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Full text of "The Cambridge natural history"

7 



THE 

CAMBRIDGE NATURAL HISTORY 



EDITED BY 



S. F. HARMER, Sc.D., F.R.S., Fellow of King's College, Cambridge; 
Superintendent of the University Museum of Zoology 



AND 



A. E. SHIPLEY, M.A., Fellow of Christ's College, Cambridge; 
University Lecturer on the Morphology of Invertebrates 



VOLUME VI 




JUL 1 - 1P1 



,9- 




INSECTS 



PART II. Hymenoptera continued (Tubulifera and Aculeata), Coleoptera, 
Strepsiptera, Lepidoptera, Diptera, Aphaniptera, Thysanoptera, 
Hemiptera, Anoplura. 



By DAVID SHARP, M.A. (Cantab.), M.B. (Edinb.), F.R.S. 



MACMILLAN AND CO., LIMITED 

NEW YORK: THE MACMILLAN COMPANY 
I 9O I 



All rights resen.<ed 



"Men are poor things; I don't know why the world thinks so 
much of them." Mrs. Bee, by L. & M. Wintle. 







First Edition 1899 
Reprinted 1901 



CONTENTS 



PAGE 

SCHEME OF THE CLASSIFICATION ADOPTED IN THIS BOOK . vii 



CHAPTER I 

HYMENOPTERA PETIOLATA COXTINUED SERIES 2. TCBULIFERA on CHRYSI- 
DIDAE. SERIES 3. ACULEATA GENERAL CLASSIFICATION DIVISION I. 
ANTHOPHILA OR BEES . 1 



CHAPTER II 

HYMENOPTERA ACULEATA CONTINUED DIVISION II. DIPLOPTERA OR WASPS 
EUMENIDAE, SOLITARY TRUE WASPS VKSPIDAE, SOCIAL WASPS 
MASARIDAE 



CHAPTER III 

HYMENOPTERA ACULEATA CONTINUED DIVISION III. FOSSORES OR 
FOSSORIAL SOLITARY WASPS FAMILY SCOLIIDAE OR SUBTERRANEAN 
FOSSORES FAMILY POMPILIDAE OR R.UNNERS FAMILY SPHEGIDAE OR 
PERFECT-STINGERS 90 



CHAPTER IV 

HYMENOPTERA ACULEATA CONTINUED DIVISION IV. FOKMICIDAE OR 

ANTS 131 



CHAPTER V 

COLEOPTERA OR BEETLES STREPSIPTERA . ... 184 

CHAPTER VI 
LEPIDOPTERA, OR BUTTERFLIES AND MOTHS . . 304 



VI CONTENTS 



CHAPTEI! VIT 

PAGE 

DIPTEKA, on FLIES APHANIPTERA, OR FLEAS THYSANOPTERA, OR THRIPS 438 



CHAPTER VIII 

HEMIPTEUA, OK BUGS ANOPLTJRA . . . 532 



NOTES . 602 

INDEX * 603 



SCHEME OF THE CLASSIFICATION ADOPTED 
IN THIS BOOK 



Order. 



Sub-order, Division, 
or Series. 



Family. 



Sub-Family or 
Tribe. 



Group. 



HYMENOPTERA ( Petiolata. 

(continued from - (continued 
F). (from Vol. V). 

Tubulifera | CHRYSIDIDAE (p . 



Vol. 



HYMENOPTERA 

(continued) 



Aculeata 

(P- 4) 



AXTHOPHILA 

(p. 10) 
APIDAE (p. 



FOSSORES (p. 90) 

SCOLIIDAE 

(p. 94) 



-: 



IArcliiapides (p. 21). 
Obtusilingues (p. 22). 
Andrenides (p. 23). 
Denudatae (p. 29). 
10) Scopulipedes (p. 32). 
Dasygastres (p. 35). 
s Sociales (p. 53). 

DlPLOPTERA 

(P- 71) 

( EUMENIDAE (p. 72). 
| VESPIDAE (p. 78). 
I MASAIUDAE (p. 88). 

Mutillides (p. 94). 

Thynnides (p. 96). 

Scoliides (p. 97). 

Sapygides (p. 99). 

Rhopalosomides (p. 100). 
POMPILIDAE (p. 101). 

' Sphegides (p. 107). 

Ampulicides (p. 114). 

Larrides (p. 116). 

Trypoxylonides (p. 118). 
SPHEGIDAE I Astatides (p. 119). 
(p. 107) ] Bembecides (p. 119). 

Nyssonides (p. 123). 

Philaiithides (p. 124). 

Mimesides (p. 127). 

Crabronides (p. 128). 

'Camponotides (p. 144). 
Dolichoderides (p. 157). 

