceive the great disadvantages attendant upon the employment of a single character in our attempts to arrange zoological objects, since the order Aptera regarded as equivalent in rank with the orders Coleoptera, Lepidop- tera, &c., comprised not only apterous hexapod, and polypod insects, but also Crabs and Spiders, which more recent anatomical investi- gations have proved to be respectively groups of equal rank with all the winged orders united together. The elaborate researches of Cuvier in comparative anatomy fully proved, that the Crabs and other allied animals (genera— Cancer, Oniscus, and Monoculus, Zinn.) could not be retained amongst insects, inasmuch as they possessed a totally distinct system of respiration, breathing by means of bronchiz or gills, as well as a complete system of circulation: of these, therefore, he formed the class Crustacea, which has been regarded as distinct by all subsequent entomologists. Lamarck on similar grounds removed the spiders, and some other species of Apterous insects (genera — Aranea, Scorpio, and Phalangium Zinn.), constituting them into the class Arachnida, including therein, however, the Mites (G. Acarus, Linn.), which breathe by means of tracheze; tie Centipedes (G. Scolopendra and Iulus Linn.), the spring-tailed insects (G. Lepisma, and Podura Linn.); and the Lice (G. Pediculus Linn.*) These four last-mentioned groups have much _ perplexed systematists, none of whom are agreed as to their location, and by several of whom their situations have been frequently altered. By Dr. Leach+ the Mites were raised to the rank of a distinct class, under the name of Acari; the Centipedes were also elevated to the rank of a class under the name of Myriapoda Latreille ; whilst the spring-
* The two remaining Linnzan genera are, Pulex (the flea), subject to metamor- phosis ; and Termes, which belongs to the order Neuroptera. t+ In Samouelle’s Comp. p. 75.
INSECTS IN GENERAL. 3
tailed insects (as an order named Thysanura from Latreille), _ and the lice (as an order named Anoplura or the Parasita Zatr.) were united with insects as a subclass under the name of Ametabolia.* Mr. Curtis has adopted the group of insects as thus pence by Dr.
Leach. By Latreille the spring-tailed insects and nites have been constantly regarded as belonging to the class of insects, from their possessing six legs, in common with the metamorphotic insects; and the mites have been constantly regarded as portion of the class Arachnida, from possessing in common with the spiders eight legs. The Centipedes, however, have been variously regarded by Latreille, either as forming a distinct order of insects, under the name of Myriapoda (as in the 2d edition of the Regne Animal), or as con- stituting of themselves a distinct fourth Annulose class (as in his last work, Cours d’ Entomologie, 1831.)
Messrs. Kirby and Spence have adopted a classification totally dif- ferent from that of all preceding or subsequent systematists, in which three Annulose classes are formed, namely, Crustacea, Arachnida, and Insecta; but the latter is augmented by the hexapod spring- tailed insects and lice, the octopod mites, and the polypod centipedes, all of which are regarded as forming a single order, Aptera, charac- terised by respiring by trachez, and having no system of circulation. (Introd. iii. p. 22.) The incongruous character of the groups thus associated together ; the possession of pulmonary sacs by Thelypho- nus, which is thus placed amongst the mites; the recent researches of M. Dugés on the respiratory organs of Dysdera and Segestria, read before the Academie des Sciences on the 9th February, 1835, (clearly demonstrating that the respiratory system in the Arachnida is not entitled to pre-eminence as a character of the class); and, lastly, the admission of Messrs. Kirby and Spence themselves (Jntrod. iv. 383.), that their order Aptera is not a natural, but merely a pro- visional one, and that the hexapod insects are to be regarded as more peculiarly entitled to the denomination of Insects (Jntrod. iii. 22.), will, I trust, be considered as suflicient to authorise me in not adopt- ing their views.
Mr. Stephens, in addition to the ordinary metamorphotic groups, has added the lice (Anoplura) to the class of insects, without, how- ever, offering an argument for this deviation from all previous systems. The general construction of the hexapod antenniferous body of the
* Edinb. Encyel. vol. ix. B 2
4 MODERN CLASSIFICATION OF INSECTS.
lice approaches very closely to that of such of the very few true insects, which undergo an imperfect metamorphosis, without their wings being developed; as the bed bug, certain apterous Orthoptera, &e.
Dr. Hermann Burmeister has, however, introduced both the spring- tailed insects and the lice amongst insects; placing the suctorial lice (Pediculidz ) in the order Hemiptera, and the mandibulated lice (Nir- midz) and the spring-tailed insects in an order with Libellula, Ephe- mera, Termes, and Psocus. (Comp. his Handb. der Ent. vol.i. § 352., and vol. ii. p. 39., and De Insect. Syst. Nat. 1829.) He appears to have been led to adopt this arrangement by giving too slight a weight to the developement of organs of flight; citing, in support of his views, the occasional apterous conditions of certain true insects, and even the occasional absence of wings in one of the sexes, as in the glow-worm. In these instances, however, I can see but exceptions to a general rule, for which allowances ought to be made, and, conse- quently, as not warranting the introduction of entire groups of Apte- rous animals into the class.
It only remains for me to mention the system of Mr. MacLeay ; which, from the philosophical manner in which the subject has been treated, merits particular notice. In this system, the metamorphotic Annulosa are regarded as the true insects; the classes Crustacea and Arachnida are introduced, with the limits given to them by Latreille ; but the four other groups, which have already occupied so much of our attention, are, together with another group of Annulose animals, formed into a separate class, under the name of Ametabola, “ having no metamorphosis in the usual sense of the word, or only that kind of it, the tendency of which is confined to an increase in the number of feet.” (Linn. Tr. xiv. p.66.) All these Ametabola are destitute of wings, but their respiratory system is similar to that of the true insects. It would occupy too much space were I to recapitulate the arguments adduced by Mr. MacLeay, as to the propriety of the establishment of this class ; I must, therefore, refer the student to the fifth and sixth chapters of the second volume of the Hore Entomologice, premising only that I have adopted this classification, because it leaves the true winged metamorphotic insects as distinct from the other groups, and without expressing any opinion upon the Quinarian views of Mr. MacLeay, or upon the introduction of the Vermes amongst the Ame- tabola.
STRUCTURE OF INSECTS. 5
Having thus determined the limits of the class of Insects, a slight sketch of their structural characters, and their distribution into orders, will next occupy our attention.
II. GENERAL STRUCTURE OF INSECTS.
The word Jnsect, derived from the Latin (iz and seco, sectum), indicates one of the chief characters of this group of animals; the body in which is not only composed of a continuous series of seg- ments, articulating with each other, but is also often divided or cut into three very marked portions, to which the names, head (caput), thorax, and abdomen have been applied. Unlike the higher animals, insects are not internally furnished with a bony skeleton ; but, to supply its want, the external envelope is ordinarily of a corneous-like texture, to which, on its inner surface, the muscles are attached, and which encases the viscera and other internal organs.
This external Envelope is, therefore, the analogue of the skeleton of the mammalia, the external organs of locomotion being attached to it-- In its chemical composition, this envelope is peculiar, consisting of a substance which has been named Chitine, which is found only in the teguments of articulated animals. To this is added a certain quantity of albumen, an animal matter, certain salts, and an oil of variable colour, which last is generally disposed near the external sur- face, and supplies the varying colours so much admired in this tribe of animals.
The Number of Segments, whereof the body of insects is composed, appears, at first sight, to be very variable, but this is more apparent than real; being caused either by the soldering together of certain of these segments, or the rudimental state of others, in consequence of the superior developement of the adjacent segments. It is, however, to be observed that, owing to the non-developement of wings and generative organs in the larva, the body, in this state, is more regu- larly segmented, and the segments more equally sized, than in the imago state. It has been supposed that the typical number of segments is thirteen ; but some late observations upon the larva of Hymenop- terous insects, and upon the common earwig ( Zrans. Ent. Soc. vol. i. p- 157.), have led me to consider that the number is greater. From the elaborate researches of Audouin and MacLeay into the structure of the thorax of hexapod insects, it has been considered that each of
B 3
6 MODERN CLASSIFICATION GF INSECTS-
the primary segments is typically composed of four other segments ; but as it is only in the thorax that this complex structure is to be traced, I shall notice it in describing that part of the body.
The Distribution of the Segments of the body of insects into three distinct regions (head, thorax, and abdomen), although not observable in the early states of existence of many species (ex. gr. larvae of Lepidoptera), wherein the body is composed of a series of nearly equal-sized continuous rings, is so essentially indicated in the last and highest state of perfection of the animal, that we may, with propriety, adopt the Linnzean division of the body, into head, thorax (¢runcus a) and abdomen; but the limbs (artus), being attached to the segments forming the thorax, are to be regarded as appendages thereto, and not as distinct primary parts of the body, in the same manner as the antenne and trophi are appendages of the head ; and the ovipositor, sting, caudal forceps, &c., appendages of the abdomen. These three primary divisions of the body appear to be adapted for distinct ob- jects of equally great importance in the physiological relations of the animal. The head comprises the organs of sense, and consists of a single segment. The thorax comprises the organs of locomotion, and is composed of the three following segments; and the abdomen, which contains the generative organs, comprises the remaining segments.
The Head is a generally corneous, and often somewhat globose, skull; having an opening in its anterior part for the reception of the organs of the mouth (¢rophz), and a similar opening at its posterior part, where it is attached to the following segment, and through which the cesophagus or other anterior part of the digestive canal passes. On each side of the head is fixed an immoveable eye, of large size and complex structure, between which are occasionally two, or oftener three, minute simple lenses (ocelli). Towards the front of the head, but behind the mouth, are also attached two move- able organs, of endless variety of form, size, and structure, which are
_termed antennz, and of which the precise uses have not been deter- mined. The front margin of the head is often separated by a suture from the remainder of the skull, and is then termed the Clypeus. The under surface of the head is the Jugulum K. (Gula, Strauss, and Burm. ), the swollen anterior margin of which is the “piece prébasilaire” of Strauss and Burmeister (Manual, pl. iii. f. 12. d, d), being the stipes of MacLeay, and the mentum and insertion of Newman. It is to the front margin of this swollen part that the true mentum is attached.
STRUCTURE OF INSECTS. 7
The form of the head and of its various organs is exceedingly varied ; the variations of the mouth are of the highest importance, serving for the primary divisions into orders.
The Antenne are generally elongated, and consist of a number of rings attached together, but varying in structure, not only in almost every species, but also in the sexes of the same species; those of the males being often much more complex than those of the females. These organs, from their existence only in insects and cther articu- lated animals, as well as from their diversity of structure, have at- tracted much attention, and have been employed as excellent distinc- tive characters of genera, &c. They have received the names of filiform (thread-like), setaceous (bristle-like), moniliform (necklace- shaped), cylindrical, prismatic, ensiform (sword-like ), fusiform (spindle- shaped), aristate (terminated by a hair), dentate (toothed), serrated (saw-like), pectinated (comb-shaped), flabellated (fan-like), ramose (branched), furcate (fork-like), geniculated (elbowed), perfoliated (with a knob, composed of loosely attached joints), lamellate (with a plate-like knob), plumose (feathered), verticillate (with whorls of hair), &c., according to their various form and clothing.
The Composite Eyes are exceedingly varied in their size; sometimes, as in the male hive bee, nearly occupying the whole surface of the head: they are composed of a number of minute transparent corneous hexagonal facets, representing the cornea, and being more or less gibbous, according to the carnivorous habits of the species. The number of these facets is equally varied; thus in the ant there are but fifty, whereas in the eyes of the butterfly, according to Geoffroy, there are not fewer than 34,650. From the elaborate anatomical re- searches of Muller and others, it is evident that each of these facets operates asa distinct organ of vision; and from an interesting me- moir lately presented by Mr. Ashton to the Entomological Society, it appears, that in some species (Libellula) the upper facets are of a larger size than the inferior ; whence a distinct kind of vision may be supposed to be possessed by these different sized facets, according with the habits of the insect. A similar observation was made by Hooke in Tabanus.
The Simple Eyes (ocelli)are generally three in number, and arranged in a triangle on the crown of the head; they are of a simple and semiglobular form. The eyes of larva, spiders, and some other Annulosa are simple ocelli, arranged in groups.
B 4
8 MODERN CLASSIFICATION OF INSECTS.
The Organs of the Mouth (upon the variations in which Fabricius constructed his system) are, notwithstanding al) their variety of form, reducible to one type of structure. They consist of six principal - organs, of which four are lateral and disposed in pairs: the two others are opposed in the opposite direction : thus, :.:, filling up the space left by the two other pairs above and beneath. The upper single piece is the upper lip (labrum): the upper lateral pair of organs are the mandibles or upper jaws; the lower lateral pieces are the max- illa, or lower jaws, and the under single piece is the under lip. The three lower organs are furnished with articulated appendages (palp/). The first principal variation in the structure of the mouth originates in the mode of action of the various organs. Thus, when the lateral pieces are short, inserted at a distance apart, and have a horizontal motion, the action is that of biting; when, on the other hand, the lateral pieces are elongated, originating near together, and having a longitudinal motion (by means of strong and elongated muscles at their base), the action is. that of sucking; the ascension of fluids in the latter case being produced by the gradual approximation of the pieces of the mouth (Latr. Cours. d Entomol. p. 206.), which thus form a siphon or haustellum. The insects having the former action are termed Broyeurs by the French, and Mandibulata by the English, and the latter Suceurs, or Haustellata; but as these,terms are liable to ob- jection (the insects composing the latter group possessing mandibles, although in an altered form), it would be convenient to employ some other names. The terms Dacnostomata and Antliostomata express the properties of the two groups, without involving the contradiction suggested by the terms ordinarily in use.
In the Biting Insects, the upper lip is generally a simple and flat- tened plate closing the mouth above; the mandibles are generally horny, and more or less toothed (the teeth being, however, portions of the jaw itself), serving for gnawing in pieces the particles of food; the maxillz are more complicated, being furnished at the side with an articulated appendage like a short antenna, but of which the number of joints is never more than six. The maxilla itself is termi- nated by two lobes, the exterior of which is sometimes articulated, representing an internal palpus as in Carabus, sometimes formed into a helmet-like appendage ( G'alea), protecting the inner lobe as in Locusta, and sometimes soldered to the inner lobe. The lower lip is still more complicated: its base is a horny plate (mentum, or the labium of
STRUCTURE OF INSECTS. 9
Fabricius): this is followed by a generally membranous organ (labiwm or the ligula of Fabricius), near to the external base of which is attached a pair of short articulated palpi; within this terminal la- bium is to be seen a fleshy organ, often forming its internal coating, but which in the dragon-fly and the grasshopper is a separate piece (lingua), and of which the lateral anterior angles (paraglosse) are oc- casionally prominent. Such is the general character of the mouth of the Coleoptera, Orthoptera, Neuroptera, and Hymenoptera. In the bees, however, the three inferior organs of the mouth are elongated, so as when in action conjointly to form a sucking apparatus; the mandibles, however, retain their ordinary form. Latreille, notwith- standing, terms this form of mouth Promuscis, a term long previously given by Kirby and Spence to the mouth of the Hemiptera. Amongst the Swetorial Insects, there is a much greater diversity of structure in the mouth. In the Hemiptera and Homoptera, the upper organ is very short and pointed; the four lateral pieces are elongated and transformed into slender lancet-like organs (the max- illary palpi being obsolete), enclosed within the equally elongated fleshy and articulated lower lip, which is turned upwards at the sides, forming a canal, up which the fluid food of the insect ascends; the labial palpi are also here obsolete. This variation of the mouth is termed by Kirby and Spence, promuscis; but by Fabricius, Olivier, and Latreille, rostrum. In the Diptera, the five upper organs, to- gether also with the internal tongue, are elongated into lancet-like organs, the maxillary palpi being attached at the base of the maxillz. These six organs are inclosed in a fleshy thickened piece (which is the lower lip), terminated by two large fleshy lobes which act as suckers. In many species, however, some of these lancet-like organs are obsolete. This kind of mouth is termed by Kirby and Spence, Linnzus and Fabricius, a proboscis. In the Lepidoptera, the three upper organs are almost obsolete, but the maxilla are greatly elongated into a delicate instrument, which when at rest is spirally folded up and hidden from sight, but when in action, is extended and thrust to the bottoms of flowers. At the base of these maxilla, a pair of minute palpi is often to be found. The lower lip is soldered to the head, but it is furnished with a pair of large palpi clothed with scales, which serves for the defence of the spiral maxilla. By Kirby and Spence, this kind of mouth is termed antlia; by Fabricius, lingua; and by Latreille, spirignatha (or more properly speirignatha). In the Apha-
10 MODERN CLASSIFICATION OF INSECTS.
niptera, the organs are all exposed, the upper lip, mandibles, and tongue elongated, the maxilla and labium short and furnished with articulated palpi. It is called rostrulum by Kirby and Spence, and rostellum by Latreille, which latter name had been previously em- ployed by Kirby and Spence for the mouth of the Pediculus, but which Latreille has termed Siphunculus.
The Thorax, on account of its being the chief seat of the various organs of motion, is extremely complicated and variable in its struc- ture; and it is only within a few years that its investigation has been philosophically entered upon, or a concise nomenclature of its parts, founded upon such investigations, proposed. It is the truncus of Linnzus, and comprises the three segments following the head, which have been respectively termed prothorax, mesothorax, and meta- thorax, which were originally proposed by Nitzsch. The first of these segments bears the anterior pair of legs, the second supports the middle pair of legs and the anterior pair of the organs of flight, and to the third are attached the posterior pair of legs and the pos- terior pair of wings. A binary division of the thoracic segments has been proposed, founded upon the nature of the organs of motion; thus the anterior of the three segments is the manitrunk of Kirby, the collum of Knoch, and the corselet of Strauss; whilst the two pos- terior wing-bearing segments are united into the “segment alifére” of Chabrier, the alitrunk of Kirby, the pectus of Knoch, and the thorax of Strauss. The composition of each of the thoracic segments (and indeed of every segment according to the views of Audouin) is essentially similar, consisting of four dorsal subsegments, namely, the Preescutum, Scutum, Scutellum, and Postscutellum; and the Paraptera, Sternum, Episterna, and Epimera, which are lateral or ventral pieces. In the Hymenoptera, Diptera, and Lepidoptera, the prothorax is almost evanescent or reduced to a simple collar; but in the Coleo- ptera, the pronotum (its upper surface) is the very large piece suc- ceeding the head, and which has from the days of Linnzus been ordinarily but incorrectly termed the thorax, and its subsegments are entirely confluent, this segment being destitute of wings; indeed, it is only in the prothorax of some Locusts that the dorsal subsegments of the prothorax can be traced. But in the Mesonotum and Meta- notum (or dorsal parts of the meso- and metathorax) the subseg- ments are much more conspicuous, indeed the scutellum of the mesothorax was noticed by Linnzeus as one of the chief component
STRUCTURE OF INSECTS. 1 F
parts of the trunk or thorax, being the small triangular piece which is seen at the base of the suture of the elytra of beetles. The chief ventral or pectoral piece in each of the three thoracic segments is the Sternum, which varies considerably in its size and form: thus in the Elateride the prosternum is elongated into a point extending between the middle legs, whilst in the large Hydrous piceus the Mesosternum and Metasternum are soldered together between the middle legs, the former projecting in a point between the anterior legs, and the latter extending far beyond the base of the hind legs. The modifications of form in the various thoracic segments result from the complicated machinery requisite for the due performance of the two chief kinds of insect locomotion, namely, leg-movements of various kinds, and wing-movements ; but more especially from the great diversity of the wings and the occasional transfer of wing-motion to a single pair of wings, this pair being either the anterior, as in Diptera, or the pos- terior, as in Coleoptera: thus, whilst in insects which have the four wings of nearly equal size, the two aliferous segments of the thorax being also of nearly equal size, and their subsegments similarly developed (ex. gr. Hemerobius) ; those tribes of insects which haye the segment to which such pair of wings is attached consequently increased in size: thus in Diptera the mesothorax nearly occupies the entire thorax, there being only mesothoracic organs of flight. In the Hymenoptera, the posterior wings exist, but of a small size; the me- tathorax is therefore much larger than in the Diptera, but much smaller than the mesothorax. In like manner, in those orders which have the posterior pair of wings enlarged, the mesothorax is di- minished, and the metathorax equally enlarged: this is especially the case in the Beetles, but still more strikingly so in the Strepsiptera,
“are as destitute of fore wings as the Diptera are of hind ones.
* The Wings, or organs of aerial progression, (upon which the Linnzan arrangement is principally founded, ) bear no resemblance to those of birds, being, on the contrary, more analogous to the wings of bats, consisting ordinarily of a delicate double membranous plate, traversed by more or less numerous tubes, which Dr. Leach regarded as repre- senting a system of bony air vessels (Pterygostea), but which the recent microscopical observations of Mr. Bowerbank and others have proved to be veins. These organs of flight, with respect to their consistence, are termed elytra, hemelytra, tegmina, membra-
12 MODERN CLASSIFICATION OF INSECTS.
naceous wings, halteres or pseudo-halteres. In the Coleoptera the upper pair of wings is transformed into ‘a pair of corneous cases (elytra), generally extending to the extremity of the body, and, when unemployed, shutting closely together by a straight suture down the back. Thus united, they become a shield of great strength, defending the posterior pair of wings, which, when at rest, are transversely folded up beneath them, and which, when the insect is on the wing, are its only effective organs of flight. This lower pair of wings is membranous, and traversed by various corneous veins. In the orders Orthoptera and Homoptera the anterior wings (teg- mina), when at rest, form a roof-like shield to the posterior wings, although much less effective than that of the beetles, since they are of a much more coriaceous texture. The posterior wings themselves are membranous, of a large size, and, when at rest, they are longitu- dinally folded. During flight, both pairs assist in locomotion. In the Heteroptera, the anterior wings (hemelytra) are coriaceous through- out their basal half, and membranous throughout their apical portion ; the posterior wings are of moderate size, membranous, folded lon- gitudinally at rest, and defended by the hemelytra, which shut hori- zontally, the membranous portion of the one folding upon the same part in the other. In the Neuroptera the wings are generally of equal size: they are of a membranous texture, and the posterior are seldom defended when at rest by the anterior, and they are consequently not folded up: whilst in the Hymenoptera and Lepidoptera the ante- rior wings are larger than the posterior, which are likewise never folded up. We thus see, that where the posterior wings take a large share in the act of flight, it is necessary that, being of a larger size, they should be folded up and defended by more or less powerful wing-cases while unemployed; when, on the other hand, the anterior wings take the largest share of action during flight, the poste- rior are so much reduced as to require no defence; in other words, where we find large and strong wing-cases or shields, the active organs of flight are membranous, large, and folded up. The veins, which I have already noticed, are extremely variable in their number: thus, whilst the wings of the Dragon-fly are covered with cells or meshes formed by the conjunction of these veins, the wings of some Hymeno- ptera are almost destitute of them. The numberand position of these veins and cells are of great use in determining the genera, especially of Hymenopterous and Dipterous insects. The wings of
STRUCTURE OF INSECTS. 13
the Lepidoptera are clothed with exceedingly minute feather-scales, which are rubbed off at the slightest touch, and which are somewhat analogous tothe scales of fishes. In the Trichoptera and some Diptera the wings are more or less clothed with hairs. In the Diptera there is a pair of slender clubbed organs attached at the sides of the metathorax termed halteres, and which have been considered as representing the hind wings, whilst in the Strepsiptera a somewhat similar pair of twisted organs are placed at the side of the thorax, in front of the wings : these have been termed pseudo-halteres, and are the analogues of the fore-wings. In addition to the foregoing appendages, the pro- thorax is occasionally armed with a pair of moveable spines (a«mbones K. and S.), and in the Lepidoptera there is a pair of scales (patagia K. and S.) clothed with hair, distinct from the tegula, with which they are confounded by Burmeister (Manual, p.77.). The latter organs acquire a large size in the Lepidoptera, and are the pieces clothed with hair, which repose upon the base of the wings. The Diptera are also furnished with a small membranous appendage (alu/a) attached to the posterior base of the wing, which Kirby and Spence regard as the true analogue of the posterior pair of wings: they are, however, decidedly portions of the fore wings ; there exists a similar pair of winglets at the internal base of the elytra of the Dyticida, and which I have discovered equally developed in Hydrous piceus.
The Legs, or organs of terrestrial or aquatic progression, are six in number, attached in pairs to the three thoracic segments: tliey are arti- culated with the sternum of each segment, and are composed of a series of articulations, united together in such a manner as to permit the re- quired movements ; thus when the insect is pre-eminently cursorial, the basal articulations are freer than in those species which being nata- torial, the movement is confined to a simple working forwards and backwards of the leg. In other species, which are saltatorial, the hind legs are elongated and thickened, for the purpose of giving support to the strong muscles by which leaping is performed ; in other species, which are raptatorial, the forelegs are formed into a prehensile organ, whilst, in the fossorial species, the same legs are altered, so as to serve for burrowing or scratching in the sand. The chief divisions of the legs are the coxa, trochanter, femur, tibia, and tarsus: the coxa is the gener- ally large and flattened piece which articulates with the sternum, of a variable form, and which assumes its maximum degree of development in Dyticus and Haliplus : the trochanter is a small piece connecting the
14 MODERN CLASSIFICATION OF INSECTS.
preceding with the femur, which, in general, is the thickest part of the legs, and generally extends beyond the sides of the body. In leaping insects it is especially incrassated and often toothed beneath. The next piece is the tibia, which is generally nearly equal in length, but more slender than the femur; it is an important piece of the leg, and is often armed with various appendages, especially at its tip, where they generally assume the appearance of spurs, or acute points (calcaria). The tarsus is a jointed piece, armed at its extremity with one or two slender curved hooks (ungues), and often accompanied by membran- ous or fleshy cushions (plvilli). The number of joints in the tarsi varies from two to five. This part of the leg affords important characters for generical and family distinctions.
The Abdomen consists of a series of segments, for the most part destitute of any appendages, which last are to be found only in a few species, where they appear either as the external organs of genera- tion, or as filaments, or other analogous pieces attached to the ex- tremity of the body. These segments are attached together by membranes, and either meet at the edges or slide into each other like the tubes of a telescope; each of them is formed of two arcs, or semi-segments, one of which is dorsal, and the other ventral. The typical number of segments is rarely to be found in the perfect state, some of them being ordinarily employed in the construction of the organs of generation ; in the male Earwig, however, nine distinct ab- dominal segments, exclusive of the anal forceps, are to be observed. The abdomen is attached to the posterior thoracic segment, either by its whole breadth, when it is said to be sessile, or by a slender portion of its base, which is called a peduncle: this latter mode of articulation is especially to be observed in the majority of the Hymeno- ptera, and some other orders. The chief appendages of the abdomen are the ovipositor, which is extremely varied in the different tribes, (being sometimes replaced by a powerful sting,) and the elongated setz, forceps, or other analogous organs.
With respect to the Internal Anatomy of insects, a few brief notes must here suffice. They arrange themselves under the head of the sensitive, digestive, circulatory, respiratory, muscular, and generative systems.
The Senses of insects are dependent upon the action of the nerves, the system whereof in insects consists of a pair of medullary threads, exhibiting a series of knots or ganglia, whereby they are united
STRUCTURE OF INSECTS. 15
together at certain distances, but which ganglia are often confluent, especially in the imago state: from these knots or ganglia, an infinity of nerves ramify to the various organs, endowing them with the various senses of which insects are possessed. These are, vision, the external organs of which, the eyes, have been already shortly de- scribed ; hearing, which insects clearly appear to possess, but upon the precise organ of which naturalists are not agreed; smell, of which the external organ is also unascertained; taste, the seat of which resides in the parts of the mouth; and feeling or touch, whereof it has been generally supposed that the antennz were the chief organs, but of which the tarsi and the palpi have been respectively regarded as the instruments by various eminent physiologists.
The Digestive Organs consist of an elongated canal, composed of several distinct portions, which have been termed the pharynx, im- mediately connected with the mouth; the cesophagus, the craw, the gizzard, the stomach, and the intestines, terminating at the anal orifice : there are moreover a number of delicate elongated tubes, or biliary and salivary vessels opening into the digestive canal, the secretions whereof assist in the conversion of the food. The length of the canal varies greatly, being short in the carnivorous species, and often several times longer than the body in the herbivorous ones, whereof an ex- cellent example will be found noticed in the family Coccinellide.
The Circulatory System has only recently been proved to exist. It had been long noticed that a series of large reservoirs, easily to be perceived beneath the transparent dorsal skin of many caterpillars, termed the dorsal vessel, and supposed to be analogous to the heart, underwent alternate contraction and dilatation, but it was supposed that no aperture existed from these reservoirs, and consequently that there was no circulation. This, however, at length appears to have been de- cidedly proved, by the researches of Carus, Strauss-Durkheim, Bur- meister, and especially by the powerful microscope of Mr. Bowerbank, whose two papers in the Entomological Magazine sufficiently prove the existence of a circulation of a cold transparent and nearly colour- less fluid, not only in the larvae of ephemerz, &c., but also in the veins of the wings of the perfect Hemerobius.
The Respiration of insects is effected by means of two great canals (tracheee) running along the sides of the body, beneath the outer sur- face, and communicating with the atmosphere by means of numerous short tubes terminating at or near the sides of the body in breathing
16 MODERN CLASSIFICATION OF INSECTS.
pores (spiracles or stigmata), varying in their number ; internally the trachez emit an infinite number of ramifications, PSEQN Oy to all parts of the body like the branches of a tree, and thus oxyginating the cir- culatory fluid which is diffused from the series of hearts throughout the various organs.
The Muscles of insects are not unlike those of the higher animals, being however attached to the inner surface of the external covering of the animal, but being far more numerous than in the human bedy, Lyonnet having discovered more than four thousand in the caterpillar of the goat moth, whereas there are in man only 529; they are at- tached to the various organs intended to be brought into motion by their assistance, being either flexors or extensors, levators or depressors.
Of the System of Generation it will be sufficient to observe, that in insects the sexes are constantly distinct, each sex being provided with its own peculiar generative organs ; those of the female consisting of numerous ovaries and egg passages, and an ovipositor of varied con- struction, and frequently external ; and that the impregnation of these eggs is produced by the union of the sexes. It is also to be observed, that amongst certain tribes of insects which are social in their habits, and where a certain number of individuals are required to perform the duties of the community distinct from those of reproduction, this effect is produced by the peculiar nature of the food given to these individuals whilst in the larva state, which has the extraordinary effect of retarding the development of their generative organs; moreover, there are other tribes (Aphides) in which a single impreg- nation suffices for several subsequent generations.
Shortly after impregnation, the female commences the laying of her eggs, an operation not performed without the utmost solicitude on the part of the parent for the welfare of her future offspring. The various means adopted for the purpose of placing the eggs in such situations as will ensure a due supply of food to the young when hatched, are amongst the most extraordinary instances of instinct with which we are acquainted.
When the young insect first makes its escape from the egg, it appears in a form generally totally unlike that of its parent, although in a few instances its form is that which it will retain through life. It is now termed a caterpillar, grub, or maggot (larva), and has a series of moultings to undergo, during the last two or three of which its form is more or less altered. In the earlier stages
STRUCTURE OF INSECTS. 17
of its growth it is termed a larva, but the state immediately pre- ceding that in which it assumes its final or imago form is termed the pupa. The variations in the form of these several states in the different orders have been employed in the construction of the meta- morphotic distribution of insects, to be subsequently noticed, which will render it unnecessary to enlarge upon them at any great length in this place. Insects, in respect to the relation of the larva with the imago, may be divided into two principal divisions which I have termed Heteromorpha, or those in which there is no resemblance between the parent and its offspring; and Hotremorpha, or those in which the larva resembles the imago, except in the absence of wings. In the former the larva is generally worm-like and articulated in its form, of a soft and fleshy consistence, and furnished with a mouth, and often with six short legs attached in pairs to the three segments succeeding the head. In the latter, including the Orthoptera, He- miptera, Homoptera, and certain Neuroptera, the body, legs, and antenne are nearly similar in their form to those of the perfect insect, but the wings are wanting.
The Period of the Larvag State is that in which the operations of _ feeding are especially carried on, and during which, owing to the rapid growth of the insect, whereby the outer envelope from time to time becomes too small for the creature, it is necessary that a succession of moultings should take place. When, however, the insect has at- tained its full size as a larva it ceases feeding, prepares for itself a retreat formed of various materials, but often composed of silk drawn from the spinnerets of the lower lip, and which is termed a cocoon. Within this retreat the insect again throws off its skin and appears as a pupa or chrysalis, in which the rudiments of the limbs of the imago are more or less distinctly seen. In the nomorphous division, however, the pupa continues active, differing only from the Jarva in its increased size, and in having attained short rudimental wing- cases upon the back of the mesothoracic and metathoracic segments.
The Pupa is far more variable in its form in the different primary groups than the larve; and hence Fabricius* employed the cha- racteristic name of the variation of the pupa to designate the general nature of the metamorphosis, whereby the metamorphosis of an insect is incorrectly said to be zxcomplete in Scarabeeus and Apis, where the larva is quite unlike its parent, and the pupa pedate, but quiescent ;
* Philosophia Entomologica, p. 56. § 6. c
i8 MODERN CLASSIFICATION OF INSECTS.
whereas this kind of metamorphosis is the most complete of any of the changes which insects undergo. This impropriety has resulted from the improper application of the Linneean terms employed for the designation of the pupa: (Pupa completa, Spiders, &c. ; semicompleta, Grasshoppers, &c.; incompleta, Bees, Ants, &c.; obtecta, Butterflies and Moths; and coarctata, two-winged Flies.)
When the insect has remained a certain period in the pupa state it again sheds its skin and appears in its perfect and final state; at first, indeed, its external envelope is humid and soft, but it soon dries and hardens by exposure to the air, acquiring at the same time its various colours. These modifications in the outer forms of insects are also accompanied by equally important changes in their internal structure as well as by the modifications in the economy of the insect consequent thereon.
III. DrstrrisputTion or INSECTS INTO ORDERS.
This branch of Entomological Science has attracted much of the notice of naturalists in all ages; but it was not until the era of Swammerdam and of Ray, that philosophical principles were introduced and made the basis of the distribution of insects into primary groups. As, however, most of the systems of Entomology have been noticed at considerable length in the fourth volume of the Jntroduction to Entomology, I must refer the reader thereto, deeming it, however, essential to offer a few observations thereon.
Previous to the age of Swammerdam, the most vague ideas were entertained with respect to insects, of which the fanciful hypothesis of spontaneous generation was one of the most absurd. Hence, it is not to be wondered at, that Swammerdam (who together with Redi and Ray completely overthrew this system by establishing the theory of the universal developement of animals, ex ovo) was led to consider the principle of metamorphosis which he had so successfully employed against the spontaneous generationists, as of primary importance for the classification of insects. Indeed it is evident that the very nature of his studies, by which he had obtained so great a victory, could have had no other effect than to have produced the Metamorphotie System of Insects, detailed in his surprising and still invaluable Biblia Nature, and in which four primary divisions are established. j
1. Insects subject to a change of skin, but undergoing no change of form. (Spiders, Lice, Woodlice, and Centipedes.)
DISTRIBUTION OF INSECTS INTO ORDERS. 19
2. Hexapod insects subject to metamorphosis, having an active pupa, in which the rudimental wings and wing-covers are exposed. (Bugs, Grasshoppers, Dragonflies, Mayflies.)
3. Hexapod insects undergoing metamorphosis, in which the pupa state is quiescent, either having the limbs enclosed in distinct cases (Beetles, Bees, Wasps, &c., and part of the orders Neuroptera and Diptera), or covered in an entire case (Butterflies, Moths).
4. Hexapod insects undergoing metamorphosis, the pupa state having neither motion nor wings, but enclosed in an ovate case (most two-winged flies and some other insects, whose transformations were not clearly understood by Swammerdam).
Various other classifications of insects, from the nature of their metamorphosis, have been from time to time proposed by different authors, amongst which it will be sufficient to notice the two follow- ing: first, that of Lamarck, in which insects are divided from the structure of the pupa as follows : —
1. Those having an inactive pupa (Chrysalis), enclosed and entirely concealed in an opake covering. (1. a.) Chrysalis signata. Lepidoptera and some Diptera. (1. b.) Chrysalis dolioloides. Diptera (coarctata. ) 2. Those having an inactive pupa (Mumia), covered by a thin skin, through which the limbs of the insect are discernible.
Coleoptera. 2 Tumi : 3 (2. a.) Mumia coarctata Hymenoptera. (2. b.) Mumia pseudonympha - Neuroptera in part. Orthoptera.
3. Those having an active pupa(Nympha) + Hemiptera, —— Neuroptera in part.
The other proposed metamorphotic system is that given by Mr. Newman in an ingenious treatise published in the 9th Number of the Entomological Magazine, in which winged insects are divided as follows : —
Amorpha, in which the penultimate state is provided neither with mouth nor organs of locomotion; consequently it neither eats nor moves; neither does it bear any resemblance to the perfect state. Lepidoptera, Diptera.
Necromorpha, in which the penultimate state is provided with mouth and organs of locomotion detached from the body; but so
c 2
4
20 MODERN CLASSIFICATION OF INSECTS.
enveloped in a case that it can employ neither. The resemblance to the perfect state is very considerable, excepting in total want of motion. Hymenoptera, Coleoptera.
Isomorpha, in which all the stages are active and voracious, and of similar form. Orthoptera, Hemiptera.
Anisomorpha, in which appear the Amorphous, Necromorphous, and Isomorphous characters, together with a typical and distinct character. Neuroptera.*~—
It will be at once perceived, that these two systems are virtually almost identical. Mr. Newman’s divisions of the Amorphous Ader- mata and Dermata are also identical with Lamarck’s groups, Chrysalis signata and Chrysalis dolioloides, and the varied character of the Neuroptera is indicated by MacLeay. (Hore Ent. and Linn. Trans. xiv. p. 68.) A
Besides the five variations of metamorphosis constituting the diagnostics of Lamarck’s five groups or sub-groups mentioned above, there are others of minor importance, which ought not to be omitted in a system established exclusively on Metamorphosis; thus some of the aquatic pupe of the Adermatous Diptera are locomotive, whilst some of the Dermatous Diptera are nourished within the bodies of the parent, the Hippoboscide not being excluded from them until their arrival at the pupa state. Again, the Phryganeidz, Libellulide, Ephemeride, Chalcidide, Raphidiide, Coccide , and Aleyrodes, respectively offer various striking modifications in the nature of their metamorphoses; whilst in the pupa of Quedius tristis, Water- house, and some othet Staphylinideous larvee, the limbs are soldered as completely to the body as they are in the pupz of the Lepidoptera. Hence I consider that the nature of the metamorphoses, if alone relied upon, will not afford a sufficient mode of classification.
We will, therefore, next proceed to notice the Alary System, or that
vie : :
* In the Essay published by the same author, under the name of “ Sphinx vespi- formis,” this order is formed into a central circle, around which are arranged the six preceding orders, forming as many circles. The distribution of the English groups have been arranged by the same author in his Grammar of Entomology, and in a memoir inserted in the second volume of the Entomological Magazine, upon the same system; of which Dr. Burmeister has remarked — “ Die Eigenthumlichkeit dieses newen Systemes is vollkommen bezeichnet, wenn wir ein allbekanntes Urtheil darauf anwenden, indem ‘ das Gute desselben nicht neu und das neue nicht gut’ genannt werden kann.” Weigm.’ Arch. V.1. No. 4.
{ Trans, Ent. Soc. No. 1. pl. iii, f. 2. 2.
DISTRIBUTION OF INSECTS INTO ORDERS. A |
~
founded upon the structure of the organs of flight, of which Linnzeus was the founder, and by whom insects were thus classed in accordance with the gradual diminution in the number and consistency of their wings.
= horny, with a straight suture - - 1. Coleoptera. The anterior ; E ; : — semicrustaceous, incumbent - - 2. Hemiptera. 4 covered with scales - - - 3. Lepidoptera. Wings. All unarmed - - 4. Neuroptera. membranous, abdomen : ; Z with a sting - 5. Hymenoptera. 2 With a pair of halteres in the place of the posterior pair 6. Diptera. O i.e. Destitute of wings and elytra - - - 7. Aptera.
The celebrated Swedish naturalist, De Geer, proposed another distribution of insects in 1778, the primary and secondary divisions of which were formed for the most part upon the structure of the wings, which were also employed to characterise the orders. The structure of the mouth was, however, regarded as a secondary character of the latter; the consequence of which was the neces- sary and advantageous separation of the Caddice-flies and Ephemeree from the other Neuroptera, under the name of Elinguia*; and the division of the Linnzean Hemiptera into three orders — Siphonata * (Cicada, &c.), aga Dermaptera *, having a suctorial mouth (Cimex and Nepa), and Hemiptera* having a mandibulated mouth, (Mantis, Gryllus, Blatta, and Forficula.) The genera Coccus and Pulex were also respectively raised to the rank of distinct orders. The genera Termes and Psocus, together with the spring-tailed insects and lice, formed another order of the remainder of the Linnean Aptera, which were divided into two others, thus forming fourteen distinct orders.
This was a great step made in the distribution of insects; and the partial success with which the employment of the variation of the mouth was attended, probably induced Fabricius to construct his Cibarian system, founded upon the characters of the Trophi alone. This was as follows : —
* Mandibulated Insects. A. Two pairs of jaws. a. The lower pair Palpigerous. 1. Lleutherata (Coleoptera L., Beetles.) Maxilla naked, free.
* These names were proposed by Retzius, the commentator of De Geer ( Gen. et Spec. Ins. Lips.1783.) That of Dermaptera has, by some accident, been misapplied by English entomologists for the Earwig, instead of the Cimicide, for which it was proposed.
c 3
bo iS?)
MODERN CLASSIFICATION OF INSECTS.
2, Ulonata (Orthoptera Ol. &c. Grasshoppers, &c.). Maxillze covered with a blade.
3. Synistata (Neuroptera partly Z. and Thysanura Latr.), Maxille geniculated, and connected with the lower lip.
4. Piezata (Hymenoptera L., Bees, &c.) | Maxilla corneous, com- pressed, and often elongated.
5. Odontata (Gen. Libellula Z., Dragon-flies). Maxillee corneous,
toothed.
b. The lower pair of jaws, not palpigerous. Mitosata (Myriapoda, Centipedes).
>
B. One pair of horny jaws, armed with a claw. Unogata (Arachnida, Spiders, Scorpions).
Sa
C. More than two pairs of jaws. Polygonata (Isopod Crustacea, &c.). Jaws within the lower lip. . Kleistognatha (Brachyura, Crabs, &c.). Jaws outside of the lower lip. 10. Exochnata (Macrura, Lobsters, &c.). Jaws external, covered by palpi.
o ©
** Suctorial Insects. 11. Glossata (Lepidoptera, Z., Butterflies and Moths). A spiral tongue between palpi. 2. Ehyngota (Hemiptera Latr., Bugs, &c.) A rostrum enclosed in a jointed sheath. 13. Antliata (Diptera £., two-winged flies, &e.). Mouth with a soft exarticulate haustellum.