Myrmicini 

(p. 159). 
Attini (p. 165). 

ilvrmicides Pseud oniyrmini 

(p. 158) (p. 168)._ 

Cryptocerini 
" (p. 169). 
Ponerides (p. 170). 

I Ecitonini 

Doiylides (p. 174)] D J/^ 

(p. 177). 
Amblyoponides (p. 180). 



HETEROGYXA 
(p. 131) 

FOBMICIDAE 

(p. 131) 



VI 11 



SCHEME OF CLASSIFICATION 



Order. 



Sub-Order, Division 
or Series. 



Sub-Family or 
Tribe. 



COLEOPTERA 

(p. 184) 



Lamelli- 
cornia 

(p. 190) 



Adephaga or 
Caraboidea 

(p. 200) 



Polymorpha 

(p. 213) 



I 
(Continued on the nextp<i<f>. 



Family. 

'PASSALIDAE (p. 192). 
LUCANIDAE p. 193). 

j'Coprides (p. 195). 
SC-ARABAEIIKVE I ^elolonthides (p. 198). 



(p. 194) 



- Rutelides (p. 198). 

Dynastides (p. 199). 
^ Cetoniidi 



Cetoniides (p. 199). 

ClCINDELIPAE (p. 201). 

fCarabides (p. 206). 
CARABIDAE | Harpalides (p. 206). 

(p. 204) "I Pseudomorphides (p. 206). 
I. Morraolycides (p. 206). 

AMPHIZOIDAE (p. 207). 

PKLOBUDAE (p. 207). 

HALIPLIDAE (p. 209). 
DYTISCIDAE (p. 210). 

( PAUSSIDAE (p. 213). 
GYRINIDAK (p. 215). 
HYDROPHILIDAK (p. 216). 
PLATYPSYLLIDAE (p. 219). 
LEPTINIDAE (p. 220). 
SILPHIDAE (p. 221). 

Sc'YDMAENIDAE (p. 223). 
(iXn.STIDAE (p. 223). 
PSKLAI'HIDAE (p. 223). 

STAPHYLINIDAE (p. 224). 
SPHAERIIDAE (p. 227). 
TRICHOPTERYGIDAE (p. 227). 
HYDROSCAI-HIDAE (p. 228). 

CoRYLOPHIDAE (p. 228). 
Sl'APHimiDAE (p. 229). 

SYKTELIIDAE (p. 229). 

HlSTERIDAE (p. 230). 

PHALACKIHAE (p. 231). 
NITIDULIDAE (p. 231). 
TROGOSITIDAE (p. 232). 

(Jnl.YDIIDAE (p. 233). 
l!ll YSODIDAE (p. 234). 

CrcrjiDAE (p. 234). 
CRYPTOPHAGIDAE (p. 235). 
HELOTIDAE (p. 235). 
THORICTIDAE (p. 236). 

KlIiiTYLIDAE (p. 236). 
M YCF.TiiPHACIKAK (p. 237). 
CiiCCIXELLIDAE (p. 237). 
KXDOMYCHIPAE (p. 239). 

MYCKTAKIHAK (p. 239). 
l,\Ti:ii)iii>AK (p. 240). 
AIUMERIDAK, (p. 240). 
DKUMESTIDAK (ji. 241). 
P,VI;KHIDAE (p. 242). 
CYATHOCEUIDAE (]i. 243). 

( 1 I ui;\ ssl DAK i p. 2 l:>). 
HETEUni'llUlliAK 'p. 213). 

I'AKMD \i: |>. 243 . 
LDERODONTJDAE (p. 21 



SCHEME OF CLASSIFICATION 



IX 



Order. 



Sub Order, Division, 
or Series. 



COLEOPTERA 

(continued) 



Polymorpha 

(continued] 



Heteromera 

(p. 262) 



Phytophaga 

(p. 276) 



Rhyncho- 
phora(p. 288) 



Strepsiptera 

(p. 298) 



Family. 



ClOIDAE (p. 245). 

SPHINDIDAE (p. 245). 

BOSTRICHIDAE (p. 246). 



Sub- Family or 
Tribe. 



Jf Ptinides (p. 246). 
Anobiides (p. 246). 
(P- 248) 



Lampyrides (p. 248). 
Telephorides (p. 248). 



TINIDAE 

(p. 246) 

MALACODEK- 

MID4E 

948^ 

MELYRIDAE (p. 252). 
CLERIDAE (p. 253) 
LYMEXYLONIDAE (p. 254). 
DASCILLIDAE (p. 255). 
RHIPICERIDAE (p. 256). 

' ' Throscides (p. 260). 

Eucnemides (p. 260). 

Elaterides (p. 260). 

Cebrionides (p. 260). 