The harsh nomenclature of this system, joined with its many errors and the incongruous manner in which insects, the most widely apart, were brought into the same groups, producing altogether a most artificial and unnatural classification, gained for it but few followers.
The primary division, however, into mandibulated and suctorial insects, together with the adoption of some of De Geer’s more natural groups, were further steps gained in the approach towards a natural distribution.
Entomologists had, however, by this time learned from the advan- tages and disadvantages connected with the Metamorphotic, Alary and Cibarian systems, but more especially from the more decisive
DISTRIBUTION OF INSECTS INTO ORDERS. 95
views opened to them by the researches of comparative anatomists, that it was necessary to have recourse to the entire characters afforded by the insect in all its stages, and with reference also to its internal as well as external organization. Here, however, we are met, as might indeed be naturally expected, by difficulties of a far greater weight than were caused by the employment of a single character in its various modifications ; since a very slight acquaintance with any extensive group of insects will be sufficient to prove, that many characters which we find in one group, exhibiting a constancy of character, vary in the greatest degree in another, showing that although with the former they might, and indeed ought to be re- garded of the highest importance, in the latter they acquire but a secondary consideration; thus, whilst some groups which agree in their wings, disagree in the structure of the mouth; others agree- ing in metamorphosis, vary in their organs of flight.
The first attempts made towards the establishment of this, which has been called, the Helectic System, were of course partial ; thus Olivier, in 1789, proposed a system in which insects were divided into eight orders, distributed primarily from the wings, from which, as well as, secondly from the mouth, the characters of the orders were derived. Clairville on the contrary, in 1798, divided the winged insects into two groups, Mandibulata and Haustellata, from the structure of the mouth, whether furnished with jaws or a proboscis, characterising the orders from the wings. But it is to Latreille that we are indebted for the great advances made towards the perfection of this system. For nearly forty years was this author unceasingly occupied in im- proving the classification of insects; and it was by him that the introduction of family groups was effected, from which so great ad- vantages were derived, by greatly limiting the number of the secondary groups ; which from the cutting up of the more unwieldy Linnean genera had become very numerous. It is true that his numerous suc- cessive publications exhibit variations in the classification of some of the orders ; but this was the natural result of his labours, which were closed by the publication of his introductory work, the Cours d’ Ento- mologie, in which the Hexapod Annulosa formed his fourth Class Insecta, distributed as follows : —
24 MODERN CLASSIFICATION OF INSECTS.
A. Without wings, eyes generally simple.
a. Without metamorphoses. Ord. 1. Tuysanoura (Spring-tailed Insects). Mouth with jaws, tail forked. Ord. 2. Parasira (Lice). Mouth with a rostrum, tail simple.
b. Metamorphoses. (Pupa incomplete.) Ord. 3. SrpHonarTERA (Fleas).
B. With wings, eyes facetted, and occasionally also with ocelli. a. (Elytroptera.) Wings two, covered by two horny or coriaceous wing covers. * Mouth with four jaws. Ord. 4. CoLtrorreraA (Beetles). Wing-covers horny. (Pupa incom- plete.)
Ord. 5. DermMAprera (Earwigs). Wing-covers horny. (Pupa semi- complete.)
Ord. 6. Orruorrera (Locusts, &c.). Wing-covers horny. (Pupa semicomplete.)
** Mouth, an articulated proboscis, enclosing sete. Ord. 7. Hemiprera (Bugs and Cicade, &c.). (Pupa semicomplete.) b. Gymnoptera. Wings four, or two naked. * Wings four. + With four jaws. Ord. 8. Neuroptera (Dragonflies, &c.). Wings reticulated. (Me- tamorphoses various. ) Ord. 9. HymenoprEerA (Bees, &c.). Wings veined. (Pupaincom-
plete.) ++ With a spiral tongue between two palpi. Ord. 10. LeprpopterRA (Butterflies). Wings powdery. (Pupa ob- tecta.) ** Wings two. Ord. 11. Raipiprera (Bee-parasites, &c.). Balancers, two in front of wings.
Ord. 12. Diprera (Flies). Balancers, two behind the wings.
There are, however, various objections to this system, which it will be seen has for its primary character the variation of the wings ; thus, for instance, whilst the semicomplete-metamorphosed Orthop- tera and Hemiptera were brought into contact, the mandibulated Neu- roptera were separated from the Orthoptera, to which they are so
DISTRIBUTION OF INSECTS INTO ORDERS. 25
nearly allied. Mr. MacLeay has offered some other objections against Latreille’s system, which he describes as one “ expressing more importance to the aerial organs, and the texture of the body, than to the modifications of those organs upon which the very existence of the animal depends. ‘ Ainsi,’ says Lamarck, ‘les carac- téres si importans de la bouche ne furent nullement considérés, et cédérent leur préeminence aux organes si variables dela locomotion dans lair’ ”’—Hore Ent. p. 360. I can, however, by no means agree with Mr. MacLeay, in the inferior rank given to those organs, which, as before said, are the very organs which, by their extraordinary deve- lopement, prove the winged insects to be the centre or types of the annulose animals. Moreover, the variableness attributed by Lamarck to the locomotive organs is not less striking in the oral organs of some groups, as in the Lepidoptera, Hymenoptera, and Neuroptera ; thus evincing the correctness of the remark already made upon the insufficiency of a single character, when attempted to be relied upon as an infallible key to the classification of this class of animals. This is not denied even by those authors who are disposed to admit the or- ganisation of the mouth, as of the highest importance in the classifica - tion of insects; thus Savigny approves of the Lamarckian divisions into “ broyeurs”’ and ‘ suceurs;” but adds, ‘ je ne pense pas qu’on puisse tirer le caractére de ces divisions de la présence ou de l’absence des mandibules,” because he considers the mouths of all insects to be “ essentiellement composée des mémes ¢lémens.” (Mémoires, ch. 1.). Latreille also has suggested the division of insects into two other groups, Gymnostomes, or those which have the parts of the mouth naked, and Thecostomes, or those in which some of them are bristle- like, enclosed in a sheath. The Lepidoptera, as well as the Hymenop- tera, enter into the first of these divisions. (Fam. Nat. 334. 417.) In like manner, Dumeril (Cons. Gen. p. 9.) has remarked, that the lower lip and jaws of some of the Hymenoptera form an apparatus, having “ le double faculté de broyer les alimens, et de les pomper par une sorte du succion;” and Messrs. Kirby and Spence (Jntr. 111, 417.) remark, that “ If the mode in which insects take their food be strictly con- sidered, it will be found that in this view they ought rather to be regarded as forming three tribes ; for the great majority of the Hyme- nopterous order, and perhaps some others, though furnished with mandibles and maxilla, never use them fer mastication, but really lap their food with their tongue: these, therefore, might be denominated
26 MODERN CLASSIFICATION OF INSECTS.
9
‘lappers.’” And at a subsequent page (vol. iv. p. 366.), they regard Clairville’s Mandibulata and Haustellata as secondary groups*, “ be- ing convinced from the numerous characters they possess in common, notwithstanding the different mode in which they take their food, that they form one connected primary group; in which opinion they are further confirmed by the variations that take place in their mode of feeding in their different states ; some from masticators becoming suctorious (Lepidoptera), and others from being suctorious becoming masticators, (Myrmeleon, Dytiscus) ; which shows that this character does not enter the essential idea of the animal.”
Much of the difficulty which has arisen upon this branch of our sub- ject has been produced by regarding the relations of insects as exhibit- ing themselves only in a linear series. It is not necessary for me here to advocate either the quinary or the circular disposition of groups ; but when I perceive that, by quitting the linear series we gain a decidedly greater number of points of contact, whereby groups, which in such linear series must be kept widely apart, are brought into juxta-posi- tion, I cannot but think, that some other than an undeviating chain of linear affinity is the correct mode of viewing the productions of nature. Now, Mr. Macleay, by his system (which may be termed the Representative System), has obviated more of the difficulties already pointed out, than could by possibility be effected by any of the previous systems. Here, indeed, as he observes, in the Hore Ento- mologice, “a beautiful regularity is perceivable, comprising those dis- tinctions of Trophi, insisted upon by Fabricius, Cuvier, and Lamarck ; those relations of metamorphosis, constituting the leading principle of De Geer, Olivier, and Latreille; and, finally, those characters from the organs of locomotion, upon which orders were founded by Aris- totle, Ray, and Linnzus.”
By reminding the student, that in the following tables the first and fifth order in each series are supposed to be as nearly related to each other as any other two contiguous orders (each series returning into itself and forming a circle), an idea may be gained of Mr. MacLeay’s mode of distribution. The names printed in italics are
* Mr. MacLeay, in order to establish a quinary distribution of the Annulosa, con- sidered the Mandibulata and Haustellata as classes, each possessing an equal rank with the Crustacea, Arachnida, and Ametabola. I cannot adopt this view; which, indeed, Mr. MacLeay has himself partially modified in his Memoir in the 14th volume of the Linnean Transactions,
DISTRIBUTION OF INSECTS INTO ORDERS. a7 |
“a
either those of the osculant groups connecting the orders, or of those insects which point out the connection of the orders. Distribution of metamorphotic winged Insects from the “ Hore Entomo- logice,” ii. p. 367., and “ Linnean Transactions,” xiv. p. 67. MANDIBULATA. Relations of Analogy. HAUSTELLATA.
1. Tricuorrera ? K. Metamorphosis obtected. (Phryganea, Perla,{ Larva with membranous $1. Lepinoprera Linn. Tenthredo. ) legs, Ord. Bomboptera M. L. G. Pterophorus. (G. Sirex.) G. Psychoda. DMEF oea we Lins Metam. incomplete or f © pen, De coarctate. ; 2. Dierera Arist. ‘ Z Larva apod, or vermiform. O. Strepsiptera K. Fam. Hippoboscide. G. Atractocerus. G. Nycteribia, leading to Arachnidous class. 8. Corrortera Arist. Metam. incomplete. 3. Aprera (the Flea). Larvz various. (Larva apod, but of Co-
leopterous structure ), Q. Dermaptera Leach. ( Earwig. )
4 UG opens Ole f Metam. semicomplete. ]|4. Hemrrrera Linn,
| Larva resembling Imago. J (G. Cimez. ) G. Mantispa. Fam. Hydrocorisa. O. Raphioptera M. L,
(G. Boreus. )
5. Homoptera.
Metam. various, includ-) - 5, Neurorrera Linn. hs ing subsemicomplete. } (G. Cicada, &c:) arve hexapod. O. Megaloptera (G. Flata limbata Fab. & Sialis). Perlides, \ead- G. Aleyrodes, leading ing to Trichoptera. to Lepidoptera.
The point of connection of the two circles, formed by the returning of these two series into themselves, is supposed to exist between the Trichoptera, and Lepidoptera, being effected by such species of the latter as have very long antenne, like the Adele, and such as have the trophi obsolete, as Aglossa.
That objections existed against this mode of classification on various points of view, Mr. MacLeay was well convinced, and of which indeed he has enumerated several in the sixth Chapter of the Hore Entomologice, besides which it may be urged that the minor details, and especially the construction of the majority of the osculant groups and orders are quite untenable Still, however, as a whole, it must be admitted that the various Metamorphotic, Alary and Cibarian cha- racters have never before been brought to bear together with such weight as in the system now before us; which, far from regarding as all-perfect, I would rather consider as the first stepping-stone towards a natural system not founded upon a linear series of affinities, but
28 MODERN CLASSIFICATION OF INSECTS.
which it will require far greater investigations and much more elaborate research to bring to perfection than it has hitherto received.
Modifications of the system of Mr. MacLeay have been published by Messrs. Kirby and Spence, and Mr. Stephens. The former authors (Introd. iv. p. 868.), distribute the Metamorphotic Insects in the following manner.
* Orders in which the ordinary Trophi all occur, or the mouth is perfect.
1. Coleoptera. 2. Strepsiptera. 3. Dermaptera. 4. Orthoptera. 5. Neuroptera. 6. Hymenoptera.
** Orders in which all the ordinary Trophi do not occur, or the mouth is imperfect. a 7. Hemiptera (divided into Heteroptera and Homoptera as sub- orders). 8. Trichoptera. 9. Lepidoptera. 10. Diptera. 11. Apha-
niptera (the Flea).
Here we find the Strepsiptera interfering to prevent the passage between the Coleoptera and Orthoptera effected by the earwigs, the Neuroptera and Trichoptera far asunder, and other equally great objections.
Mr. Stephens (Illustrations Brit. Ent. Mand. i. p. 2., Haust. i. p. 2.), adopts the divisions, Mandibulata and Haustellata, placing in the former the orders, 2. Strepsiptera, 3. Coleoptera, 4. Dermaptera, 5. Orthoptera, 6. Neuroptera, 7. Trichoptera, 1. Hymenoptera ; and in the latter the orders, 13. Hemiptera, 14. Homoptera, 8. Lepi- doptera, 9. Diptera, 10. Homaloptera, 11. Aphaniptera, and 12. Ap- tera (or the lice which I have excluded, as above mentioned). Like Mr. MacLeay, Mr. Stephens regards these two groups as forming circles, the extremities of each being allied together.
Against these and other classifications *, which it would be too tedious to detail, and in which the number of the Linnzan orders is much in- creased, objections exist on the part of some authors who consider it more natural to place the Strepsiptera, Earwigs, Forest-flies, Fleas, Thrips, &c., in some of the old orders. Such groups, it is true, are of far more limited extent, and also of less decisive characters than the great groups; but it appears to me to be as erronecus to force them
* Amongst these, that proposed by M. Laporte, in his Etudes Entomologiques, ought not to be omitted ; in which Termes, Libellula (with Perla and Ephemera), Aphis, and Coccus, are raised to the rank of distinct orders,
DISTRIBUTION OF INSECTS INTO ORDERS, 29
into the larger groups as it is to regard them as orders equivalent with the great ones, such as Coleoptera, Lepidoptera, &c. It is indeed impossible not to be convinced that in many of these groups the structure of one order is partially lost, and that of the adjacent one assumed, whence it would be as correct to place the insects in the one as in the other order. Indeed, it appears to me to be the strongest evidence of the forcing of nature to a preconceived system, when we see introduced amongst a number of animals, characterised by a constant peculiarity of structure in some important organs, an animal, or group of animals, which does not possess such character, and find such a step attempted to be justified on the plea that this organ is variable in other groups,
With a view to avoid some of the inconveniences which still exist in the distribution of insects, I would propose the following classifi- cation in which the approach of the Hymenoptera to the Trichoptera, and of the Lepidoptera towards the Diptera, as pointed out by Mac Leay, together with the direct passage between the Trichoptera and Lepidoptera, and the descending series in the first, and the ascending series in the second columns must be borne in mind.
CLASS OF HEXAPOD METAMORPHOTIC INSECTS.
Subclass, Mouth with jaws. Subclass, Mouth with a sucher. (Dacnostomata VW.) (Antliostomata W.) Ord. HyMENOPTERA. Ord. Diptera. ? Osculant Ord. Strepsiptera. —_? Osculant Ord. Homaloptera. Ord. CoLEOPTERA. ? Osculant Ord. Aphaniptera.* Osculant Ord. Euplexoptera (Earwig). Ord. ORTHOPTERA. Ord. Hereroptera (includ- ing the Water Bugs). ce Ghhrips': )- ? Ord. NEUROPTERA. Ord. Homoprera. ? ?
Ord. Tricuoprera (Phryganea alone). Ord. LeprpopTera. ? ? Moreover, the relationships existing between the opposite orders must not be overlooked. Since, in some instances, as between the
* The relationship of the Flea with the Hymenoptera, insisted upon by Strauss, does not appear to me to be well founded.
+ Order Thysanoptera Haliday, by whom an elaborate paper upon this tribe of remarkable insects has been published in the Lntomo!. Mag. No.15. April, 1836.
50 MODERN CLASSIFICATION OF INSECTS.
Orthoptera and Heteroptera, they are so strong as to become direct affinities according to some authors. As, however, it has been the fashion to commence the series of orders with the Coleoptera (which it is true is the greatest in extent), I have followed the general plan, the effect of which will be the introduction of the Hymenoptera and Strepsiptera between the Trichoptera and Lepidoptera. With this explanation we take our leave of the modern distributions of the orders of insects, which from the nature of the present work it would have been in- consistent to have passed over in a more concise manner; the reader having been put into possession, not only of the modes of classification adopted by all the most recent authors of any eminence, but also of the principles upon which such distributions have been proposed.
4
IV. BiBLioGRAPHICAL REFERENCES. (COLEOPTERA.)
(A. Cotrortera IN GENERAL. )
Fabricius. Syst. Eleutheratorum, 2 vols. Kil. 1801. 8vo.
Olivier. Entomologie, 8 vols. Paris, 1789. 4to.
Schonherr. Synon, Insect. Holm. 1806—27. 4 vols. 8vo.
Dejean. Catal. des Coleopt. Paris, 1821. 8vo. 2d edit. Paris, 1833. 3d edit. 1836.
Dejean and Latreille. Hist. Nat. Col. d’Europe, livr. 1—3. Paris, 1822. 8vo.
Dejean and Boisduval, Hist. Nat. Col. d’Europe, livr. 1—. Paris, 1829—. 8vo.
Germar. Insect. Noy. Spec. Coleopt. Halle, 1824. 8vo.
Jablonsky and Herbst. Natursystem aller Bekannten Insecten. Berlin, 1783—95, 10 vols. (40 cah.) 1783—95. 8yo. with atlas of plates.
Voet. Descript. et icones Coleopt. La Haye, 1766, 2 vols. 4to. pl.; ditto, Ger- man edition by Panzer, 4to. Erlang, 1793.
Schwartz. Nomenclatur uber die Roselschen und Kleemanschen insecten. Erste Abth, Kafer. Nurnberg, 1793. 4to.
Knoch. Beitrage zur Insecten geschichte. Leipz. 1781. 8vo.
Knoch. Neue Beitrage zur insecten kunde. 1801. 8vo.
Thunberg. Numerous Memoirs in the Swedish Trans. (Vet. Acad. n. Hand. 1781. 1804. 1806. 1814. 1816. and in Noy. Act. Upsal.)
Bilberg. In Nov. Act. R. Soc. Upsal, t. 7.; and in Mem. Acad. Se. St. Petersb. t. 7. 1820.
Miiller. P. W. G. in Germar’s Magazin. vol. iii. and iv.
Schaller. Neue Ins. (Coleopt.) in Naturf. Gesellsch. Hale, b. i.
Laporte. Etudes Entomologiques. Paris, 1834. 8vo.
Laporte and Gory. Hist. Nat. des Coléopteres. Paris, 1835. S8vo. in parts.
Audouin and Brullé. Histoire Nat. des Insectes. vol. iy. 5. 6. Coleopt. Paris, 1834. 8vo.
Bonelli. Obsery. Entomol. ‘Turin, 1809, 4to.
Vigors, in Zool. Journ. Nos. 3, and 4,
Kirby, in ditto, vols. i. ii, iii,
BIBLIOGRAPHICAL REFERENCES. , 31
Phe. Cole oftar ats Man wal . Fa. 112-3. Ladbor~ fo. 16-357 ffir
Hope. Descriptions new Coleopt. in Zool. Soc. Trans. vol. i. 4to.
Duncan. Naturalist’s Library. Beetles. Edinb. 1835. Small 8vo.
Léon Dufour, Recherchés Anatomiques sur les Carabiques et sur plusieurs autres Insectes Coléopteres in Annales des Soc. Nat. 1824—.
B. Correorrerous Faunists. a. Great Britain.
Martyn. The Engl, Entomologist, exhibiting all Coleopt. Insects, &e, Lond. 1792. Martin Lister. De Scarabzxis Britan. in Hist. Insect. Raii. Lond. 1710. 4to. Marsham. Entomologia Britannica, tom. i. Coleopt. Lond. 1802. 8vo.
Stephens. lustr. Mandibulata, vols. i—v. 1828—. Hacasl Britt bclreplon Lena
Burrell, in Trans. Ent. Soe. vol. i. (old series. )
Wilson and Duncun. Entomol. Edinensis. Edinb. 1834. 8vo.
Dillwyn. Observations on the Coleopt. of Swansea. Swansea, 1829. 8yo. Pri- vately printed.
b. Other Countries of Europe.
Boisduval and Lacordaire. Faune Entomologique de Paris. 1 vol. 12mo. 1836.
Illiger. | Verzeichniss der Kafer Preussens, &e. Halle, 1798. 8yo.
Iiliger. Supplement to ditto, in his Magazine. vol. i. 1802.
Kugellan. Kafer Preussens in Schneider’s Ent. Mag. vol. i.
Erichson. Kafer der Mark Brandenburg. vol. i. 8vo. Berlin, 1837.
Creutzer. Entomol. Versuche. Wien, 1799. 8vo.
Kunze. Entomolog. Fragmente. Halle, 1818. 8vo.
Nicolai. Dissert. Col. sp. Halensis. Halle, 1822. 8vo.
Sturm. Deutschland Fauna. Kafer. 1805—-38. 10 parts. 18mo.
Harrer. Beschreibungen zu Schaeffer’s Naturl. Ausgemahlten Abbildung. Re- gensburg. Insecten. Regensb. 1791. to.