Perothopides (p. 260). 
. Cerophytides (p. 260). 

. BUPRESTIDAE (p. 261). 



ELATERIDAE 
(p. 256). 



TEXEBRIOXIDAE (p. 263). 
CISTELIDAE (p. 264). 
LAGRIIDAE (p." 264). 
OTHXIIDAE (p. 265). 
AEGIALITIDAE (p. 265). 

MONOMMIDAE (p. 265). 

NILIOXIDAE (p. 265). 
MELANDKYIDAE (p. 265). 
PYTHIDAE (p. 265). 
PYROCHROIDAE (p. 266). 
AXTHICIDAE (p. 266). 
OEDEMERIDAE (p. 266). 

MORDELLIDAE (p. 267). 

CAXTHARIDAE (p. 269). 
TRICTEXOTOMIDAE (p. 275). 

BRUCHIDAE (p. 276) 



( Eupoda (p. 280). 
| 



CHRYSOMEL- | Camptosomes (p. 281). 
IDAE (p. 278)1 Cyclica (p. 282). 

Cryptostomes (p. 282). 



CERAMBYCIDAE f . 

- Cerambycides (p. 






- . 

( Lamiides (p. 287). 



/' ANTHRIBIPAE (p. 290). 

| CURCULIONIDAE (p. 290). 
SCOLYTIDAE (p. 294). 

BRENTHIDAE (p. 295). 
f AGLYCYDERIDAE (p. 297). 
\ PROTERHINIDAE (p. 298). 



STYLOPIDAE (p. 298). 



SCHEME OF CLASSIFICATION 



Order. 



LEPIDOPTERA 

(p. 304) 



lib-Order, Division, FT mil \- Sub-Family or 


or Series. Tribe.' 


r 


[ Danaides (p. 344). 




Ithomiides (p. 346). 




Satyrides (p. 347). 




XYMPHALIDAE , Morphides (p. 348). 




(p. 343) " Brassclides (p. 349). 




Acraeides (p. 350). 


Rhopalocera 


Heliconiides (p. 351. 


(p. 341) " 


v Xyinphalides (p. 352). 




ERYCIXIDAE f Eryciiiides (p. 355). 




(p. 354) \ Libytheides (p. 355). 




LYCAEXIDAE (p. 356). 




PIERIDAE (p. 357). 




PAPILIOXIUAE (p. 359). 




^HESPERIIDAE (p. 363. 




CASTXIIDAE (p. 371). 




XEOCASTXIIDAE (p. 372). 




SATURXIIDAE (p. 372). 




BRAHMAEIDAE (p. 374). 




CERATOCAMPIDAE (p. 375), 




BOMBYCIDAE (p. 375). 




EUPTEROTIDAE (p. 376). 




PEUOPHORIDAE (p. 377^. 




SPHIXGIDAE (p. 380). 




CUCYTIIDAE (p. 382). 




XOTODOXTIDAE (p. 383). 




CYMATOPHORIDAE (p. 386). 




SESIIDAE (p. 386). 




TlXAEGERIIDAE (p. 387). 




SYXTOMIDAE (p. 388,. 




ZYGAEXIDAE (p. 390). 


< 


HlMAXTOPTERIDAE (p. 392). 




HETEROGYXIDAE (p. 392). 




PSYCHIDAE (p. 392). 




COSSIDAE (p. 395). 




ARBELIDAE (p. 396). 




CHRYSOPOLOMIDAE (p. 396). 


Heterocera 

(p. 366) 


HEPIALIDAE (p. 396). 
CALLIDULIDAE (p. 400). 
DREPAXIDAE (p. 400). 




LlMACODIDAE (p. 401). 




MEGALOPYGIDAE (p. 404). 




THYRIDIDAE (p. 404). 




LASIOCAMPIDAE (p. 405). 




EXDIMIMIDAE (p. 406). 




PTEROTHYSAXIDAE (p. 406). 




LYMAXTRIIDAE (p. 406). 




HYP.SIDAE (p. 408). 




ARCTIIDAE (p. 408). 




AHARISTIDAE (p. 410). 




GEOMETRIDAE (p. 411). 




XoCTl'IDAE (p. 414). 




EPICOPEIIDAE (p. 418). 




UUAXIIDAE (p. 419). 




Kl'iri.KMIHAE (p. 420). 




PYRALIDAE (p. 420). 




I'l 1 Hi'l'lliiKIIiAE (p. 426). 




A 1,1 ii i in \i; p. 126). 




TORTRICIDAK (p. 427). 




TIXKIDAE (p. 428). 


I 


EllIOCEPHALIDAE (p. 'I:;.'! . 


I 


. MlCRdl'TERYGIDAE (p. K!.~> 



SCHEME OF CLASSIFICATION 



XI 



Order. 



Sub-Order, Division, 
or Series. 



Family. 



DIPTERA 

(p. 438) 



Sub-Family or 
Tribe. 



Orthorrha- 
pha Nemo- 
cera 

(p. 455) 



Orthorrha- 
pha Bra 

chycera 
(pp. 455, 

478) 



Cyclorrha- 
pha As- 
chiza 

(pp. 455, 

494) 



Cyclorrha- 
pha Schi- 
zophora 

(pp. 456, 
503) 



I 



' CECIDOMYIIDAE (p. 458). 
MYCETOPHILIDAE (p. 462). 
BLEPHAROCERIDAK (p. 464). 
CULICIDAE. (p. 466). 
CHIRONOMIDAE (p. 468). 
ORPHXEPHILIDAE (p. 470). 
PSYOHODIDAE (p. 470). 
DIXIDAE (p. 471). 

TiPULiDAF ( pt y ch I' terinae (P- 4 " 2 )- 
T^jS \ Limnohiinae (p. 473). 

( Tipulinae (p. 475). 

BlBIONIDAE (p. 475). 
SlMULlIDAK (p. 477). 

(p. 478). 



STRATIOMYIDAE (p. 478). 
LEPTIDAE (p. 479). 
TABANIDAE (p. 481). 
ACANTHOMERIDAE (p. 483). 
THEREYIDAE (p. 484). 
SCEXOPIXIDAE (p. 484). 
NEMESTRINIDAE (p. 484). 

BOMBYLIIDAE (p. 485). 
ACROCERIDAE (p. 489). 
LOXCHOPTERIDAE (p. 490). 

MYDAIDAE (p. 491). 
ASILIDAE (p. 491). 
APIOCERIDAE (p. 492). 
EMPIDAE (p. 492). 
_ DOLICHOPIDAE (p. 493). 

' PHORIDAE (p. 494). 

PLATYPEZIDAE (p. 496). 
PIPUXCULIDAE (p. 496). 

CONOPIDAE (p. 497). 
I SYRPHIDAE (p. 498). 

MtTSCIDAE ACALYPTRATAE (p. 503). 
AXTHOMYIIDAE (p. 5*06). 

TACHIXIDAE (p. 507). 
DEXIIDAE (p. 510). 
SARCOPHAGIDAE (p. 510). 

MUSCIDAE (p. 511). 

OESTRIDAE (p. 514). 



Pupipara ( 

(pp. 456, - 
517) 



HIPPOP,OSCIDAE (p. 518). 
BRAULIDAE (p. 520). 



j STKEBLIDAE (p. 521). 
I XYCTERIBIIDAE (p. 



521). 



APHANIPTERA | PULICID , E ( , 

d<l<. 456, 522) \ 



THYSANO- f Terebrantia (p. 531). 

PTERA (p. 526) \ Tubulifera (p. 531). 



XII 



SCHEME OF CLASSIFICATION 



Order. 



Sub-Order. 



Series. 



HEMIPTERA 

(p. 532) 



Heteroptera 

(pp. 543, 
544) 



GYMNOCER- 

ATA (p. 

544) 



Homoptera 

(Pp. 543, 



' Anoplura 
(p. 599.) 



CRYPTOCER- 

ATA (p. 

562) 



Family. 
f PENTATOMIDAE (p. 545). 

COREIDAE (p. 546). 

BERYTIDAE (p. 548). 
LYGAEIDAE (p. 548). 
PYRRHOCORIDAE (p. 549). 
TINGIDAE (p. 549). 
ARADIDAE (p. 550). 
HEBRIDAE (p. 551). 
HYDROMETRIDAE (p. 551). 
HENICOCEPHALIDAE (p. 554). 
PHYMATIDAE (p. 554). 
REDUVIIDAE (p. 555). 
AEPOPHILIDAE (p. -559). 
CERATOCOMBIDAE (p. 559). 
CIMICIDAE (p. 559). 
ANTHOCORIDAE (p. 560). 

POLYCTENIDAE (p. 560). 

CAPSIDAE (p. 561). 
', SALDIDAE (p. 562). 

(GALGULIDAE (p. 562). 
NEPIDAE (p. 563). 
NAUCORIDAE (p. 565). 
BELOSTOMIDAE (p. 565). 
NOTONECTIDAE (p. 567). 
CORIXIDAE (p. 567). 



ClCATMDAE (p. 568). 
m I FULGOP.IUAE (ll. 574). 

TRIMERA (p. | MEMBKACIDA ^ (p . 5 f 6) . 

CERCOPIDAE (p. 577). 
I, JASSIDAE (p. 578). 



f PSYLLIDAE (i). 578). 