Preysler. Verzeichniss Bohmischen Insecten. Prag. 1790. 4to.
Hoppe. Enumeratio Insect. Elytratorum cirea Erlangam indig. Erlang. 1795. 8yo.
Duftschmidt, Fauna Austriz, oder Beschreibung der Osterreichischen Insecten &c. Linz. 1803—1825. 3 vols. 12mo.
Beck. Beitrage zur Baierschen Insecten Fauna, &e. Augsburg, 1817.
Gistl. Enumeratio Coleopt. Monacensis. Munich, 1829.
Waltl. Verzeichniss Kafern in Spanien. &c. Isis, 1832.
Panzer. Beschreibung seltener Kafer (in Der Naturforsch. 24. stu.)
Frihlich. Ditto in ditto. stu. 23. and 26.
Ahrens. Beitr. Deutscher Kafer. in Neue Schr. Naturf. Gesells. Halle, y. ii.
Malinowsky, in ditto. vol. i.
Eschseholtz, Zoologischer Atlas. - fol. Ditto, in Mem. Act. Nat. Mose. vol. vi. Ditto, in Mem. Acad. Soe. St. Petersburg. vol. vi.
Palliardi. Beschreibung zweier Decaden. 8yo. Wien, 1825.
Clairville. Entomologie Helvetique. Zurich, 1798—1806. 2 vols. 8vo. ( Pre- daceous Land and Water Beetles and Weevils).
Laicharting. Der 'Tyroler Insecten. Zurich, 1781—8vo.
Bassi, in Annal. Soc. Ent. de Paris. 1834.
Sturm and Hagenbach. Coleoptera Alpina (Noy. Act. Cas. Acad. vol. xii.)
Villa. Coleoptera Europe Dupleta, Milan, 1833; with suppl. of n. sp.
a . 4 a al Ae { X 4tdo Shu a) Shacetdan 1b. Gale fo ate o YS belrrwrceales . / /
i ut
Clu oe Se ar /3 teed xs bbi.tén. clos
b
oh
tan
/yvl.§ vo-/F.
Ava 1ORF
] Part J. 4939.
32 MODERN CLASSIFICATION OF INSECTS.
Géné, in Act. Acad. Turin. tom. 39. (Italian Coleoptera).
Detharding. Disputatio de Insectis Coleopt. Danicis. Buetzovii, 1763. 4to.
Pauli. Dissert. de Coleopteris Danix. 4to. Butzov. 1763.
Paykul!, Fauna Suecie, 8vo. Upsal, 1798. 1800.
Zetterstedt. Insecta Lapponica. Leipz. 1838. 4to. vol. i. fase. 1. pp. 191. f. 2.
Hoffmansegg. Portugeis Kafer, in Hliger’s Magaz. vol. ii. and vi.
Gyllenhal. Insecta Suecica. Classis 1. Coleopt. Seara. 1808. 1827. 4 vols. 8vo.
Sahlberg. Dissert. Entomol. Insecta, Fennica enumerans Coleoptera. Abo. 1817. - 8vo.
Eschscholtz. Nova genera Coleopt. Fauna Europe. Bull. Soc. Mose. vol. ii.
Zoubhoff, Krynicki, Mannerheim, Faldermann, Steven, Gebler, Hummel, Bohemann, and Fischer. Descriptions of many detached Russian Coleoptera by these Authors in the Memoires and the Bulletin Soc. imper. des Naturalistes de Moscow.
Pallas. cones insect. Rossiz Siberieque. Erlang. 4to. 1781.
CapAsia.
Germar. Mem. sur la Faune Ent. de Kolywan (Siberia) in Isis. 1825.
Faldermann. Coleoptera of China, Mongolia, &e. Ditto Coleopt. Persico, Ar- meniz (in Mem. Acad. imp. St. Petersb, tom. ii. 1835.)
Perty. Observationes nonnulle in Coleoptera Indiz Orientalis. Monachii, 1831. Ato.
Wiedemann. Several articles in his Zoologisches Magazin. vol. i. and ii.
Hope. Synopsis of Nepaul Insects, Coleoptera (in Gray’s Zoological Miscell. No. i. London, 1831. 8vo.)
MacLeay. Annulosa Javanica. London, 1824. 4to.
d. Africa. Goldfuss. Enumerat. Insect. Eleuth. Capitis Bone Spei, totiusque Africe. Erlang, 1804. 8vo. Hausmann. Ins. Cap. Bon. Sp. Iliger’s Mag. vol. vi. Wulfen. Deser. quorund. Capens. Ins. 4to. Erlang, 1794. Klug. ‘Bericht tuber eine auf Madagascar sammlung Coleoptera. Berlin,
1833. 4to. (From the Abhandlung. der Kongl. Akad. der Wissenschaft, zu Berlin. )
e. Australasia.
Schreibers. Descriptions of some New Holland Coleoptera (in Linn. Trans. ), vol. vi. Hope, in Trans. Ent, Soc. Lond. vol. i. pl. 1, 2.
Sf. America.
Say. Descriptions of Coleoptera of North America, in Journ. Acad. Nat. Sci- ences, Philadelph. vol. iii. and y. Contributions to the Maclurian Lyceum of Philadelphia, in American Phil. Trans. vol. iv. New series. And in the Journal of the Nat. Hist. Soc. Boston, vol. i. 1836. ;
Leconte, in Annal. of Lyceum of New York. December, 1824.
Harris, in &4OQ Val. Heat. Sec, Hartinr, vol-l,
Hentz, in Journ. of Acad. Nat. Science. Philadelphia. vol. v.
Payhull, Yaune Insect. Amer, Boreal. Prodromus, 4to. Erlang. 1794.
COLEOPTERA. 33
Chevrolat. Coléoptéres du Mexique. Strasbourg, 1834. 12mo. in parts.
Klug. Entomologiz Brasiliz Specimen, and Specimen alterum (in Nova Acta Nature Curios. vol. x. and xii. ).
Perty. Delect. An. Articul. Brasiliz. fol. Munich, 1830.
Sahlberg. Periculi Entomologici (chiefly Brazilian Coleoptera). 4 pl. 1823. 8vo.
Lacordaire. Observations on the Habits of the Beetles of Brazil (in Annales des Sci, Nat. for 1830, and in Nouy. Ann. du Muséum).
Orver COLEOPTERA Aristotle.
Cuar. Anterior wings (elytra) horny or leathery, concealing the posterior wings when unemployed, horizontal, united down the back by a straight suture.
Posterior wings membranous, longitudinally and transversely folded when unemployed.
Mouth with transversely moveable jaws.
Pupa incomplete.*
This order comprises the extensive tribes of beetles, and, in respect to the size of some of these insects, or the number of individual species, must be regarded as occupying the foremost rank amongst insects. From our earliest childhood, when, at school, we learned the cruel trick of putting a pin through the tail of a Cockchafer to see it “spin ;” or caught the pretty ladybirds, and watched them take their flight from our hand, exclaiming, in the words of the childish couplet —
“ Ladybird, ladybird, prythee begone ; Thy house is on fire, and thy children at home,”
we have been familiarized with the leading character of this order of insects, derived from the structure of the wings and elytra, from whence, indeed, the name of the order, signifying “wings in a case,” was given to them by Aristotle. In the earlier editions of the Sys- tema Nature it was even employed as its sole characteristic ; so that the Grasshoppers, Cockroaches, and Earwigs were included in it; and,
* It has been usual to apply the character of the pupa to designate the peculiar nature of the metamorphosis in general. This is, however, very incorrect; since the Coleoptera are thereby defined to haye an incomplete metamorphosis, whereas their metamorphoses are complete, in the ordinary acceptation of the word, the pupa being on the contrary incomplete. Moreover, Linnzus applied this and other similar terms to the pupa, and not to the metamorphosis; the confusion originating in their misappropriation by Fabricius,
*D
34 MODERN CLASSIFICATION OF INSECTS.
in the last edition of this work, the last-named insect was still retained, from the straight suture by which the upper wings are united when unemployed. These insects, however, although nearly agreeing with the beetles in the structure of the mouth, are widely separated by the nature of their metamorphoses, which consist in a simple cast- ing of the skin from time to time, the larva nearly resembling the perfect insect, the pupa being also similar, but furnished with rudi- mental wings, and being active; whereas, in the beetles, the larva is quite unlike the beetle, and the pupa inactive. So that, on the whole, this may be considered as one of the most definitely marked groups of the animal kingdom. Mr. MacLeay has indeed (Hore Entomol. p- 42].) considered its insulation as the result of accident, although it is a group of that precise kind which he has since proved (in his Letter upon the Dichotomous System) to be a perfectly natural one.
The number of species of this order with which entomologists are actually acquainted cannot be less than 35,000*; and it is more than probable that, when the number of species of foreign climes shall have been collected, the number will be doubled, if not trebled, as we may, indeed, conceive by noticing the great proportion of European species already in the lists. In this surprising number we find the most brilliant colours and most singular forms, with a variation of size from the most bulky to the most minute of the insect tribes. Hence, it is not surprising that these insects should have attracted the great, and even absolute, attention of so many authors and ama- teurs; and which has probably been increased by the comparative largeness of size and superior consistence of their external covering, whereby they are rendered so much easier in collection, preservation, and examination.
In addition to the characters given above the following may be men- tioned : — The head is furnished with a pair of antenne, of which the form varies very greatly, even in the sexes of the same species, but which, except in a few instances, consist of eleven joints. The eyes are large and lateral ; and, in a few species, each is divided by a horny sinus (being part of the skull), so that in these four eyes appear to exist, an example of which occurs in Gyrinus. ‘The simple ocelli are here wanting, except in Paussus bucephalus, and some of the small Staphylinidz, in which Dalmann observed two tubercles on the crown of the head, which he considered to be ocelli, but Latreille
* The Berlin Museum alone contains 28,000 species.
COLEOPTERA. 35
considered them as mere tubercles. Curtis and Brullé have, however, recently discovered a small frontal tubercle in some of the Dermes- tide, which appears to be an ocellus, as it is indeed named by the former. The mouth consists of an upper lip, generally transverse ; a pair of horny jaws; a pair of lower jaws of less firm consistence, furnished with an articulated palpus, and a lower lip, also furnished with a pair of similar but shorter palpi; the lip itself (labium) arises from a transverse horny plate (mentum), articulated at its base with the head. The second segment is by far the largest ; it is generally termed the thorax, but, strictly, it is only the first thoracic segment (prothorax), having the fore-legs articulated to it on the under side. Latreille has claimed the discovery of a pair of spiracles on the inner membrane of this segment; Messrs. Kirby and Spence had, however, previously noticed it. (Introd. iii. p. 43.) The second thoracic segment (mesothorax) is very short, and is scarcely visible from above when the insect is at rest, the only ex- posed part consisting of a triangular plate (scutellum, or, more strictly, scutellum of the mesothorax), on each side of which the elytra are attached ; this segment also bears on its under surface the middle legs. The third thoracic segment (mesothorax) is larger in size than the preceding, bearing on its upper surface, near the anterior angles, a pair of membranous wings, and on its under surface the pos- terior legs, of which the basal joint is large and transverse. The ab- domen is intimately attached by its entire breadth to this last tho- racic segment, and generally consists of six or seven segments (the typical number being nine), each having a pair of spiracles at the sides, or upper surface ; the tarsi vary in the number of the joints, from two to five.
In an order of such extent as the present, it is not surprising that some few variations from these typical characters should exist ; thus in some, as the females of the Glowworm and Drilus, the organs of flight are entirely wanting. Many genera of Carabide, Curculionidae, &c. have elytra, but no wings: in some the elytra are soldered together; others again, as Molorchus, Buprestis, &c. have the wings only longitudinally folded; others, as Meloe, have the elytra lapping over each other; and in some the elytra are narrowed, and do not meet in a straight line down the back, — Sitaris, &c. The only character which prevails throughout the order is that derived from the metamorphosis, which is of that species which has been termed
D2
36 MODERN CLASSIFICATION OF INSECTS.
(but erroneously), incomplete, the term being applicable only to the pupa state. The darve are active, and generally resemble short thick worms, with the head and back of the anterior segments of the body scaly, and generally with a pair of short jointed legs attached to each of the first three segments, representing the three thoracic segments and the six legs of the Imago. The head is furnished with the same organs as that of the perfect insect, except that they are more rudimental; thus the eyes are represented by several small ocelli-like tubercles observable at the side of the head, and the antennze are very short ; the parts of the mouth are, however, more developed, as might be expected from the circumstance of the larva state being that in which the chief amount of nourishment is taken. The more inactive and retired these larve are, the more they resemble worms or maggots: thus the Nut Weevil, which is borne in the midst of its food, is destitute of legs. The larvae of the carnivorous species are the most active, being furnished with robust legs; whilst some of the larvee of the herbivorous species have the legs replaced by fleshy tuber- cles, which are even occasionally wanting. The last segment of the body is often furnished with horns or other appendages, and its under side provided with a fleshy retractile tubercle, which is employed as an additional leg, and even, as is asserted respecting the larva of the Glowworm, as an instrument ‘employed in cleansing the body when soiled. Until the publication of the Hore Entomologice, entomologists had neglected the study of these larvae since the time of De Geer. Within the last two or three years, however, the attention of several authors has been directed to this branch of the subject, including Messrs. Waterhouse, De Haan, and Hammer- schmidt. I have also been long engaged in the like manner, having observed the transformations of many species, as more fully to be mentioned in the various families. The pupa is inactive, the insect taking no nourishment in this state, it is of a dirty white colour, and often enclosed in a case composed of bits of earth or chips of wood, united by silken threads or viscid matter formed by the larva previous to undergoing its change. In this state the parts of the future beetle are plainly perceivable, being incased in distinct sheaths; the head is applied against the breast; the antenne lie along the sides of the thorax; the elytra and wings are short, and folded at the sides of the body, meeting on the under side of the abdomen; the two anterior pairs of legs are entirely exposed,
COLEOPTERA. 37
but the hind pair are covered by the wing-cases, the extremity of the thigh only appearing beyond the sides of the body. In some few species of Brachelytra, as already mentioned, the limbs are so firmly soldered to the body, and apparently under a single sheath, that the pupa might almost be termed an obtected one. Some pupz are enclosed in the cases in which the larve resided; but the habitations, economy, habits, and period of duration in the various states of these insects vary exceedingly.
Naturalists are indebted to Aristotle for the establishment of this order of insects, which embraced, according to his definition, all insects with encased wings, “ ocx to mrepoy eyes ev koAew,” and which has been adopted by subsequent authors. Fabricius, however, changed its name to Eleutherata, and Clairville to Elytroptera.
Linnzus described 891 species of these insects, which he divided into 29 genera, distributed into the three following sections : — *Antennis clavatis extrorsum incrassatis, * * Antennis _ filifor- mibus, and * * * Antennis setaceis. Geoffroy proposed numerous additional generic groups, and adopted a different mode of classifi- cation, dividing the order into two primary sections, from the elytra entirely or partially covering the abdomen, and subdividing these sections according to the numerical variations in the joints of the tarsi. This latter character was subsequently adopted by Olivier to distin- guish the primary sections of the order, which were thence named as follows : — 1. Pentamera, in which all the tarsi are 5-jointed; 2. Heteromera, in which the four anterior tarsi are 5-jointed and the two posterior 4-jointed ; 3. Tetramera, which have four joints in all the tarsi ; and 4. T7imera, having only three joints in all the tarsi. To the four above-mentioned. groups Latreille added another, named Dimera, composed of the minute family Pselaphide, which were supposed to have only two joints in all their tarsi. He, also in the Regne animal, proposed a sixth section, Monomera, for the reception of the Clambus armadillo, supposed to possess but a single joint in the tarsi; but the real affinities of these insects (the former with the Staphylinide,' and the latter with the Agathidiide), as well as the real. structure of the tarsi, do not warrant the establishment of these two sections, the Pselaphide having three joints, and the Clambus, as I have ascertained, having four joints at the least in the tarsi. Cuvier, in his Tableau Elementaire, divided the order into 13 principal divisions, characterised by the forms of the antenne, tarsi, palpi, and elytra.
x DS
Welkedag in Leotegeeat Jornal cay S. / C :
‘a
38 MODERN CLASSIFICATION OF INSECTS.
Fabricius added a great number of genera, and distributed the order into ten divisons according to the form of the antenne. The Zarsal System of Olivier has, however, been almost universally adopted, chiefly in consequence of Latreille having employed it in his numerous works. It would appear, however, from the observations of the last- named author, that he has always doubted whether its rigid adoption might not be inconsistent with nature. Thus, in several of his most valuable works, and especially in the Genera Crustaceorum, &c., we find him questioning the series of Coleoptera founded thereon ; adding, “ Coleoptera palpis sex, robustiora, agiliora, primates ordinis*; rhyn- cophora e contra instrumentorum cibariorum exiguitate, ore ros- trato, debiliores, Mammalia edentata hic quodam modo emulantes. Articulorum tarsorum progressio numerica decrescens in methodo naturali non admittenda.” (Vol.i. p.172.). In his Considérations Générales, he has entered more fully into this subject, discussing the distributions resulting from the employment of the antenne, tarsi, and trophi, as leading characters, again giving the preference to the latter, but admitting his inability to propose a natural system, and therefore adopting the tarsal system.
Against this system it may be advanced, that by rigidly employing the numerical gradations of the tarsal joints, the real affinities of cer- tain insects (such as the Pselaphide and Clambus, above mentioned) are disturbed. It appears to me, however, that these ought rather to be regarded as exceptions, such as constantly oppose the adoption of every rule, especially when founded upon a single character. I do not, therefore, with Mr. MacLeay +, regard these instances as affording in themselves a sufficient ground for overthrowing the tarsal system. But another and more forcible objection has been raised by Mr. MacLeay, namely, that the Tetramerous Beetles are, in fact, Pen- tamerous, and that the Trimerous are Tetramerous; each of these groups possessing a minute joint, observable between the two lobes of the supposed penultimate joint. It appears to me, however, that this objection can only be maintained against the names of these groups, and not against the groups themselves, which, as it seems to me, are
* Mr. Kirby more recently, on account of their perfection and symmetry of general structure, the lightness and velocity of their motions, and brilliancy of colours, has regarded the Cicindelide as the analogues of the Lion amongst quadru- peds, and as the typical and most perfect group of Coleopterous insects,
+ Linn. Trans. vol. 15.
COLEOPTERA. 39
characterised by a strongly marked peculiarity of structure which it is impossible to overlook. * Mr. MacLeay is, however, of a different opinion ; and, rejecting the tarsal system entirely, has proposed a division of the Coleoptera, founded upon their larva state, of which he describes the five following types :—
1. A carnivorous Hexapod larva, with an elongated linear and flattened body, having a large head armed with two sharp falciform mandibles, and furnished with six granular eyes on each side. Ex- amples, Carabus, Dyticus. [ Analogy with Scolopendra. ]
2. An herbivorous Hexapod larva, with along and almost cylindri- cal body, so fashioned that the posterior extremity being curved under the breast, the animal, when at rest, necessarily lies like an Iulus on its side. Examples, Petalocerous Larve. [Analogy with Iulus. ]
3. An Apod larva, having scarcely the rudiments of antenna, but furnished, instead of feet, with flat fleshy tubercles, which, when con- tinued along the back and belly, give the animal a facility of moving in whatever way it may be placed. Examples, Curculio or Cerambyx. [Analogy with Vermes.]
4, An Hexapod and distinctly antenniferous larva, with a subovate rather conical body, of which the second segment is longer and of a different form from the others, so as to give the appearance of a thorax. Examples, Coccinella or Chrysomela. [Analogy with Pediculus. ]
5. An Hexapod antenniferous larva, of oblong form, having, like the former, vestiges of a thorax, besides two or more articulated or un- articulated appendages to the last segment of the abdomen. Example, Meloe. [Analogy with Podura. ]
«Each Coleopterous larva, I have observed, may be assimilated to one or other of these types, which it is scarcely possible to look at without being reminded of the primary types of the Ametabola. Indeed, it occurred to me almost immediately, that I had Chilopodi- form, Chilognathiform, Apod or Vermiform, and Anopluriform larve, with a fifth form (Thysanuriform), of which, even now, I know little,
* Several French entomologists (Solier, Chevrolat, Brullé) have very recently rediscovered this minute joint, to which they have attached as much importance as Mr, MacLeay. The advantages resulting from the discovery are not very apparent when the Longicorn Beetles are separated from the other Tetramera, and brought into the Pentamera, as proposed by Mr. Brullé. M. Solier also (Ann. Soc. Ent. Fr. 1834, pp. xlii.), considers that all the Coleoptera are in fact Pentamerous.
pd 4
40 MODERN CLASSIFICATION OF INSECTS.
except from the examination of two or three collected by myself, together with the marvellous descriptions of Goedart and De Geer.” (Hore Entomol. vol. ii. p.422.) Subsequently the same author states, that “ the types of Chilopodiform larvee are those carnivorous insects with four maxillary palpi, forming two groups, Carabus and Cicindela Linn., and Hydrocanthari Latr. To the same tribe, Hydro- philus Geoffr. ought to be ascribed, from which, by Spheridium, we enter among the Chilognathiform larvae, the type of which is ap- parently Scarabeus Linn. The tendency of Chilognathiform larve is herbivorous, having among them Lucanus, Byrrhus, Ptinus, Hister, Elater, Buprestis, and part of Tenebrio and Dermestes. By means of Bostrichus Geoffr. we probably quit the Chilognathiform for the Apod or Vermiform larve, including Bruchus, Curculio, Attelabus, Cerambyx, Leptura, Necydalis, and part of Tenebrio and Dermestes. With the help of Donacia (nearly allied to Leptura), we quit this tribe for the Anopluriform larve, including Chrysomela, Cryptoce- phalus, Coccinella, Cassida, and Hispa. These last lead to the Thysanuriform larva, including Meloe and Mordella, with many Heteromera Latr., and perhaps Clerus. I suspect that some of the Malacoderma Latr., find a place in this group, but certainly Staphy- linus Linn., reconducts us from these insects to the Chilopodiform tribe of larva. Some groups I dare scarcely venture on, such as Melasomes, Taxicornes, and Stenelytres Latr.” We are then informed of the important fact, that Pentamerous insects chiefly belong to Chi- lopodiform or Chilognathiform larve; that Apod larve become Tetramerous ; Anopluriform larve either Tetramerous or Trimerous ; and Thysanuriform larvee either Heteromerous or Pentamerous. * In the Annulosa Javanica, Mr. MacLeay declared his views relative to the Chilopodiform type or tribe as follows : —
Ex, Typic. Families.