DlMEKA (P-l ApHIDAE(p : 1 581). 

[ ALEURODIDAE (p. 591). 



PEDICULIDAE (p. 599). 



CHAPTEE I 



HYMENOPTERA PETIOLATA COXTIXL'ED 

SERIES 2. TUBULIFERA OH CHRYSIDIDAE SERIES 3. ACULEATA- 

GENERAL CLASSIFICATION DIVISION I. ANTHOPHILA OR BEES 

THE First Series Parasitica ; of the Sub-Order Hymenoptera 
Petiolata was discussed in the previous volume. We now pass 
to the Second Series. 

Series 2. Hymenoptera Tubulifera. 

T/'ochanters undivided; the hind-body consisting of from three 
t<> Jive visible segments; the female with an ovipositor, 
usually retracted, transversely segmented, enveloping a Jine, 
pointed style. The larvae usually live in the cells of other 
Hijinenoptera. 

The Tubulifera form but a small group in comparison with 
Parasitica and Aculeata, the other two Series of the Sub-Order. 
Though of parasitic habits, they do not appear to be closely allied 
to any of the families of Hymenoptera Parasitica, though M. du 
Buysson suggests that they have some affinity with Proctotrypidae ; 
their morphology and classification have been, however, but little 
discussed, and have not been the subject of any profound investi- 
gation. At present it is only necessary to recognise one family, 
viz. Chrysididae or Ruby-wasps. 1 These Insects are usually of 
glowing, metallic colours, with a very hard, coarsely-sculptured 
integument. Their antennae are abruptly elbowed, the joints 
not being numerous, usually about thirteen, and frequently so 

1 Systematic monograph, Mocsary, Budapest, 1889. Account of the European 
Chrysididae, R. da Buysson in Andre, Sjiec. gen. Hym. vol. vi. 1896. 

VOL. VI IE B 



HV.MEXOPTERA 



CHAP. 



connected that it is not easy to count them. The abdomen is, 
in the great majority, of very peculiar construction, and allows 
the Insect to curl it completely under the anterior parts, so as to 
roll up into a, little ball ; the dorsal plates are very strongly 
arched, and seen from beneath form a free edge, while the ventral 
plates are of less hard consistence, and are connected with the 
dorsal plates at some distance from the free edge, so that the 
abdomen appears concave beneath. In the anomalous genus 

Cleptcs the abdomen 
is, however, similar 
in form to that of 
the Aculeate Hynien- 
optera, and has four 
or five visible seg- 
\ ments, instead of the 
three or four that 
are all that can be 
seen in the normal 
Chrysididae. The 

larvae of the Euby- 
iiies have the same 
number of segments 
as other Hymenoptera Petiolata. The difference in this re- 
sprct of the perfect Chrysididae from other Petiolata is due 
to a greater number of the terminal segments being indrawn 
so as to form the tube, or telescope-like structure from which 
the series obtains its name. This tube is shown partially 
extruded in Fig. 1 ; when fully thrust out it is seen to lie 
segmented, and three or four segments may be distinguished. 
The ovipositor proper is concealed within this tube ; it appears 
to be of the nature of an imperfect sting ; there being a very 
sharply pointed style, and a pair of enveloping sheaths ; the style 
really consists of a trough-like plate and two tine rods or spiculae. 
There are no poison glands, except in t'leptes, which form appears 
to come very near to the Aculeate series. Some of the Cbrysi- 

didii i occasions use the ovipositor as a sting, though it is only 

capaldc, of iiillid.ing a very minute and almost innocuous wound. 
Although none of the Huby-flies attain a large size, they arc 
usually very conspicuous on account of their gaudy or brilliant 
colours. They are amongst the most restless and rapid of Insects ; 




FIG. 1. Clirysis -iyiiita, ?. 



England. 