1. Normal group, consisting of Cicindelide. insects haying linear or 1. G oe Carabide.
‘setaceous antenna, with (*° 7POPPPHAGA. Carabus. < Harpalide. ‘the outer process of the Scaritide,
maxille palpiform. Brachinide. (ADEPHAGA Clairville.) J 2. Hypraprruaca., Dyticus. ei yticide.
* Hore Ent. p. 464, 465.
COLEOPTERA, 41
Ex. Typic. Families. Heteroceridz ? Parnide. (3. Purtuypriva. Hydrophilus.~ Helophoride. Hydrophilidz. Spheeridiide ? Seaphidiide.
2. Aberrant group, consisting of insects having their antenne clavate, or at
‘ bat Silphide. elem tee src 4, Necrorpnaca. Silpha. Nitidulide. the external lobe of the ae i maxill is not palpiform, Rise a
[RYPOPHAGA Steph. } Pe aaa Stenide. 5. Bracuetyrra. Staphylinus. 2 Omalide. Pselaphide. Tachyporide.
The circularity of this tribe is supposed to be effected by the ap- proximation of Lesteva amongst the Brachelytra to the Carabidae. It must be borne in mind that the analogies existing between these five types of Coleopterous larvee and the five groups into which Mr. MacLeay divides his class Ametabola, constitute Mr. MacLeay’s chief argument in support of their correctness.* The above is the sub- stance of Mr. MacLeay’s hitherto published views upon this subject, whence the impossibility will be seen of following them out through the entire order, and which is indeed the more increased from our great ignorance of the larvae of many important groups. We have seen also that Mr. MacLeay himself admits the existence of a variation in the tarsal structure concurrent with the variation in the form of the larva, a circumstance dependent, as it seems to me, upon the principle that modifications of the preparatory states of an insect ought merely to be regarded as indications of corresponding pe- culiarities in the final state, the former modifications being subordinate to those observed in the imago, having in fact been undergone with a direct view to the perfection of the insect. We might indeed carry the subject still further. Thus, whilst the intimate connexion ex- isting throughout the whole of the Tetramerous Beetles cannot be denied, yet Cerambyx has a subvermiform and Chrysomela an
* Tt is to be observed, that Mr. MacLeay’s views respecting the Ametabola have been adopted by no subsequent author, and that Messrs. Kirby and Spence (who regard Mr. MacLeay’s proposed distribution as requiring much more consider- ation and investigation than have hitherto been given to it— /ntrod. iii. 161.) have pointed out numerous other and equally striking analogies’ existing between the Coleopterous larvae and the Crustaceous and Arachnidous classes (Introd. iii. 166.)
49 MODERN CLASSIFICATION OF INSECTS.
anopluriform larvee ; whilst the latter and Coccinella (Mr. MacLeay’s two examples of the Anopluriform stirps), although agreeing in the larvee, are totally different in the habits and in the structure of the tarsi of the imago. |
For these and other reasons which want of space prevents me from here detailing, but which I have embodied in a paper read before the Entomological Society, I have not considered it advisable in this work to adopt Mr. MacLeay’s views until they shall be more fully developed *, and shall therefore follow (with proper restrictions
* Mr. Stephens has endeavoured to adopt MacLeay’s views in the distribution of the Coleoptera (Catalogue of Brit. Ins. p. viii.), and it cannot be denied that some of the relations thus elucidated are very strong affinities ; others, however, appear equally slight, not being more than distant analogies. It is to be regretted, that want of space has prevented him in his Illustrations from developing his own views so fully as could have been wished. I need not do more than mention the classifications proposed by Mr. Newrnan (Ent. Mag. No. 9.), and by M. Laporte (Etudes Ent. No 1.), these authors not having thought fit to detail the principles upon which their classifications (which certainly do not appear to be very consistent with nature) are founded. Mr. Kirby, however, in his Fauna Boreali- Americana (published subsequently to the preparation of the following pages relative to the Coleoptera), has proposed various important modifications in the general classifi- cation of the order, founded upon and accompanied by observations of their affinities, and of which it will be useful in this place to give a slight review. Many of these are of too great value to be overlooked; but not a few appear to me to be un- founded in nature. Rejecting the Tarsal System of' Latreille, by a too great reliance whereon “ groups evidently nearly related are scattered far and wide through his various sections,” and also that of Mr. Macleay, which he considers impossible to adopt through the “ mazy labyrinth” of nature, he: regards it as impossible ‘* either to conceive or delineate it so as to maintain all its connections undisturbed and unbroken. We must do it in a series, which can only be a series of mutilations and dislocations.” (Pref. p. Xxiii.—xxv. )
Commencing with Cicindela, and passing through the remainder of the Adephaga (Carabus, Dyticus, and Gyrinus Linn.), he next enters into various anatomical arguments relative to the respective relations of the Brachelytra with the Adephaga (Latreille’s method, but rejecting Latreille’s location of the Buprestide, &c. ), and of the Philhydrida with the same group (MacLeay’s method), from which he con- siders that the Brachelytra by means of Lesteva (analogous to Lebia), and Stenus (analogous to Cicindela), are intermediate between the terrestrial Adephaga and the Necrophaga, whence he passes to the Philwidrida, which bear a stronger relation to the aquatic Adephaga, and thus a circular distribution of these groups is com- pleted. In this distribution the groups Geodephaga and Hydradephaga are made subsections, whilst the genus Necrophorus is raised to the rank of a section, equivalent to the Adephaga, Brachelytra, &c., under the name of Entaphia, a step surely not adyiseable. The Byrrhide are introduced amongst the Necrophaga, and it is.from this group that we are instructed to approach the Lamellicorn beetles by means of the Histerida. The Spheridiide are also regarded as another transition
COLEOPTERA. 43
and allowance for occasional exceptions), the Tarsal System, which,
from being in general use and of easy application for every practical
purpose, is superior to any of the other modes of classification yet
proposed.
The Coleoptera are therefore divisible into the four following
sections : —
1. Pentamera, in which all the tarsi are 5-jointed, the fourth being of ordinary size.
2.. Heteromera, in which the four anterior tarsi are 5-jointed, and the two posterior 4-jointed.
from the Philhydrida to the Histeride, and as consequently completing the circle of the Chilopodomorpha of MacLeay, Hister being an osculant group. Passing the Histeride, which he considers to be as much allied to the Copride as to the Lucanide, he proceeds through the Lamellieorns, and returns to the Histeride to take up the Lucanide, some of which (Passali, &c.) appear to diverge by Bostrichus to the Weevils. He then takes up, without indicating their affinities, the Sternoxi (Elater and Buprestis), which he regards as entitled to a station somewhere between the Ligniyorous Lamellicorns and the Capricorns; and, after simply hinting at the relation of the Sternoxi with the Malacoderma by means of Cebrio, he considers that there are two routes,—one from Lucanus to the Capricorns (through Cucujus, Pytho, and Trogosita), and thence directly to the Eupoda and Chrysomela; the other from Passalus through the G. Phrenapates K. Sinodendron, F., and an osculant tribe of timber-borers (Xylotrypa K.), including Ptinus L. (out of the direct line), Scolytus, Hylesinus, Bostrichus, &e., directly to the Weevils. This arrangement “ keeps together the great body of Lignivorous Coleoptera.” By means, moreover, of Anthribus and Clythra a connection is regarded as established between the Weeyils and the Cyclica Latr. The Coccinellide next follow, but their natural place is not yet satisfactorily ascertained. To these succeed the Heteromera in their ordinary course, merely “ because one of that subdivision, Meloe, appears evidently to be one of the links that connect the Coleoptera with the Orthoptera,” — “ which is proved not only by its galeate maxilla, but by its elytra lapping over each other, and its vertical and often inflexed head, in this respect re-
>
sembling the Blattina.” We are lastly to retrace our steps to take a fresh route from the Sternoxi by Cebrio to the Malacoderma, which, “in the soft substance of their elytra, and their reticulations in some species of Lycus,” seem to make an additional approach to the Orthoptera ; these relations being regarded as of higher value than that of Staphylinus and Forficula, adopted by Mr. Stephens for the like purpose.
It is admitted, that there are numerous links wanting to complete many of these proposed passages ; and it is also to be observed that it is only upon a consideration of the perfect state that these views have been founded ; but at the same time it will be conceded, that if the difficult task of arranging the Coleoptera upon a review of the relations of the various groups be our object, these suggestions of Mr. Kirby will be entitled to great weight, from the evident justice upon which so many of them haye been made.
44 MODERN CLASSIFICATION OF INSECTS.
3. Pseudotetramera* (or Subpentamera, Tetramera Latreille, Crypto- pentamera Burm.), in which the tarsi are 5-jointed, but the fourth joint is exceedingly diminutive, and concealed between the lobes of the preceding.
4. Pseudotrimera (or Subtetramera, Tetramera Laér., Cryptotetramera Burm.), in which the tarsi are 4-jointed, the third jomt being very diminutive, and concealed between the lobes of the pre- ceding.
Sect. i. PENTAMERA.
It is observable that although Linneus and Latreille considered very different groups as entitled to precedence amongst Beetles, by both Pentamerous insects were selected as fitted to commence the classi- fication of the order; the former, probably influenced by their su- perior size and strength, employing the giant Lamellicorns for that purpose; and the latter, in consequence of the superior developement of the manducatory organs and their agility, together with their internal organization, giving the precedence to the predaceous Tyger-beetlest and Carabi. Mr. Macleay, indeed, does not consider the latter in- sects as entitled to peculiar rank on these grounds; and, in fact, it only becomes a matter of importance with what insects we ought to commence our arrangements, when we have ascertained what are the precise animals composing the group immediately connected with or preceding them in a natural system, and by means of which the most easy passage between the two groups can be effected.
The section Pentamera seems to be divisible into two tribes, cor- responding with Mr. MacLeay’s groups Chilopodomorpha and Chi- lognathomorpha. I retain them, however, as merely provisional.
The tribe Chilopodomorpha, in like manner, seems to be divisible into two subtribes, viz. : —
1. The Adephaga Clairville. 2. The Rypophaga Stephens.
The former, constituting the more typical portion, is again divisible into
* J had elsewhere proposed the terms Subtetramera and Subtrimera for Latreille’s Tetramera and Trimera; but as the preposition sub implies a deduction from the quality of the word to which it is prefixed, and as these two groups require, in effect, a designation implying a slight addition to the character originally imputed to them, I have proposed for them the terms of Pseudotetramera (or Subpentamera), and Pseudotrimera (or Subtetramera).
+ See ante, p. 11. note *, p. 38. and Dahlbom, Clavis Noy. Hym. Syst. Pra- mon. p. iy.
COLEOPTERA. — ADEPHAGA. 45
two stirpes, Geodephaga and Hydradephaga, the insects being very predaceous in their habits; and the latter, or the aberrant portion, composed of three stirpes, Philhydrida, Necrophaga, and Brachelytra, and feeding upon dead animal or vegetable matter.
The circularity and natural progression of this tribe is effected, according to Mr. MacLeay, in the following manner : — Commencing with the Geodephaga we arrive at the Hydradephaga by means of Omophron, Hydrous in like manner connecting the latter stirps with the Philhydrida; whence, by the assistance of Helophorus, we arrive at the Necrophaga, which are united with the Brachelytra by means of Micropeplus; Lesteva, from its analogy with Lebia, and Stenus, from its analogy with Cicindela (as suggested by Mr. Kirby), form- ing the links which complete the circle, by uniting the latter stirps with the Geodephaga.
In the following pages I have adhered to this arrangement, although by introducing the Brachelytra at the end of the Necrophaga, the series of affinity between Dermestes, Anthrenus, Byrrhus, Hister, and Lucanus is broken.
The subtribe ADEPHAGA is distinguished by the outer lobe of the maxille being distinct and articulated, or palpiform*, so that these insects have been ordinarily stated to possess six palpi, one pair being attached to the lower lip, and a pair to each of the maxilla, as though the gluttony of these insects required an additional organ. The inner edge of their lower jaws is also armed with strong spines along its entire length, the extremity being also terminated by an acute hook. The antennz are long and slender, the legs of more than ordinary length, and the anterior tarsi generally dilated in the males. The variations occurring in this last-mentioned character have been employed by the French entomologists to distinguish several of the groups com- prising this subtribe, but it is to Mr. Kirby that we are indebted for the most satisfactory remarks upon them. (Faun. Bor. Amer. p. |.) In this respect, the Adephaga may be divided into — |. Those without dilatation; 2. Those having the two anterior tarsi dilated; 3. Those which have the four anterior tarsi dilated. Instances of simple tarsi in both sexes occur in Manticora, the Scaritides, many Carabides, and some Brachinides, with Haliplus; on the other hand, in some Cicin- delidz, Colliuris, &c., and in certain Brachinides, the tarsi are dilated
* Tt is to Knoch (and not to Latreille nor MacLeay) that we are indebted for the first intimation of the true nature of this organ.
46 MODERN CLASSIFICATION OF INSECTS.
in both sexes. The Bimani (Dimani A.), or those with the two an- terior tarsi of the males dilated, are most numerous amongst the Geodephaga, and exist also in Hydaticus and Acilius, whilst the Quadrimani are more numerous in the Hydradephaga, and also in the subfamily Harpalides. The number of dilated joints varies from one to four, the basal joint exhibiting the greatest degree of dilatation. There is also considerable variation in the form of the dilated joints, and which (as in the Patellimani) has been employed to characterise divisions. An equally important character exists in the lining of the sole of these dilated joints, which Mr. Kirby describes (without being apparently acquainted with the observations of Messrs. L. Dufour and Audouin, subsequently alluded to), and divides the Adephaga into three sections thereupon, namely : —
1. SarroTHROPODA, or those with the dilated joints clothed beneath with a dense brush of stiff hairs: Cicindelidae, Carabides, Chlanius and its allies. (See also Aud. and Brullé, Hist. Nat. Ins. Col. vol. i. p. 395. and 444.) Others have only a few hairs at the sides of the joints of the tarsi.
2. CystTopropaA, or those with the foot-cushions formed of little mem- braneous vesicles or cysts variously arranged : Brachinus, Agonum, and various Harpalides. In Helluo and Anthia the dilated joints are furnished with lateral brushes, and in the middle with vesicles.
3. Pyxrptopopa, or those with the foot-cushions formed of pedun- culated cups or suckers: Dyticide.
In the figures accompanying the various groups of Adephaga, I have illustrated the variations in these foot-cushions, which will thence be seen to be more numerous than stated by Mr. Kirby.
These insects are extremely active, and eminently predaceous, devouring other insects with great avidity, and occasionally not even sparing their own species. This subtribe is divisible into two* stirpes, according to the element in which the insects reside —
The Ist, Geodephaga, being terrestrial
caress seh The 2d, Hydradephaga, being aquatic fin reir habits
The former very far exceeds the latter in the number of its species.
* Mr. Kirby (Faun. Bor. Am. p. 6.) has proposed another stirps, Hygradephaga, for those Carabideous insects delighting in humid places (the Acutipalpi,Subulipalpi, and Elaphridz), constituting an intermediate or transition group between the Geode- phaga and Hydradephaga. I cannot, however, find any decisive characters for such a separation of insects so closely allied together as the Acutipalpi, &c., and the other Geodephaga.
COLEOPTERA. — CICINDELIDZ. 4.7
The first stirps, GEODEPHAGA, or predaceous Land-beetles, have the legs long and formed for running, the four posterior being placed at equal distances apart, and capable both of vertical and horizontal motion ; the jaws are long, horny, and curved, not being concealed by the upper lip; the terminal internal lobe of the maxilla is straight, except at the tip, which is hooked; the antenne are long and slender ; the body is generally oblong, seldom oval; and the eyes prominent. This is one of the most extensive groups of insects; Dejean enume- rating 2500 species, and Stephens describing about 450 as inhabitants of this country. From their numbers, therefore, they must be emi- nently serviceable in keeping in check the tribes of other insects which, if undisturbed, might become noxious. They are chiefly found beneath stones, under clods of earth, &c.
This stirps corresponds with the Linnean genera Cicindela and Carabus, which appear to constitute the two chief divisions of the stirps. Mr. MacLeay has indeed divided the Carabi into four groups, and Mr. Stephens into six, each of which is considered of equal rank with Cicindela. Messrs. Kirby and Spence, however, follow Linnzus and Latreille, adopting only two chief divisions, proposing for Ci- cindela the name of Eupterina (in allusion to their powers of flight), and for Carabus that of Eupodina (from their power of running). I however prefer regarding these two divisions as families, and of equal rank : —
Ist, Cicindelide, or Tiger-beetles (maxille armed with a moveable claw).
2d, Carabide, or Ground-beetles (maxilla not armed with a move- able claw ).
Tue first family, CicinDELIDA Leach*, is distinguished by the terminal hook of the maxilla being articulated at its base (fig. 1. 3.):
* Bisriocr. RererENCEs TO THE CICINDELIDA, Dejean. Spécies Général des Coléoptéres, vol. i. and App. to vol. ii. and v. Paris, 1825—1831. Latreille and Dejean. Coléoptéres d’Europe, 8vo. Paris, 1822. 7 Van der Linden. Memoir upon the Inseets of Java, part 1. Bruxelles. 4to. 1829. MacLeay. Annulosa Javanica, part 1. 4to. London, 1825. Laporte de Castelnau. Etudes Entomologiques. 8yo. Paris, 1834. — Memoirs in Silbermann Rey. Entomol. tom. i. and ii. — Hist. Nat. des Anim. Articulés.
Paris, 8vo. 1837. ‘ F . , J ' x 08 a OA tateert etlilign of Hew lubrotuc fa mr é ite Sines ca TS 435 ile a Q o On Ppt es a
2
48 MODERN CLASSFICATION OF INSECTS.
the head is large, and broader than the thorax (fig. 1.1. Cicindela campestris, the common green Tiger-beetle): the eyes are very large
Thunberg. Museum Upsaliense, p. 51,52. (Cicind. Cap. Bon. Sp.)
Gory. Centurie, &c. Ann, Soc. Ent. France. t. ii.
Audouin and Brullé. Histoire Naturelle des Insectes Coléopteres, vol. i.
Lund. Trycondyla aptera; Schrift. Natur. Gesellsch, vol. i. Copenhagen, 1793.
Mathes. Mem. Soc. Imp. Natur. Mose. tom. ii, p. 311. (Cicindela gracilis. )
Kirby. Century, Linn. Trans. vol. xii.
Perty. Delect. An. art. Brasilia, ut supra.
Griffith. Animal Kingdom. Insects. Pl. 29.
Fischer. Entomogr. de la Russie. 4to.
Say. In Trans. Amer. Phil. Soe. vol. i. New series, 1818 (containing a Monogr. on the N. Am. Cicindele); and vol. iv. — Ditto, in Journ. Acad. Nat. Se. vol. i. 1817, vol. iii. 1823. Philad.—Amer. Entomol. vol. iii. 8vo. 1824—28.
Klug, in Wiegmann’s Archiv. vol. i. (Stenocera, Pogonostoma.) — Ditto, in Mem. Madagascar Col. in Kon. Akad. Wissensch. Abh. 1832. — Ditto, in Ehren- berg’s Symbola Physicee. — Jahrbucher for 1834. Berlin. 8vo.
Gory and Laporte. Hist. Nat. Coleopt. (Stenocera Monogr.)
Gistl, Syst. Insectorum Monachi. 1837. Part 1. containing a Synopsis of the Species. 8vo.
Guérin. Mag. Zool. (various detached species. )
Brullé, in Silberm. Rev. Ent. Nos. 9. and 18.
Mannerheim, in No. 2. Bullet. Soc. Imper. Nat. Moscow, 1837. (Including the Description of a new Genus, Oxygonia, from Columbia. )
Erichson. Kafer Brandenburg. part 1. Berlin, 8vo. 1837.
Westwood, in Ann. Sciences Nat. vol. xxii. (Mem. Hist. Nat. Cicindel.) — Ditto, in Zool. Journ. No. 17. (On Ctenostoma. )— Ditto, in Jardine’s Mag. Zool. and Bot. vol. i. (Comprising the Description of a new Genus, Distipsidera. )
A. A. Gould, in Boston Journ. Nat. Hist. 8vo. No. 1. 1834. (Cicind. of Massa- chusetts, with 1 pl.)
Eschscholtz. Zool. Atlas. fol. Berl. 1831. (Genus Omus.)
Kollar, in Annal. Weiner Mus. der Naturgesch, b. 1. 1837.
And the general works of Fabricius, Olivier, Herbst. &c.
COLEOPTERA. — CICINDELIDZ. 49
and prominent: the mandibles (fig. 1.2. the head,) very large, acute, and armed with several strong teeth; the palpi are not longer than the mandibles: the lower lip is concealed by the broad mentum (jig. 1. 4.); and the labial palpi (which are three-jointed, and appear to arise from a moveable base, and are densely clothed with white hairs), as well as the legs, tarsi, and antenne, are long and slender: the anterior tibia are not notched on their inner side, and the penultimate segment of the abdomen is often notched in the males: the anterior tarsi of the males are often broader than in the females (fig. 1. 5., anterior tarsus Cicindela ¢ — 1.6. ditto? ). Of these characters, that afforded by the slight development of the labium is, perhaps, the most valuable, although that of the articulated terminal maxillary hook has been chiefly relied upon; but M. Audouin has recently observed, not only that in the genus Ctenostoma the hook is entirely wanting —as, indeed, I had previously noticed by dissection (fig. 1. 10.) —but also that, in the Carabideous genus Trigonodactyla, the hook is distinctly articu- lated. In the genus Stenocera Brullé (Pogonostoma Alug), the hook is also wanting.
The Cicindelide generally frequent hot sandy districts, where they may be observed flying in the sunshine with great velocity; whence Messrs. Kirby and Spence have termed them Eupterina. Their flight is, however, of short duration ; but they immediately take wing again on our approaching them. From the brilliancy of their colours they have been termed sparklers, whilst the name of Tiger-beetles has also been conferred upon them, not only from the spots and stripes with which they are ornamented, but also from their savage propensities, preying upon every insect which they can overcome, their extreme agility at the same time rendering escape impossible to their luckless prey. In the warmer climates of the New World some of the species of Cicindela, Iresia, Euprosopus, &c., appear to lose some of the habits of their congeners of more moderate climes; since it is upon the leaves and trunks of trees that they are generally found; where, like their terrestrial relatives, they carry on a ferocious war against other insects; flying from leaf to leaf with the agility of flies, and darting upon their prey with great quickness: some, however, make less use of their wings, running rather than flying, and seeking the most arid spots of ground for their abode.
The only larvae hitherto observed in this family are those of several species of the genus Cicindela. Geoffroy (Hist. des Insectes, vol. i.),
E
50 MODERN CLASSIFICATION OF INSECTS.
Desmarets (Bull. Soc. Phil., 1801 —1805, No.190.), Ratzeburg (Forst. Ins. 1837. pl. 1.), Latreille (Nowe. Dict. d Hist. Nat. 2d ed., &c.), and Messrs. Kirby and Spence (Jntrod., vol. ili. pl. 17. f. 13.), have given accounts and figures of these larvee; and in the Annales des Sciences Naturelles for March 1831, ina Memoir by myself, containing a detailed account of the habits of the insects of this family, both in the pre- paratory and perfect states, I have minutely described the structure of these larvee, which in Cicindela campestris are long, subcylindric, and of a whitish colour and of a fleshy consistence (fig. 1. 7.); the head (jig. 1. 8. head and prothorax seen beneath) is very large, obscurely coloured, horny and hollow above, with a pair of large and powerful, sickle-shaped jaws, two short 4-jointed antenne, and, as it appeared to me, six small simple eyes. The maxilla and labium are re- presented at jiy. 1.9. The first segment of the body is alse horny, flat, and of a semilunar form, being, as is also each of the two following segments, furnished with a pair of strong articulated legs. The eighth segment of the body is the largest, and is armed with a pair of bent hooks placed upon a fleshy retractile tubercle. The four remaining segments are narrowed and terminated by a conical anal process. These larvae burrow cylindric retreats in the earth to the depth of a foot or more, employing their legs and jaws in loosening the particles of sand and earth which they, carry to the surface upon their broad saucer-like head, ascending by the assistance of the two hooks upon the back, somewhat after the fashion of a sweep going up a chimney. Having completed this burrow, they station themselves by means of their legs and dorsal hooks, at its mouth, their large flat- tened head and first segment fitting the hole; here they lay in wait for such insects as may be crawling about, seizing them with their jaws, by suddenly throwing the head backwards and then dragging them to the bottom of the burrow where they may generally be found in the position represented in fig 1.7. At the least approach of danger they also immediately slide down to the bottom of their retreats. When ready to assume the pupa state, it is stated that they close the aperture of the cells. Mr. Curtis (Brit. Ent. No. 1. 2d ed.) has suggested that the hooks on the back of the larva “may, by being brought into contact with the head, enable it to hold its prey.” This, however, I conceive can scarcely be correct, since, at the period when the larva seizes its prey, its hooks are employed like anchors, to sup- port it at the mouth of the burrow. Moreover, in the larve which I
COLEOPTERA. — CICINDELID®. 51
kept alive, I never saw these organs employed in the manner suggested by Mr. Curtis, although I repeatedly saw them seize their prey. Nothing is known respecting the transformation of the exotic species of this family; Latreille, however, considers that they are similar to those of the European ones, neither have the pupz been observed. Respecting the name given to this group of insects, it is to be ob- served that our Latin dictionaries, and Cowper’s pretty poem, Ad Ci- cindelam, supported by the authority of Mouffet ( Theatr. Ins. cap. xv. De Cicindela), and some others of the old entomologists, would lead the entomological tyro to suppose that the Glow-worm was the insect under consideration. Such, however, is not the case; Linnzeus, who at first described that insect as a species of Cantharis, having with evident impropriety applied the name of Cicindela to the Tiger- beetles. Geoffroy, however, erroneously considering the latter as not even generically distinct from the Linnean Carabi (which he regarded as the Buprestis of the ancients), improperly gave the name of Ci- cindela to the Soldier-beetles ( Telephorus, or the Cantharides of Linneus), applying at the same time the name of Cantharis to the Blister-fly (Canth. vesicatoria), and to the Glow-worm, the old Greek name of Lampyris. In the subsequent editions of the Systema Nature, Linnzus adopted the latter alteration, and the Glow-worm is now known by the name of Lampyris; but for the Tiger-beetles, which he correctly considered as distinct from the Carabi, he still re- tained the name of Cicindela, which, notwithstanding its evident impropriety, has been universally adopted. The Soldier-beetles, he named Cantharis, and the real Cantharis of the shops or Blister-fly he called a Meloe, whilst to complete the confusion Fabricius con- sidered the Blister-fly as distinct from Meloe, and gave it the name of Lytta. Such is one of the instances (of which it is to be regretted that too many are to be found in the works of the earlier entomo- logical nomenclaturists) of that want of unity which has resulted from the great deficiency of some fixed rules of nomenclature in Natural History. In the instances before us, this confusion still exists ; for although the two insects which gave rise to it, the Tiger-beetle and the Glow-worm, have at length by general scientific usage acquired a title to the names of Cicindela and Lampyris, modern entomologists are still at issue respecting the names of the Blister-fly and the Soldier-beetle, the former being called by some of the most recent authors (Dejean, Gyllenhall, &c.,) Lytta; and by others (Latreille, E 2
52 MODERN CLASSIFICATION OF INSECTS.
Stephens, &c.) Cantharis; whilst the Soldier-beetles are by some writers termed Telephorus, and by others Cantharis.
The number of insects belonging to this family scarcely exceeds 250 (or one tenth of the family Carabide*); of these by far the greater portion are exotic, and the Baron Dejean has described about 200 of them as belonging to the genus Cicindela. They are generally of a moderate size, the majority varying from half an inch to an inch in length: none are beneath one third of an inch long; their ferms, as well as the disposition of their colouring, are generally very elegant.
The affinities of these insects with the Carabidae are very varied and interesting ; thus whilst Manticora and Anthia (as indicated by Latreille, Col. d Eur. No. 1.) are strikingly allied together, Elaphrus seems not less nearly to approach Cicindela. In like manner Colliuris Latr. closely resembles some of the genera of the Brachinides, espe- cially Colliuris De G. (Casnonia Laér.), and Agra (see Kirby, Faun. Amer. Bor. p.5.); and M. Audouin has lately placed Trigono- dactyla (one cf the Brachinideous genera) at the head of the Cara- bide, from the identity of structure of its maxilla and those of the Cicindelide. Mr. MacLeay also appears to consider that the Cicindelidee and typical Carabide are allied, from the identity in the unnotched anterior tibia. From these varied points of affinity of the Cicindelidz with the various divisions of the Carabidz, it is impossible to arrange them in a series, or to describe them in the order in which they are arranged by nature. A more distant relation of the Cicindelide with the Brachelytra, by means of Stenus has been suggested by Mr. Kirby (Fauna Bor. Amer.).
Several modes of distribution of the genera of this family have been proposed by Latreille, MacLeay, Brullé, &c.+ ; but these authors have chiefly selected isolated organs, and as the genera are entirely
* Tt is a remarkable circumstance in the distribution of insects, that whilst some families abound in species, such, for instance, as the Carabidae, &¢. others intimately allied to them, as the Cicindelide and Dyticide, possess but very few. The same observation may be made of generic groups: thus, Dejean enumerates 163 species of Carabus, whilst of the more widely distributed allied grovp, Calosoma, he only describes 28.
+ Mr. Kirby divides the family into Longicollia (Colliuris), Brevicollia (Cicin- dela), Fissicollia (Manticora), forming them into a cirele with some of the Brachi- nides, namely, Agra as the counterpart of Colliuris, Anthia of Manticora, and Calleida of Cicindela. These Brachinideous genera appear, however, to me to be in this manner unnaturally separated from their legitimate allies, the remainder of thee Brachinideous group.
COLEOPTERA. — CICINDELIDZ. 53
exotic, with the exception of Cicindela, which is the only British, and, indeed, European, genus, the discussion of these arrangements may be here dispensed with.
The species of the G. Cicindela are generally of a green or bronzed colour, ornamented with white spots and streaks upon the elytra. * The typical species C. campestris Zinn. the common green tyger-beetle (fig. 1.1.) is by far the most abundant species in this country, being generally distributed throughout England. It is rather more than half an inch Jong, of a deadened green colour, with several white spots on the elytra, varying in number from three to six; the upper lip also is white ; the female has also two additional spots, but of a dusky colour, near the base of the elytra. I have found this species as early as the middle of February at Coombe Wood, and have noticed, that when handled, it exhales a pleasing scent, something like that of roses. Messrs. Kirby and Spence have observed, respecting another British species (Cic. sylvatica Linn.), that it flies with a considerable hum- ming noise.
Another species of this genus, which has only recently been ascer- tained to be indigenous, is the true Cicindela hybrida of Linneus, respecting which great confusion exists in modern entomological works. See Stephens (Jl/lust. Brit. Ent. vol.i. pp. 8. 18.175. and vol. v. p. 366.); Curtis (Brit. Ent. No.1. second edition); Au- douin and Brullé (Hist. Nat. Ins. Coleopt. 1. livr.1.). Laporte (Sil- berm. Rev. Ent. No.7. and Hist. Nat. An. Artic.); Klug. (Zahr- bucher, 1834); Erichson (Kafer Mark Brandb.). I am therefore happy in being able to clear up the doubts which have been raised, by giving a figure from the specimen of this species, preserved in the Collection of Linnzeus, now in the possession of the Linnean Society. The insect attached to the label of Cie. hybrida, in the handwriting of Linneus himself, is that to which the French entomologists have assigned (but doubtingly) that name, and which Mr. Stephens has de- scribed under the name of Riparia. Moreover the Linnzan Cabinet does not contain a specimen of the species which Mr. Sowerby figured
* M. Audouin has taken occasion, in describing a beautiful species of Cicindela from the East Indies, ornamented with golden yellow spots, to make some obsery- ations upon the general uniformity and prevalence of particular colours in particular groups of animals, giving the Cicindelx as an example ; the species under descrip- tion of course forming an exception to their prevailing hue. — Guérin, Mag. Zool. Ins. pl. 18. ;
a
E J
54: MODERN CLASSIFICATION OF INSECTS.
under the name of Hybrida, and which has been described by the French authors under that of C. maritima.
The smallest British species is the C. Germanica Zinn., which is only five lines long, of a dark green colour with four small spots, and a terminal lunule of a white colour on the elytra. It has been found in great abundance at Blackgang Chine in the Isle of Wight, and elsewhere ; and instead of frequenting hot sandy places, like the other species, it is observed running amongst grass and low herbage in moist situations. It has also been observed, that it makes but little use of its wings: this, and some other circumstances, induced me to suggest the establishment of a subgenus (named Cylindera, from its cylindric neck), for the reception of it, and some allied species ; in the majority of its structural characters, however, it agrees with the common species.
Some of the exotic species of the genus Cicindela are generally found upon the leaves of trees (Westermann in Silberm. Rev. Ent. No. 3.; Lacordaire, Annales Sci. Nat., June, 1830, &c.). These species have been separated by M. Laporte, under the name of Odontocheila : they are of a narrower form than the other species, and have the upper lip more advanced and toothed; M. Brullé, moreover, having examined the structure of these tree insects, discovered that in some of them the anterior tarsi are channelled down the centre beneath, whilst some have a style or bristle at the tip of the internal max- illary palpi; but these characters, net being permanent throughout the group, he is induced to regard them as merely indicative of a sectional division, of which the Cic. cylindricollis, Cayennensis, &c. are examples. The Cic. nodicornis Dej. also belongs to the same group, but is distinguished by the curious structure of the antennze of the males, which have the basal joint terminated in a thick knob.
We are informed by M. Chevrolat, that the Mexican species, Cic. curvata, burrows in moist sand, whenever the wind blows or the sun is obscured, and that the natives macerate it in water or spirit of wine to prepare a medicinal liquid (St/berm. Rev. Ent., No. 6.). According to M. Westermann, the Cic. 6-pustulata, bicolor, and 20-punctata are found upon the young rice plants in the East Indies (Silb. Rev. Ent. No. 3.). Mr.W. W. Saunders has published an account of the first of these three species, as well as of another new species, which he has termed Cic. limosa, which he observed near Calcutta, and which are attracted by the light of lamps entering apartments,
COLEOPTERA. — CICINDELID®. 55
or even ships lying at anchor by night, thus totally differing from their European brethren (Zrans. Ent. Soc. London. No. 1.). M. Guerin has figured a remarkable species from Cochin China, in his Lconographie, in which the legs are excessively long and slender ; and M. Barthelemy has described and figured another curious species ( Cie. Audouinii) from the coast of Barbary, in which the last joints of the antenne are much thicker than the rest (Ann. Soc. Ent. France, 1835, pl. 17. f.1.). The species had, however, been previously de- scribed by Mr. Vigors in the Zool. Journ., under the name of C. Ritchii. {It forms the genus Laphra, in Dejean’s new catalogue.
The largest insectsin the family belong tothe genus Manticora, of which the type is the M. tuberculata De G. (maxillosa Fabr.). It is about two inches long, an inhabitant of the arid sandy plains of Southern Africa, entirely of a black colour, and destitute of wings, the elytra being soldered together: hence the insect is confined to the ground ; it runs with great agility, and secretes itself under stones. Mr. G. R. Waterhouse has described a second species of the genus (M. lati- pennis) brought from the interior of Southern Africa by Dr. Andrew Smith, by whom it was discovered in a clump of dead trees (Mag. Nat. Hist. Sept. 1837). Other genera (Ctenostoma, Aptema, Dromica, Platychile) are also apterous, or furnished only with rudiments of wings. The species of Megacephala, also owing to the great length of their legs, and their somewhat less developed wings, are more accustomed to running than flying than the other winged species of the family; and according to M. Lacordaire, Meg. sepulchralis makes no use of its wings, but runs with great agility through the grass growing in sandy situations in the forests of Brazil. This species emits a strong smell of roses, but which shortly after death becomes fetid and disagreeable.* Another Brazilian species of the same genus hides itself beneath dry dung in the burrows of the On- thophagi and Coprides, where it remains during the heat of the day, endeavouring to defend the mouth of its hole, if an attempt be made to drag it forth. If this be persisted in, the insect soon descends to the bottom of its burrow, whence it may be drawn in the same manner as the larva of our common species, by introducing a straw, which itimm ediately seizes, and will then allow itself to be drawn up forcibly, rather than
* This species differs so materially from the other species of the genus in its pro- duced labrum and short antenna, as to require a distinct subgenus for its reception.
E 4
56 MODERN CLASSIFICATION OF INSECTS.
let go its hold. According to Olivier and Lacordaire, the insects of this genus make a noise somewhat like a grasshopper, by rubbing the hind thighs against the margins of the elytra. M. Laporte has pub- lished a monograph upon this genus in Silbermann’s Revue Ento- mologique, No.7. Another of the exotic genera, Oxycheila De7., is distinguished by having the upper lip very long, pointed in front and notched at the sides. According to M. Lacordaire, the species of this genus fly more heavily and run slower than the Cicindele, and hide themselves under stones during the heat of the day: they also emit an acute sound when handled, by rubbing the posterior femora against the elytra, as does also the species of which the genus Hu- prosopus is composed. The upper lip is also greatly developed in my genus Distipsidera (fig. 1. 12.).
The genus Therates Latr. (Eurychile Bonelli) is remarkable in having the internal maxillary palpi rudimental (zg. 1.11.) and the tarsi with the penultimate joint dilated and heart-shaped (fig. 1. 13.). From this circumstance Bonelli (who had made the interesting observation, that in general those insects which creep upon the ground have simple tarsi, whilst in those which frequent plants they are more or less di- lated,) was led to suppose, that the Therates seeks its prey upon plants in the same manner as the Calosoma. I have, however, elsewhere shown, that in this family, at least, such an observation does not entirely prevail (Annales Sc. Nat.): the genera Euprosopus and Iresia, for instance, which are known to be tree insects, have the tarsi, even in the males, but slightly dilated, as is indeed the case in Calosoma, whilst some of the apterous, and consequently terrestrial species in the family (Tricondyla, Ctenostoma,) have the anterior tarsi in the males greatly dilated. The structure of the tarsi is very different between those insects in which the dilatation is merely sexual, and confined to the fore legs, and the true herbivorous Coleopterous insects, in which all the tarsi in both sexes are dilated and cushioned, and which is also the case even in the genus Collyris (fig. 1. 14. anterior tarsus, f . 1.15. ditto @.). Of the exotic genera, Collyris Fadr. (Colliuris Lat.) Tricondyla, Therates, Procephala, and Ctenostoma are distinguished by their very narrow form, and by their thorax, which is globose in the middle. Of the last named genus I have published a detailed account in the Zoological Journal, No. 17. The Brazilian genus Iresia is distinguished from all the other genera by having the terminal joint of the labial palpi longer than the penultimate. The
COLEOPTERA. — CARABIDA:. 57
genus Stenocera Brullé (Pogonostoma Klug.) is confined to Mada- gascar, and is remarkable for the great length of the antenne, palpi and legs, and the cylindrical body.
The second group of predacious land Beetles or the family Cara- BiD& Leach*, comprises the entire genus Carabus of Linneus with a
* Brisriocr. REFERENCES TO THE CARABID&.
Dejean. Spécies Général des Coléoptéres, vol. i—y. 8vo. Paris, 1825—1831. Dejean and Boisduval. Iconogr. Coléoptéres d’Europe, 2d edit. vol. i—iy. 1829. Latreille and Dejean. Coléoptéres d’Europe, Ist edit. Nos. 1,2, and 3. 1829.
Klug. Jahrbucher for 1834.— Ditto, in Entomologische Monogr.— Ditto, in Nova Acta. Acad. Curios. t. x. — Ditto, in Ehrenb. Symbole Physicee. — Ditto, in Descriptions of Col. of Madagascar.
Audouin and Brullé. Histoire Nat. des Ins. Coléopt. vols. i. and ii. 1834.
Laporte. Etudes Entomologiques. — Ditto, Hist. Nat. An, Articulés. — Ditto, in Annal. Soe. Ent. Fr. vol. 1.
MacLeay. Annulosa Javanica.
Clairville. Entomologie Helvetique.
Say, in Amer. Phil. Trans. vol. ii. new series, and vol. iv.
Paykull. Monogr. Caraborum Sueciee, 8vo. Upsal. 1790.
Bonelli. Observations Entomologiques. Turin, 1809. 4to.
Sturm. Deutchsland Fauna. 12mo.
Guérin. Iconographie du Régne Animal.
Gory. Centurie de Carabiques Nouveaux, Ann. Soc. Ent. de France, vol. i.
Gory and Laporte. Hist. Nat. et Teon, des Coléopt. (G. Eunostus, and Nycteis. )
Various detached Memoirs by Chevrolat, Laporte, and Brullé, in Silbermann’s Reyue
Entomologique. Faldermann, in Nouy. Mem. Soe. Mose. vol. iv. Zoubkoff, Bullet. Soc. Hist. Nat. Mosc. vol. i. 1829. Bassi, in Annal. Soc. Ent. de France, 1834. (G. Cardiomera, &e. ) Buquet, in ditto, for 1835. (Various new exotic Carabide). Dufour, in Ann. Général des Se. Phys. Bruxell. 1820. Palliardi, Beschr. zw. Decad. Neu. Carab. Vienna, 1825. 8vo. Eschscholtz, in Mem. Soc. Imp. des Natur. Moscow. t. vi. Rylands, in the ‘ Naturalist,” April, 1837. (On Amara. ) Mannerheim (Mem. sur quelques Genres et Espéces de Carab.), in Bull. Soc. Imp. Nat. Moscow, 1837. Part ii.