CHRYSIDIDAE 



they love the hot sunshine, and are difficult of capture. Though 
not anywhere numerous in species, they are found in most parts 
of the world. In Britain we have about twenty species. They 
usually frequent old wood or masonry, in which the nests of 
Aculeate Hymenoptera exist, or fly rapidly to and fro about the 
banks of earth where bees nest. Dr. Chapman has observed the 
habits of some of our British species. 1 He noticed Clirysis 
iijiiita nying about the cell of Odynerus parietum, a solitary 
wasp that provisions its nest with caterpillars; in this cell the 
(. '// i't/sis deposited an egg, and in less than an hour the wasp had 
sealed the cell. Two days afterwards this was opened and was 
found to contain a larva of Clirysis a quarter of an inch long, as 
well as the Lepidopterous larvae stored up by the wasp, but there 
was no trace of egg or young of the wasp. Six days after the 
egg was laid the Chrysis had eaten all the food and was full- 
grown, having moulted three or four times. Afterwards it formed 
a cocoon in which to complete its metamorphosis. It is, however, 
more usual for the species of Chrysis to live on the larva of the 
wasp and not on the food ; indeed, it has recently been positively 
stated that Chrysis never eats the food in the wasp's cell, but 
there is no ground whatever for rejecting the evidence of so care- 
ful an observer as Dr. Chapman. According to M. du Buys- 
son the larva of Clirysis will not eat the lepidopterous larvae, 
but will die in their midst .if the Odynerus larva does not de- 
velop ; but this observation probably relates only to such species 
as habitually live on Odynerus itself. The mother-wasp of 
Chrysis bidentata searches for a cell of Odynerus spinipes that has 
not been properly closed, and that contains a full-grown larva of 
that wasp enclosed in its cocoon. Having succeeded in its search 
the Chrysis deposits several eggs from six to ten ; for some 
reason that is not apparent all but one of these eggs fail to pro- 
duce young ; in two or three days this one hatches, the others 
shrivelling up. The young Chrysis larva seizes with its mouth 
a fold of the skin of the helpless larva of the Odynerus, and 
sucks it without inflicting any visible wound. In about eleven 
days the Clirysis has changed its skin four times, has consumed 
all the larva and is full-fed ; it spins its own cocoon inside that 
of its victim, and remains therein till the following spring, 
when it changes to a pupa, and in less than three weeks there- 

1 Enf. Mag. vi. 1869, p. 153. 



HYMENOl'TERA CHAP. 



after emerges a perfect Chrysis of the most brilliant colour, and 
if it be a female indefatigable in activity. It is remarkable 

O V 

that the larva of Chrysis is so much like that of Odynerus that 
the two can only be distinguished externally by the colour, the 
Odynerus being yellow and the (.'/trysts white ; but this is only 
one of the many cases in which host and parasite are extremely 
similar to the eye. Chrysis shanghaiensis has been reared from 
the cocoons of a Lepidopterous Insect Monema flavescens, family 
Limacodidae and it has been presumed that it eats the larva 
therein contained. All other Chrysids, so far as known, live at 
the expense of Hymenoptera (usually, as we have seen, actually 
consuming their bodies), and it is not impossible that C. shamj- 
haiensis really lives on a Hymenopterous parasite in the cocoon 
of the Lepidopteron. 

Parnopcs car tic a frequents the nests of Benibex rostra ta, a 
solitary wasp that has the unusual habit of bringing from time 
to time a supply of food to its young larva ; for this purpose it 
has to open the nest in which its young is enclosed, and the 
Pa mopes takes advantage of this habit by entering the cell and 
depositing there an egg which produces a larva that devours that 
of the L'emlex. The species of the anomalous genus Cleptes live, 
it is believed, at the expense of Tenthredinidae, and in all prob- 
ability oviposit in their cocoons which are placed in the earth. 

Series 3. Hymenoptera Aculeata. 

The females (whether workers or true females) provided with a 
sting: trochanters usually 'undivided- (monotrochous). Usual/// 
the antennae of the males with thirteen, of the females with 
twelve, joints (exceptions in ants numerous^. 

These characters only define this series in a very unsatisfac- 
tory manner, as no means of distinguishing the "sting" from the 
homologous structures found in Tubulifera, and in the Procto- 
trypid division of Hymenoptera Parasitica, have been pointed 
out. As the structure of the trochanters is subject to numerous 
exceptions, the classification at present existing is an arbitrary 
one. It would probably be more satisfactory to separate the 
Proctotrypidae (or a considerable part thereof) from the Para- 
sitica, and unite them with the Tubulifera and Aculeata in 
a <_;ival scries, characterised by the fact that the ovipositor is 



ACULEATA 



withdrawn into the body in a direct manner so as to be entirely 
internal, whereas in the Parasitica it is not withdrawn in this 
manner, but remains truly an external organ, though in numerous 
cases concealed by a process of torsion of the terminal seg- 
ments. If this were done it might be found possible to divide 
the great group thus formed into t\vo divisions characterised by 
the fact that the ovipositor in one retains its function, the egg 



FIG. 2. Diagram of upper sur- 
face of Priocnemis affi-iiis 9, 
Pompilidae. o, ocelli ; 7J 1 , 
pronotuni ; R z , mesonotum ; 
B 3 , scutellum of mesonotnm ; 
B 4 , post-scutelluni or middle 
part of metanotum ; B 5 , propo- 
deuni or median segment (see 
vol. v. p. 491) ; If', combing 
hairs, pecten, of front loot : C 1 , 
first segment of abdomen, here 
not forming a pedicel or stalk : 
Z) 1 , coxa; JL>' 2 , trochauter; Lt 3 , 
femur ; /> 6 , calcaria or spurs 
of bind leg : 1 to 15, iierv- 
ures of wings, viz. 1, costal ; 
2, post -costal ; 3, median ; 
4, posterior ; 5, stigma ; 
6, marginal ; 7, upper 
basal ; 8, lower basal ; 9, 9, 
cubital ; 10, the three sub- 
marginal ; 11, first recurrent ; 
12, second recurrent; }?,, 
anterior of hind, wing; 1J, 
median ; 15, posterior : I to 
XI, the cells, viz. I. upper 
basal ; II, lower basal ; III, 
marginal : IV, V. VI, first, 
second and third sub-mar- 
ginal ; VII, first discoidal ; 
VIII, third discoidal ; IX, 
second discoidal ; X, first 
apical ; XI, second apical. 



passing through it (Proctotrypidae and Tubulifera), while in the 
other the organ in question serves as a weapon of offence and 
defence, and does not act as a true ovipositor, the egg escaping 
at its base. It would, however, be premature to adopt so revolu- 
tionary a course until the comparative anatomy of the organs 
concerned shall have received a much greater share of attention. 1 
We have dealt with the external anatomy of Hymenoptera in 
1 For new views on this subject see note on p. 602. 




HYMENOPTERA 



CHAl'. 



Vol. V. ; so that here it is only necessary to give a diagram to 
explain the terms used in the descriptions of the families and 
sub-families of Aculeata, and to discuss briefly their characteristic 
structures. 

The Sting of the bee has been described in detail by Kraepelin. 
Sollmann, Carlet l and others. It is an extremely perfect me- 
chanical arrangement. The sting itself independent of the 
sheaths and adjuncts consists of three elongate pieces, one of 
them a gouge-like director, the other two pointed and barbed 
needles ; the director is provided with a bead for each of the 
needles to run on, these latter having a corresponding groove ; 
the entrance to the groove is narrower than its subsequent 
diameter, so that the needles play up and down on the director 

with facility, but cannot be 
dragged away from it; each 
needle is provided with an 
arm at the base to which 
are attached the muscles for 
its movement. This simple 
manner of describing the 
mechanical arrangement is, 
however, incomplete, inas- 
much as it includes no 
account of the means by 
which the poison is conveyed. 
This is done by a very com- 
plex set of modifications of 
all the parts; firstly, the 
director is enlarged at the 
anterior part to form a 

PIG. 3.-Stingofhee. A, One of the needles chamber, through which the 
separated ; a, the barbed point ; 6, piston ; needles play ; the needles are 

f 1 I'l i i T? ' I" "ii t.' i'ij fijfi Oi'n i\-\ r\-\~t r\*F -flii. 




a 



B 



arm. 



B, Tr;uis\ cisc section of tic i 

sting: dd, the two needles; e, bead for each Provided With a pro- 

the needles; /, director; g, jectillg piece, wllicll, aS the 

f poison. (After Carlet.) 



channel of pois 



needle moves, plays in the 



chamber of the, director, and forces downwards any liquid that 
may be therein ; the poison-glands open into the chamber, and 
the project ions on the needles, acting after the manner of a piston, 
carry the poison before them. The needles are so arranged on 

1 Ann. Sd. Nat. (7} ix. 1890, p. 1. 



ACULEATA STING LARVA 



the director that they enclose between themselves and it a space 
that forms the channel along which the poison flows, as it is 
carried forwards by the movement of the pistons attached to 
the needles. If the needles be thrust into an object quite as 
far as, or beyond, the point of the director much poison may be 
introduced into a wound, as the barbs are provided with small 
orifices placed one above the other, while if this be not the case 
much of the liquid will flow on the outside of the object. 

According to Carlet the poison of the bee is formed by the 
mixture of the secretions of two glands, one of which is acid and 
the other alkaline ; it is very deadly in its effects on other 
Insects. We shall see, however, that the Fossorial Hymenoptera, 
which catch and sting living prey for their young, frequently 
do not kill but only stupefy it, and Carlet states that in this group 
the alkaline gland is absent or atrophied, so that the poison con- 
sists only of the acid ; it is thus, he thinks, deprived of its lethal 
power. Moreover, in the Fossoria the needles are destitute of barbs, 
so that the sting does not remain in the wound. Bordas, however, 
states l that in all the numerous Hymenoptera he has examined, 
both acid and alkaline glands exist, but exhibit considerable differ- 
ences of form in the various groups. He gives no explanation 
of the variety of effects of the poison of different Aculeata. 