Chaudoir, in ditto, part iii. — Ditto, in Ann. Soe. Ent. de France. 1836.
Newman, in Entomol. Mag. Nos. 22. and 24. Various new genera and _ species
of exotie Carab. Guérin. Mag. de Zool. Figures of various detached species.
58 MODERN CLASSIFICATION OF INSECTS.
few of his species of Cicindela. This is a family of very great extent, 2250 species having been described by Dejean, which are contained in his own collection, and their are numbers which he does not possess. It is a family which has attracted much of the attention of modern entomologists, Clairville, Bonelli, Paykull, Dejean, Leach, Latreille, &c., being amongst those whose investigations have been directed to its elucidation; hence, the structure of the perfect insects has in a great degree become well known: few observations have, however, been made relative to the natural history of the various species.
This family is at once distinguished from the Tiger-beetles by the maxille terminating in a simple point without any articulation at
Gory, in ditto. (Monograph on Pamborus.)
Lequien, in ditto, (Monograph on Anthia. )
Westwood, in ditto. (Memoir on the Genus Ozena and its allies. )
Erichson. Kafer der Mark Brandenburg.
Cadolint. Enumeratio Carabicorum Ticinensium. 8vo. 1830.
Mannerheim. Monographia Pelophilarum in Hummell’s Essais Entom. part iii. ; and Descriptions of new Species of Carabide in ditto, pp. 4, 5, 6.
Eschscholtz. Zoologischer Atlas, fase. 2.
Griffith. Animal Kingdom. Part, Insects.
Fischer. Entomographie de la Russie. 4to.
Kirby, in Linn, Trans. y. 12. (Century), and y. 14. (Pseudomorpha and Catascopus. )
Zimmerman. Monographia Amaroidum. Berlin. 8vo. — Ditto, Monographie der Carabiden, Ist Stuck. Berlin, 1831. 8vo. — Ditto, Monograph on Amara in Gistl’s Faunus, Ist band, Ist heft. Munchen. 1832.
Germar. Coleopt. Nov. Spec.
Hoppe, in Noy. Act. Acad. Curios. vol. xii.
Knoch. Neue Beitrage. (On Cychrus. )
Adams. Mem. Soc. Nat. Moscow. vol. v.
Waterhouse. Monograph on Nothiophilus, Entomol. Magazine, vol. i.
Schilling, on Holoenemis (Pteroloma Gyll.) in Beitrage zur Entomologie, Ist heft.
Hrichson, on the same genus in Weigman’s Arch. (in which the genus is regarded as belonging to the Necrophaga).
Hope, in Synopsis Ins. Nepaul (Gray’s Zool. Mise. i.) — Ditto, in Zool. Soe. Trans. vol. i. — Ditto, in Trans. Ent. Soe. vol. i. pl. 1.
Babington, in Trans. Ent. Soe. vol. i, (On Dromius. )
Solier, in Ann. Soe. Ent. de France. 1834.— Ditto, Ditomus, Aptinus, &c. —
Ditto, 1837, various genera. — Ditto, 1835, various new species.
Kollar, in Annal. der Weiner Mus. der Naturgesch. b.i. 1837. (G. Damaster, near Cychrus; G. Panagzus. )
Schiodte. Memoir on Danish Amare in Kroger’s Naturhist. Tidsskrift.
And the Works of Linneus, Fabricius, Gyllenhall, Olivier, Stephens, Curtis, Marsham, §e.
COLEOPTERA. — CARABIDE. 59
the tip* (jig. 3. 2., 4. 14. 16. 18.). The lower lip is more or less de- veloped, generally square, and sometimes with a short tooth on each side in front+ (fig. 3. 4. 4. 19.). The mentum, or horny basal portion (fig. 3. 3. fig. 4. 8.19.) is advanced into corneous lobes at the sides, the centre of the emargination being either simple, or furnished with an entire or bifid tooth. ~The mentum in some few species (Enceladus, Silphomorpha Westw.) is soldered to the under side of the skull. The head (fig. 3.1. head of Omaseus melanarius) is generally narrower than the thorax, the eyes but slightly prominent, and the various parts of the mouth are considerably advanced in front; the upper jaws are very powerful and hooked at the tip, but they are destitute of teeth or have them very short; the labial palpi (jig. 3. 4.) are 4-jointed ; the basal joint is, however, soldered to the lip, merely forming a support to the three terminal joints. The palpi are generally longer than the mandibles (fig. 4.1.11.), and smooth or but slightly clothed with bristles. The body is oblong, rarely ovate, and the posterior legs have a vertical as well as a horizontal motion, the pectoral laminez (coxz) to which they are attached, being small (fig. 4. 9.). Mr. Kirby has observed that in all those species which have the anterior tibiz notched (with the exception of those which have the elytra dis- tinguished by elevated ridges) there is usually a series of from twelve to thirty ocellated impressions near the Jateral margins of the elytra on the 8th, or between it and the 9th furrow: these impression, but not ocellated, also exist in Cicindela; the typical number of furrows is nine, but in Omophron and some others there are fifteen (Fawn. Bor. Amer. p.7.) Fig. 3. 6. represents an elytron of Omaseus me- lanarius. Many species are furnished merely with rudimental wings: these, however, in some instances are fully developed in certain spe- cimens of the same species; occasionally also the elytra are soldered together, whence these insects are distinguished from the Cicindelida, which are especially fliers, by their great activity and running powers, and whence they have been termed by Messrs. Kirby and Spence, Eupodina.
The body of these insects is of a very firm consistence, whereby they are enabled to creep about under stones, &c., as well as
* In the genus Trigonodactyla, as has been obseryed by M. Audouin, the max- ille are terminated by an articulated hook.
+ These lateral appendages, or paraglosse, as they have been termed, have been generally considered to be portions of the lip itself; such, however, is not the case, as they are evidently the exserted margins of the internal lingua or tongue (jig. 3. 4; 5.)
60 MODERN CLASSIFICATION OF INSECTS.
prevented from falling beneath the power of the insects which they attack; most of the species of this family being eminently insectivorous, prowling about on the surface of the ground, under stones, &c., or beneath the bark of trees, or in the moss growing at the roots of the Jatter, in search of their prey, which consists of larve, or of herbivorous beetles and other insects, sometimes even attacking their own species. Latreille in like manner informs us (Histoire Générale, &c., vol. viii. p. 374.), that the Scarites gigas burrows into the earth under cow-dung in sandy places by the assistance of its an- terior palmated fore legs, which singular habitat appears to be ac- counted for by the circumstance, also stated by him, that when confined with some of the stercoracious lamellicorn Beetles (Helio- cantharus sacer, &c.), they pull them in pieces and devour them; and M. Lefebvre de Cerisy, who has published some observations upon the same species, informs us that they are nocturnal insects of prey remaining in their retreats during the day, but sallying forth at night and preying upon such Melolonthe, &c., as fall in their way. These latter insects seem indeed to be a favourite food of the Carabidee in general, for the former author elsewhere states that the chief nourish- ment of the true Carabi consists of the larvee of insects and of herbi- vorous beetles, such as Cockchafers, Rosechafers, &c., which,as Mr. Dill- wyn also observes, they certainly prefer (Memoranda, p. 56.) Latreille adds that the beautiful Carabus auratus, which is probably thence termed in France “ Le Jardinier,” devours more cockchafers than all the other enemies of these destructive insects by attacking and killing the females at the period cf oviposition, and thus preventing the birth of thousands of young larve. These insects are therefore of es- sential service in keeping down the numbers of noxious insects with which our gardens and pastures might otherwise be overrun. It has been suggested to me by a celebrated zoologist, that the carnivorous quadrupeds which are the analogies of the predacious land beetles, in like manner prey upon herbivorous quadrupeds. *
* The anterior tibiz of a great number of the species are deeply notched near the tip, on the inside; and Mr. Curtis has noticed an interesting peculiarity of structure in the anterior tibia of the genus Cillenum, which are not only armed with the two ordinary spurs (one above and the other below this notch), but have, also, two additional deflexed spines at the outer extremity of the notch, between which spines he presumes the lower moveable spur is received: hence he conceives that these notched anterior legs of the Carabida are used in seizing and retaining their prey, for the limb of an insect being received into the notch, and the lower
COLEOPTERA. — CARABIDE. 61
The habits of the family are not, however, exclusively carnivorous, since some of the species generally found in cornfields are clearly ascertained to feed upon growing grain; thus Nicolai observes re- specting Broscus cephalotes “in agris occurrit frumenti vastator videtur, certe ejus larva.” (Dissert. Inaugural. &e. Hale, 1822, p.12.) La- treille also says “ J’ai vu souvent aristus bucephalus grimper sur des graminées en arracher les bales et les emporter avec ses mandibules.” ( Regne Animal, vol. iii. Ist ed. p. 190.) : hence, he was induced in the Coléoptéres d’ Europe, to consider that the Scaritides were less car- nivorous than the other Carabide. Its larva, however, resembles that of the Cicindele, both in form and habits; and as the carnivorous habits of another of the Scaritides had been previously observed upon by him as above mentioned, it seems not improbable that the circumstance noticed, of the Aristus thus feeding upon grains, was accidental. The larve of Zabrus gibbus (fig. 2. 6.) also, upon the authority of Germar and otlier German naturalists ( Magaz. der Entomol. vol. i.), which are produced from numerous clusters of eggs a short time previously deposited by the females, feed during the night upon the young shoots of wheat, occasionally doing great mis- chief, as in the instance recorded by Germar, where 230 acres of wheat in the canton of Seeburg, near Halle, were destroyed by it. During the day these larva were observed to bury themselves in the earth several inches deep; they remained about three years in this state, and attained their full size in June, when they formed for themselves an oval cavity in the earth, sometimes two feet deep, in which they became pupe, appearing in the perfect state at the end of a month, when they resumed their destructive habits by climbing up the stalks and feeding upon the grain. When confined in a box in which several ears of corn had been placed, they first fed upon the grains ; and when they had consumed the whole, they then attacked each
moveable spur being then pressed upon it, the insect would be effectually secured, whilst the spurs of the posterior legs are supposed to be employed in securing a firm footing upon the objects they wish to destroy or tear to pieces. The former of these suppo- sitions is probably correct ; indeed, a direct observation in support of it will be found in the account of the Cantharide : the latter, however, is probably, less correct, since many insects, which have the spurs of the hind legs much developed, are remarkable for their leaping and shuffling kind of motion, as Orchesia, Platyura, &c. I imagine that the claws alone are employed in retaining hold of objects by insects whose tarsi are simple.
62 MODERN CLASSIFICATION OF INSECTS.
other. These larvae were accompanied, in the proportion of about one fourth by the grubs of the Melolontha ruficornis ad. which, from belonging to a group of insects eminently destructive to the roots of vegetables, must have augmented the mischief. Messrs. Curtis and Stephens have, indeed, doubted whether any portion of the injury is attributable to the Zabrus, the latter author considering that it might rather have been entirely caused by the grubs of the chaffer, and that the larve of the Zabrus (belonging to a decidedly carnivorous group) contributed to lessen the damage by devouring the other larvee whilst the perfect insect ascended the corn for the purpose of feeding upon the insect parasites thereon, and hence questioning whether the de- struction of the Zabri would be desirable. The opinion of Mr.Stephens (Illust. Brit. Ent. Mandib. i. 140.) was adopted also by Messrs. Au- douin and Brullé in their first volume. Contrary, however, to these opinions, we have the express statements of Germar and Kollar (Naturg. Sch. Ins.): the circumstance of the imago eating the grain when confined, the comparative paucity of the grubs of the chaffer, whence it would happen, were the larvee of the Zabri really carnivorous, that they would be unable to find sufficient food to supply their ravenous appetites ; moreover, the grubs of the chaffer are subterraneous, whilst it is stated that the young growing shoots were devoured, and lastly, the circumstance that the perfect Zabri are constantly found in corn- fields and even upon the ears of corn, crawling up the stems or hovering upon the wing just above the heads of the corn, as Mr. Vigors has informed me that both sexes may occasionally be observed. The question, however, has been set at rest, at least as regards the perfect insect by the Rev. G. T. Rudd, who has stated to me that he and a friend distinctly perceived several specimens of Zabrus upon the ears of corn, opening the husks and extracting and devouring the grain. He has subsequently published an account of this circum- stance in the Entomological Magazine, vol. 11. p. 182. *
I have in like manner noticed several of the large black species of Harpalides running about pathways near cornfields, especially in the evening, and creeping from amongst the standing wheat, where it is probable that they have been feeding during the day: their appearance
* Messrs. Audouin and Brulle, in the fifth volume of their work upon the Coleop- tera, adopted this opinion, being contrary to that which they had entertained in the fourth volume of their work. Mr. Tulk has also informed me, that he has kept several Zabri alive, for many months, feeding them upon grain.
COLEOPTERA. — CARABID/ZE. 63
in the evening, although it certainly bespeaks a predacious dispo- sition, does not disprove their herbivorous habits during the day. I have also occasionally observed several of the metallic-coloured species of Harpali, Amare, &c., amongst the upper branches of umbelliferous and other plants during the autumnal months, where it is not im- probable that they ascended for the purpose of feeding upon the ripe seed. M. Zimmermann, in a valuable memoir upon the genus Amara has published an account of their habits, from which it appears that vegetable substances form their chief support; they feed upon the pith and stems of graminez, succulent roots, and likewise upon the larvee of other insects which they can master; they eat much, as may be seen from the swelling of the abdomen. Hence, as Messrs. Au- douin and Brullé (v. p. 29.) observe, this genus, together with the Zabri, compose a group of herbivorous species amongst the carnivorous family to which they belong. Signor Passerini has also published a Memoir upon the ravages occasioned to young wheat by the larvae of Zabrus gibbus and Calathus latus.
Some of the Harpalides, as we are informed by the Rev. Mr. Bird, not only fly by night, but are attracted to the light of a lamp, some- times in great numbers. (/ntomol. Magaz. ii. 41.) Mr. G. R. Water- house has also informed me that a specimen of Harpalus ruficornis flew into his chamber in which a light was placed, a short time previous to a storm; hence it is evident, not only that these insects seek their prey by night, but that their flight is high, and that they are subject to the same influence arising from lights as moths. Mr. Curtis has also recorded a similar circumstance respecting Harpalus griseus. The flight of the former insect, shortly previous to a storm, is also curious, as it is well known that many insects are peculiarly liable to be affected by sudden atmospherical changes.
Some of the species, especially the larger Carabides, exhale a fetid odour, discharging at the same time from the abdomen to a con- siderable distance a caustic and acrid fluid, whence some of them, as the Carabus auratus, already mentioned, have obtained the name of *‘ Vinaigriers” in France; indeed, as in the instance recorded by De Geer, in which some of this fluid was discharged into the eyes of his friends, it is capable of producing very violent pain for several moments.* It is owing to the rapid volatilisation of this liquid, that the genus Brachinus makes the explosions subsequently described.
* See Dufour, Recherch. Anatom., p. 204.
64: MODERN CLASSIFICATION OF INSECTS.
In like manner Anchomenus prasinus (one of the Harpalides) is said to combat its enemies with repeated discharges of smoke and noise. (K. and S. ii. 247.) They also discharge a very acrid liquid of a black colour from the mouth, when captured or whilst eating, which pro- duces similar effects on the skin to those caused by powdered can. tharides; indeed, by the ancient physicians they were employed in medicine as being highly vesicatory, whence Geoffroy considered them to be the Buprestis of the ancients, which were accounted as a dangerous poison to cattle. Latreille, however, in a valuable memoir upon these poisonous insects in the tenth volume of the Annales du Muséum, has satisfactorily shown them to have belonged to the modern family Cantharide.
Olivier informs us that a sort of soap is made by the natives in some parts of Africa from a species of Carabus, to which he gave the specific name of Saponarius, belonging to the subfamily Harpalides ; and from information given by M. Dumolin to Dejean, it appears that these insects are not employed in the manufacture as affording alka- line matter, as had been supposed, but for the fatty matter which is obtained from them.
The majority of the Carabidze inhabit the elevated regions of the southern parts of Europe, Asia, and North America. The first of these quarters of the globe is, however, their chief metropolis, none other possessing so great a number of species. In England we scarcely possess more than 400 species. The insects composing the true genus Carabus disappear as we go either to the North, or towards the tropics, whilst the allied genus Calosoma is distributed over Europe, Africa, America, and New South Wales. The genus Pam- borus is confined to the latter country, whilst Procerus and Pro- crustes are chiefly found in the countries bordering the northern shores of the Mediterranean Sea, where other equatorial genera begin to make their appearance. Some species are found upon the margins of streams or standing water, whilst some of the Carabi and Pteros- tichi, &c. are only found on the tops of the highest mountains. Again, many of the species of the Brachinides are generally found either upon the trunks or beneath the bark of trees.* These species are likewise more variegated in their colours and lest robust in their
* It is evidently owing to this circumstance that some of these insects are occa- sionally found embedded in gum anime, in which substance I have observed several beautiful species. (See also Germar, Mag. Ent. vy. i. p. 13. G. Lebina.)
COLEOPTERA. — CARABIDZ. 65
structure than the majority of the family. They are also cha- racterised in many instances by having the tarsi dilated, and the penultimate joint more or less bilobed in both sexes, a peculiarity re- sulting, as Bonelli has observed (Mem. Acad. Turin, vol. xxiii.), from the situations in which they reside, this structure being in fact ana- logous to the cushioned feet of flies as well as of those of the true herbivorous Beetles. M. Lacordaire has confirmed these observations in his account of the habits of the Brazilian Coleoptera (Ann. Se. Nat., June, 1830).
The internal anatomy of the Carabide has been investigated by M. Dufour, in the Annales des Sciences Naturelles, for August, Oc- tober, and December, 1824. Ramdohr also has published an account of the anatomy of Carabus monilis, in the Magazin Gesellsch. Na- turf. zu Berlin, vol. i.; and Dr. Burmeister’s Memoir on the Anatomy of the Larva of Calosoma, published in the Transactions of the Ento- mological Society, vol. i.. must also be mentioned.
Few observations have hitherto been made relative to the larve of this family: this may indeed, perhaps, be partially accounted for, from the obscure situations in which they are found, and from the difficulty of rearing them. Those, however, which have been noticed, are equally voracious with the perfect insects, and are found in simi- lar situations: they are generally long, with the body of equal breadth throughout, and the head of a moderate size, with the rudimental eyes equal; a scaly square prothorax, and with the eighth segment simple, and resembling the remaining joints: the terminal segment is, however, armed with two conical horny appendages, toothed in the Carabi and Calosome, but (according to Latreille, whose state- ment I am, however, inclined to doubt) long, fleshy, and articulated in the larvee of the Harpali and Licini; in the latter, also, the body is shorter and the head larger. The part where the anus is situated is prolonged into a membranous deflexed tube, which serves as a sup- port to the tail: the jaws resemble those of the perfect insect.
The larvee of Ditomus bucephalus, however, according to Latreille (Régne An. second edit. t. iv. 190.), resemble those of the Cicindelx, whilst that of Omophron decidedly approaches the form of those of the Water-beetles (Dyticidz ).
One of the most voracious larve in the family, is that of Calosoma Sycophanta, first described by Réaumur, in the third volume of his Mémoires. It is black, with six scaly legs: when full-grown it is
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66 MODERN CLASSIFICATION OF INSECTS.
an inch and a half long, subdepressed and fleshy; the upper surface of each segment and the head are, however, more scaly ; the tail is terminated by two horny spines, and the head is armed with two powerful jaws, wherewith it seizes the body of its struggling prey, which consists of the fleshy larve of | Lepidoptera. As it is very ravenous, devouring several large Cater- pillars in a day, it has the sagacity to find its way to the nests of the gregarious processionary Moths, in the midst of which it takes up its abode; and Réaumur states, that he never found a nest of these Caterpillars which was not infested with from one to six of these larva: here they feed in the midst of their prey (not even sparing them when they have become chrysalides), in so gluttonous a manner that the segments of the body become distended, and ren- der it unfit for any movement, so that it becomes an easy prey to its young and more active brethren, which, in mere wantonness, seize it with their jaws, although surrounded by their natural food. All Réaumur’s larvee died before they became perfect insects ; but there can be but little doubt, that they are the larva of the Calosome, of which, as Réaumur observes, the habits are precisely similar ; in- deed, Clairville, and the French entomologists, appear to entertain none upon this subject. And M. Boisgerard, in a communication made to M. Audouin (published by the latter, in the Hist. Nat. Ins. t. v. p. 94.), states, that having placed some female Calosome upon trees greatly infested with the Caterpillars of Bombyx Dispar, the larvee of the Calosomz were found in the following season in the nests of the Caterpillars, and that in the course of two or three years the trees were cleared. Dr. Burmeister has published a very elaborate account of the anatomy of the larvae of Calosoma Sycophanta (which is not rare in the pine forests in the vicinity of Berlin, where, both in the larva and perfect state, it devours the Caterpillars of Bombyx Dispar, and other Moths,) in the first volume of the Transactions of the Entom. Soe. of London, illustrated with two plates. This larva does not confine itself to Lepidopterous Caterpillars, for Nicolai informs us, that it is occasionally found in some profusion, in the pine forests near Halle in Germany, devouring the larve of the Saw-fly of the pine, Lophyrus Pini. (Déssert. Inaugural, p. 13.)*
M. Audouin has also published (in the Hist. Nat. Ins. vol. v. p. 99.) an account of the larva of one of the largest European species of the
* Dr. Ratzeburg has given several very characteristic figures of this larva (as well as of that of Cicindela campestris) in his Forst. Insect. (pl, 1.), just published.
COLEOPTERA. — CARABID®. 67
family, Procrustes coriaceus, found in France, and also, but very rarely, in England: the larva resembles that of the Calosoma; it is corneous, subcylindric, and terminated by two strong curved spines, each of which is furnished beneath with a smaller spine. It is about an inch and a quarter long, lives under moss and in damp situations, where it feeds upon snails and slugs. It is full grown in the spring, be- comes a pupa at the beginning of May, in a cell which it forms in the ground, and remains in that state not more than a fortnight. The perfect insect also feeds upon snails.
Dr. Heer has described and figured the larvee of the following species of Carabus and of Cychrus rostratus in his Observationes Entomolo- gice, 1836.
The larva of Carabus auronitens (fig. 2. 1.) is very similar to that of Yeti ly,
Calosoma in its general form, the mandibles sickle-shaped with a strong tooth inside at the base, and the terminal segment of the body armed on each side with a thick horny spine with a spur on each side. The antennz are 4-jointed, as well as the maxillary palpi: the eyes are simple ocelli, there being six on each side. It was found on the Ist of June under a stone, and was transformed to the pupa on the 3d, and to the imago on the 15th of the same month. The pupa (jig. 2. 2.) is of an ordinary form, with the abdominal seg- ments rounded at the sides, and furnished on each side with a fascicle of hairs; the anal segment is terminated by two conical appendages. The larvae of Carabus depressus and C. hortensis are also very similar to the preceding, differing chiefly in the greater or less breadth of the abdominal segments, and in the greater elongation of the anal fork, which are simply setulose.
Clairville also states, that the larva of one of the species of the restricted genus Carabus, which he had noticed, precisely resembled that of Calosoma, figured by Réaumur.
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68 MODERN CLASSIFICATION OF INSECTS,
Dr. Heer states, that the larva of Cychrus rostratus (jig. 3. 3.), “a Caraborum larvis capite parvulo, palporum articulis exterioribus cras« sioribus, mandibulis erectiusculis, et corporis forma breviori latiorique facile distinguitur ;”” in addition to which the prothorax is conical, and the anal segment very short, with the hinder angles prominent, and armed in the middle with two very short and simple spines.
De Geer has figured a larva (Mémoires, vol.v. pl. 12. f. 1—3.) closely
resembling that of Calosoma, which was found in the ground, and which he presumes to be the larva of one of the species of this family. He was not, however, able to rear it. _In the first edition of the Coléoptéres d Europe, M. Latreille announced a memoir upon a larva sent to him by M. Westermann from Bengal, supposed to be that of Anthia 6-guttata. This larva, since the death of Latreille, has been described and figured by M. Lequien in his Mo- nograph upon Anthia, published in M. Guerin’s Magasin de Zoologie, Insectes, pl. 41., and by Messrs. Audouin and Brullé, in their Histoire naturelle des Insectes. It is nearly three inches and half long (fig. 2. 8-) with a subcylindric scaly body, consisting of twelve segments, exclu- sive of the head (fig. 2. 9. under side of head), which is armed with strong acute mandibles, very short three-jointed antennz, and a single pair of eyes; the upper side of the head is rugose with several tubercles in front ; the terminal segment is scabrose, serrated at the sides, with a deep notch at its extremity ; this segment is furnished beneath with a single fleshy anal proleg. The second? and nine fol- lowing segments are respectively furnished with a pair of lateral spiracles ; the mentum and maxilla are long and extend almost to the base of the head. ‘The legsare short. This larva is of a black colour, with the posterior margins of the segments obscure red. It has not been stated upon what grounds this larva was considered as that of Anthia 6-guttata, and indeed upon comparing the details of its structure, and especially its trophi, with that of the known Carabideous larve, I have no doubt that it does not even belong to any of this family ; since, in its general appearance and organisation, it approaches much nearer to the larve of the Elateride.*
* Since the above was written, I have received the following communication from M. Westermann himself, relating to this larva, in answer to the inquiries which F was induced to make concerning it: “ With the greatest pleasure I shall give you all the information in my power of the circumstances which led me to believe that
the larva I brought from India, and sent to the late M. Latreille, was that of Anthia 6-guttata. M. De Haan of Leyden made a similar inquiry last year, and I informed
COLEOPTERA. — CARABIDE. 69
The larve of Zabrus gibbus (fig. 2.6.) already mentioned, are described and figured by Germar in the first volume of his magazine ; they are long and flattened, being nearly of equal breadth throughout, except towards the tail, which is gradually narrowed: the body ap- pears to be of a more fleshy consistence than in the majority of these larvee, and the body is terminated by two small spines. The pupa does not present any material characters.
A Swedish naturalist, M. Acrel, in a Memoir inserted in the sixth volume of Nova Acta, of Upsal, relating to larve and other objects sometimes found in the human stomach, has described and figured a larva which he considers to be that of Sphodrus leucopthal- mus, and which had been ejected from the stomach of one of his patients. Such, of course, is not the real habits of this insect, since Latreille considers that the larve figured by Geedart, (fig. 107. a, 6,) are referrable either to this species or to Abax striola. Certainly, Geedart’s insects, in their general appearance, strong jaws, and pre- dacious habits, coincide with the typical larvee of this family, one of them being described as preying voraciously upon softer larvae; and the other, which remained in the larva state two years, and of which he has given the transformations, feeding, not only on other larve, but also on the pupe of ants, the eggs of mole crickets, &c.
I am not certain to which species of the family the larva which I have represented in fig. 2. 4. is referable; it agrees, however, with
him that being on a visit in Burdwan in Bengal, one night returning home I ob- served by the light in a lantern the larve crawling in theroad. I immediately took it to be the larva of some large Coleopterous insect, but had not the least idea to which it belonged till the day after, when I observed at the foot of a large Banian tree several Anthia guttata, which I could not, however, secure, as they retreated into holes when I came nearthem. I therefore ordered my palankin-bearers to dig them out, when we at the same time obtained another of the lary found on the pre- ceding evening ; without being positive, as I wrote to M. Latreille, I now, naturally considered it to be that of an Anthia and concluded it was in the habit, during the night, of leaving its hole and crawling about in search of worms. M. De Haan hay- ing observed to me that this larva, whereof he had seen a figure in the French works, appeared according to his observations to belong to an Elater, I informed him that on the very tree where we obtained the larve from the holes at the roots, I found Elater fuscipes Fabr., in great abundance; this being the largest Elater which ever came under my observation in Bengal. I think this larva is much too large for such an insect, nor should I think a larva of an Elater would crawl so well on the ground. In other parts of India very large Elaters are found ; in Jaya I once took a pair as large as E. flabellicornis.” Copenhagen, July 5. 1836, F 3
70 MODERN CLASSIFICATION OF INSECTS.
the typical larve of the family. And as I have found numerous specimens of it, as wéll as others, but slightly differing from it in essential characters, I have little doubt that it belongs to one of the large common black Harpalides (perhaps Steropus madidus, or Omaseus melanarius), and do not hesitate to give it as an example — of the group. It is black and scaly, the head (fig. 2. 5. under side) is armed with two acute mandibles, two moderately long 4-jointed antennze, two long 4-jointed maxillary palpi, and two short thick labial palpi; there are six small simple eyes on each side behind the antennz, set in a circle. The under side of the head is scaly, the parts of the mouth being inserted near the front ; the legs are moderately long, and the terminal segment of the body is armed with a pair of horny bifid spines.
The larva represented in jig. 2.7. was described and figured by Desmarets (Bulletin Soc. Philomat. vol. i.), whose reasons for re- garding it as that of Omophron limbatum appear to be satisfactory ; it is of an intermediate form between that of the larvee of the Cara- bide and Dyticide, being oblong, depressed, narrowed behind, with the head very large, and broader than the rest of the body, and armed with two very robust jaws, and other usual parts of the mouth, two 5-jointed antennz, and two small eyes placed at the sides of the head ; the tail is formed of a slender 4-jointed appendage, with two hairs at its extremity; the legs are moderately long. It was gene- rally found in moist sandy places on the banks of the Seine, where the perfect insect was also met with in company with it, the food of both consisting of small sub-aquatic insects. The larva is very active, and, when touched, throws up its tail in the manner of a Staphylinus.
he anatomical observations of M. Dufour seem to confirm the intermediate location of this genus between the Carabidee and Dyti- cide ; the perfect insect, although unable to swim, cannot live except in damp situations, and, indeed, Clairville placed it at the head of the Water-beetles.
M. Zimmerman, in his Monograph upon Amara, has given some ac- count of the early stages of the insects of this genus. Considering some species as annual, and others as producing two broods in a year, and that the period of their appearance is not, like that of Moths, &c. fixed to a particular time, he adds, that the duration of life of these insects, from their bursting from the eggs to their death, never exceeds a year, whilst, in some species, it is certainly shorter. Those which
COLEOPTERA. — CARABID&. Tt
appear at the end of the summer in the perfect state, generally couple in the autumn, whilst in others this does not take place until the following spring. The eggs do not arrive at maturity until seve- ral days after coupling ; they are deposited under stones, or at a small depth under ground. The larvee soon appear; they change their skin once, and attain, previous to becoming pupe, twice the length of the imago: the larve have the same general form of those of the Zabri and Peecilli, and are so much alike that it is not possible to distin- guish the species. They remain about six or eight weeks as larvae, and half that time as pupe; the perfect insects are able to live a much longer time, especially if coupling is delayed, immediately after which they die.
Latreille is the first author who directed the attention of ento- mologists to the distribution of the numerous groups composing this family. So far indeed from abandoning the hope of effecting a natural arrangement (as asserted by Mr. MacLeay), he has proposed a classi- fication of them “ fondée sur des rapports naturels” in the Coléopteres d' Europe which appears to me to be the most natural of any given, and which I propose to adopt in the following pages. He considers that the first group having truncated elytra, and noted for their ex- plosive powers (Brachinidee MacLeay, Truncatipennes, Latr.) is con- nected on the one side with the Cicindelide by means of Anthia and some other genera; and on the other, to the second group dis- tinguished for its fossorial habits, having the elytra separated from the thorax by a wide incision (Scaritide MacL., Bipartiti Latr.) by means of Enceladus and Siagona. From this second group he pro- ceeds by Aristus and Acinopus to Zabrus, Cephalotes, &c. in the third group composed of the Harpaliens and Feroniens (Harpalidz MacL., Thoracici Latr.) distinguished by their great quickness in running; whence by means of Licinus, Panagzeus, &c. we approach the fourth group, having the inner surface of the anterior tibia not notched, and which emit an extremely acrid fluid (Carabidae MacL. Abdominales, Zaér.), from which by Nebria, Elaphrus, &c., we arrive at the fifth group, composed of the genus Bembidium, having the last joint of the maxillary palpi minute and acute (Bembidiide Stephens, Subulipalpes Laér.), which approach the Water-beetles in their sub- aquatic habit: the genus Haliplus, amongst the latter, closely re- sembling Bembidium in the curious structure of the palpi.
Other instances in support of Latreille’s arrangement may be given;
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72 MODERN CLASSIFICATION OF INSECTS.
thus Ozena hitherto regarded as one of the Bipartiti of Latreille, is now, from its explosive powers, proved to belong to the Bombardiers ; whilst on the other side Mr. Curtis has suggested that a connection between the Scaritides and Harpalides may be effected by means of the genera Miscodera (Leiochiton Curt.) and Clivina; but a more beautiful connecting link between these two subfamilies exists in a yet undescribed insect from New Holland in the collection of the Rev. F. W. Hope, which strongly partakes of the characters of both groups; whilst between the Harpalides and Carabides the genus Eurysoma (Brachygnathus Perty) having the appearance of Cychrus, may be mentioned, as well as the genus Teftlus, regarded by M. Brullé as more nearly allied to Panagzeus than to Carabus.
Latreille, indeed, was subsequently induced to adopt an arrange- ment of these groups proposed by Dejean by commencing with the Abdominales or Carabides, and in so doing has pointed out other apparent transitions; but in their more recent works, both authors have abandoned the latter arrangement and adopted that given above, with the exception that Dejean has raised these five groups, as well as two others, separated from the Harpalide, to an equal rank with the Cicindelide. Mr. MacLeay, also, in his Annulosa Javanica, has elevated the sections of the Linnean genus Carabus to an equal rank with the last-mentioned family ; but from pursuing a quinarian ar- rangement of the Geodephaga, he has been enabled only to admit four divisions of the Carabide. The group of Bembidiide is there- fore sunk, the structure of the palpi being considered insufficient for its support: if this character were indeed the only distinctive mark of the group, this step might still be questioned, as no other Carabidae possess such a structure; but the group is also characterised by the minute size, great agility and subaquatic habits of the insects of which it is composed; moreover, in point of number of species, it is not an inferior group. If even it were to be suppressed, its situation would evidently be nearer to Elaphrus than to Harpalus, which would, of course, interfere with Mr. MacLeay’s distribution, which (notwith- standing his remark upon the Latreillian distribution) is, with the exception of the suppression of the Bembidiidz, identical with that given above; the progression being merely reversed, by which means a passage is effected between the Cicindelide and Elaphrus amongst the Carabidae *, which two families he considers as forming a normal
* Mr. Curtis has adopted this view in his observations on Pelophila, although in those upon Elaphrus, he regrets that Dejean should have removed that genus and
COLEOPTERA. — CARABID#. 73
group, having entire tibize, whilst, in his three other families, which he terms aberrant, they are deeply notched.
Messrs. Kirby and Spence divide the Carabidz into two subtribes, Lamprona, named from the brilliancy of colour of many of the species and corresponding with Latreille’s Abdominales, and Amaurona or those of obscure colours, and comprising all the other groups of La- treille, the latter subtribe being divided into stirpes, as Truncipennwza, &c.; these stirpes, however, being evidently of equal value with the entire subtribe Lamprona, this primary mode of division cannot be considered satisfactory.
Mr. Stephens has proposed six families of Carabideous insects : — 1. Brachinide Mach. 2. Scaritide Mach. 3. Carabide Macl. 4. Harpalidee MacL. 5. Bembidiidze Steph., and 6. Elaphridee Steph. This distribution is, however, evidently unnatural, separating, as it does, the Harpalide and Scaritidz, and the Carabidae and Elaphride. No affinity, moreover, is traced between the Scaritide and Carabide.
Other modes of distribution have been proposed by Zimmerman (Monog. Amaroidum), Laporte (Etudes Entomol. livr. i.) and Brullé (Hist. Nat. Ins. Col. vol. i.), which it would be tedious to detail, In all these arrangements, however, the Harpalides are divided into three groups, corresponding with the Simplicimana, Patellimana, and Quad- rimana of Latreille (#égne Animal, 2d ed.). The latest arrange- ment of these insects is that of Mr. Kirby, given in the Fauna Boreali- Americana, and which is as follows :—
Euptera, (Cicindelide. ) Truncipennia. Geodephaga. f Guechanie M‘L.) Guresipcanis f Nobilia (Carabus, &e. ). Buceccha | Plebeia (Nebria, &e. ). z Sarrothropoda. | (Chleenius. ) G H Dimana ( Agonum, Calathus, _Cystopoda. and Feronia, &c. ). | Quadrimana ( Harpalus). Quadrimana. Acutipalpia Re sit as ) (Trechus, Patrobus, &c. ). Hygradephaga. Subulipalpia. (Peryphus, Bembidium, &e. ) Filipalpia. ( Elaphrus, )
Omophron (both belonging to the same group) from the termination of the Carabide, considering the latter genus as “ ordained by nature perfectly to connect the Carabide with the Dyticide.”
74 MODERN CLASSIFICATION OF INSECTS.
Mr. Kirby as well as Messrs. Laporte and Brullé have also adopted the plan of subdividing the primary groups into numerous minor sub- divisions named after the typical genus in each: thus by both the last named authors, the Brachinide are divided into six minor groups, not however corresponding with each other, as the sixth subdivision (Brachinites) of Laporte embraces Brullé’s Ist, 5th, and 6th, sub- divisions,
In considering the different primary divisions of this very natural family, as possessing a value inferior to that of distinct families, and as constituting collectively one family of equal rank with the Ci- cindelidz, I am but treading in the steps of Linnzus, Latreille, Kirby, and Leach, as well as in those, I cannot but think, of nature herself, notwithstanding the numerical majority of the Carabideous over the Cicindelideous insects. The following is a concise tabular sketch of these different groups which I have considered as sub-families, termi- nating them, for uniformity, in zdes.
A. External maxillary palpi terminated by a joint at least as large as the preced- ing (fig. 3. 2.). a. Anterior tibiz having a deep notch on the inside, near the tips ( fig. 3.192.). * Elytra generally truncated behind (fig. 3. 7.). Tarsi ] 1. Brachinides. of the g seldom dilated at the base. } ( Bombardiers. ) ** Elytra rounded at the extremity; abdomen pedun- ) 2. Scaritides. culated (fig. 3. 14.). J (Burrowers. ) ***® Elytra rounded behind; abdomen not pedunculated | 3. Harpalides. (fig. 4. 1.). } ( Black clocks.) Koen coghote bo 5 , 4. Carabides. b. Anterior tibiz net notched near the tip (jig. 4. 12.) { (Dischargers. ) 5. Bembidiides.
B. Palpi terminated by a minute conical joint ( fig. 4. 18.). { (Subaquaties.) ubaquaties.
The first sub-family BracuinIDEs, in addition to the truncature of the extremity of the elytra, are characterised by having the head and thorax narrower than the abdomen (fig. 3.7. Lebia Crux minor, 8 ungues, 9 antenne). The labium is often oval or square, and occasionally furnished with two small lateral linear iobes: in many of the smaller species the penultimate joint of the tarsi is bilobed (jig. 3.10. anterior tarsus of Cordistes), and the anterior tarsi of the males are very rarely dilated at the base; in some of the large exotic genera, as well as in the true Brachini, the body is apterous, whilst others are very narrow and subcylindric, and have been supposed to be allied to the cylindric Cicindelide.* Their colours are more
* M. Bruilé places his division Trigonodactyliens at the head of the Carabide, M. Audouin having observed that the maxille in the typical genus have the terminal unguis articulated, as in the Cicindelide.
COLEOPTERA. — CARABIDZ. 75
pleasantly variegated than in the majority of the family. Other pecu- liarities have already been pointed out.
This sub-family, as at present constituted, is perhaps the most in- congruous of all the sub-families of the Carabide, the term Trunca- tipennes applied to it by Latreille by no means indicating a constant character, as many of the species have the elytra rounded at the tips. The tarsi indeed are generally alike in both sexes, or if dilated in the males, the dilatation is of a different character from that of the other sub-families. In may, indeed, be rather regarded as a convenient receptacle for such groups as have not the bipartite and palmated structure of the Scaritides, the simple tibiz of the Carabides, the dilated male tarsi of the Harpalides and its subdivisions, or the minute conical terminal joint of the maxillary palpi of the Bembidiides. The tarsal claws are often denticulated (fig. 3. 8. claws of Lebia) ; but this structure is found elsewhere in the family.
The typical species of this sub-family are endowed with a means of defence of the most singular description, from whence they have derived the name of Bombardier-beetles, and which was first men- tioned by Rolander in the Transactions of the Academy of Stockholm for 1750. When the insect is attacked by other and larger insects, or placed in a situation of danger, it suddenly emits from the anus a fluid of so highly volatile a nature that, immediately upon coming into contact with the atmospheric air, it becomes a bluish vapour of a very pungent scent, accompanied with considerable explosion, whereby the insect is enabled to effect its escape. This it is enabled to repeat seven, eight, or even a dozen times. It is stated by an anonymous author (Ht. Mag. iii. 377.), that on immersing a specimen of Br. Grecus in boiling water, it let off one of its explosions, and the water
76 MODERN CLASSIFICATION OF INSECTS.
for about an inch around it effervesced much in the same way as a Seidlitz powder. The typical species Brachinus crepitans Linn., is not uncommon, being found in company under stones; and it is stated that these communities have been found at times to consist of at least a thousand individuals. The effect of this vapour upon the skin is very similar to that produced by nitric acid. M. Dufour has published a description of the organs by which this fluid is secreted in the Brachinus balista, which will be found in the eighteenth volume of the Annales du Muséum, as well as in the sixth volume of the An- nales des Sciences Naturelles.* Kuhn has also published an account of the habits of these insects in the thirteenth volume of Der Natur- forscher. Some authors have thrown doubts upon the statement that the explosion of these insects is accompanied by a noise; the fol- lowing circumstance, however, communicated to me by the celebrated traveller Burchell, will be sufficient (were other evidence wanting, which is not the case) to confirm the correctness of the recorded statements. Whilst resting for the night on the banks of one of the large South American rivers, he went out with a lantern to make an astronomical observation, accompanied by one of his black servant boys; and as they were proceeding, their attention was directed to numerous beetles running about upon the shore, which, when cap- tured, proved to be specimens of a large species of Brachinus: on being seized they immediately began to play off their artillery, burning and staining the flesh to such a degree, that only a few specimens could be captured with the naked hand, leaving a mark which re- mained for a considerable time; upon observing the whitish vapour by which the explosions were accompanied, the negro exclaimed in his broken English, with evident surprise, “Ah! massa, they make smoke.” ‘The late traveller, Ritchie, communicated to Mr. MacLeay some particulars respecting this secretion; his companion, M. Dupont, having taken a nest consisting of more than a thousand individuals near Tripoli. He says the crepitating matter made Dupont’s fingers entirely black