The larvae (for figure of larva of Bombus, see Vol. V. p. 488) 
are, without known exception, legless grubs, of soft consistence, 
living entirely under cover, being protected either in cells, or, 
in the case of social Hymenoptera, in the abodes of the parents. 
The larvae of Ants and fossorial Hymenoptera have the anterior 
parts of the body long and narrow and abruptly flexed, so that 
their heads hang down in a helpless manner. All the larvae of 
Aculeates, so far as known, are remarkable from the fact that the 
posterior part of the alimentary canal does not connect with the 
stomach till the larval instar is more or less advanced ; hence the 
food amongst which they live cannot be sullied by faecal matter. 
The pupa is invariably soft, and assumes gradually the colour 
of the perfect Insect. Almost nothing is known as to the 
intimate details of the metamorphosis, and very little as to the 
changes of external form. According to Packard "a period inter- 
venes between the stadium of the full-grown larva and that of 
the pupa, in which a series of changes he speaks of as semi-pupal 

1 C. . Ac. Paris, cxviii. 1894, p. 873. 



8 HYMENOPTERA 



CHAP. 



are passed through ; these, however, have not been followed out 
in the case of any individual, and it is not possible to form any 
final idea about them, but it seems probable that they are largely 
changes of external shape, in conformity with the great changes 
going on in the internal organs. Owing to the fragmentary 
nature of observations, much obscurity and difference of opinion 
have existed as to the metamorphosis of Aculeate Hynienoptera. 
Sir S. Saunders gives the following statement as to the larva of 
a wasp of the genus Psiliglossa, 1 just before it assumes the pupal 
form : " The respective segments, which are very distinctly indi- 
cated, may be defined as follows : The five anterior, including 
the head, are compactly welded together, and incapable of separate 
action in the pseudo-pupa state ; the third, fourth, and fifth 
bearing a spiracle on either side. The thoracical region termi- 
nating here, the two anterior segments are assignable to the 
development of the imago head, as pointed out by Katzeburg." 
This inference is not, however, correct. We have seen that in 
the perfect Insect of Petiolate Hymenoptera the first abdominal 
segment is fixed to the thorax, and Saunders' statement is in- 
teresting as showing that this assignment of parts already exists 
in the larva, but it in no way proves that the head of the imago 
is formed from the thorax of the larva. It has been stated 
that the larvae of the Aculeata have a different number of seg- 
ments according to the sex, but this also is incorrect. The 
difference that exists in the perfect Insects in this respect is due 
to the withdrawal of the terminal three segments to the interior 
in the female, and of two only in the male. The larva consists 
of fourteen segments, and we find this number distributed in the 
female perfect Insect as follows : one constitutes the head, four 
segments the thorax and propodeum, followed by six external seg- 
ments of the restricted abdomen, and three for the internal structures 
of the abdomen. This agrees with Corel's statement that in the 
ants the sting is placed in a chamber formed by three segments. 
The development of the sting of the common bee has been 
studied by Dewitz. 2 It takes place in the last larval stage. 
Although nothing of the organ is visible externally in the adult 
larva, yet if such a larva be placed in spirit, there can be seen 
within the skin certain small appendages on the ventral surface 
of the penultimate and antepenultimate abdominal segments 
Trans, cut. Sac. London, 1873, p. 408. - Zcitschr. iviss. Zool. xxv. 1875, p. 184. 



ACULEATAy DEVELOPMENT 




( Fig. 4, A) placed two on the one, four on the other ; these are the 

rudiments of the sting. In the course of development the 

terminal three segments are taken into the body, and the external 

pair of the appendages of the twelfth 

!>dy segment (the ninth abdominal) 

become the sheaths of the sting, and 

the middle pair become the director ; 

the pair of appendages on the eleventh 

segment give rise to the needles or 

spiculae. The sting -rudiments at an 

earlier stage (Fig. 4, C) are masses of 

hypodermis connected with tracheae ; 

there is then but one pair on the 

twelfth segment, and this pair coalesce 

to form a single mass ; the rudiments G 

of the pair that form the director are FlG . 4. Development of sting of 

differentiated secondarily from the the bee : A and C, ventral ; B, 

side view. A, End of abdomen 

primary pair ot these masses 01 hypo- 
dermis. A good deal of discussion 
has taken place as to whether the 
component parts of the sting 
gonapophyses are to be considered as 
modifications of abdominal extremities 
(i.e. abdominal legs such as exist in 
Myriapods). Heymons is of opinion 
that this is not the case, but that the 
leg-rudiments and gonapophysal rudi- 
ments are quite distinct. 1 The origin 
of the sting of Hymenoptera (and of the ovipositor of parasitic 
Hymenoptera) is very similar to that of the ovipositor of Locusta 
(Vol. V. p. 315 of this work), but there is much difference in the 
history of the development of the rudiments. 

Dewitz has also traced the development of the thoracic 
appendages in Hymenoptera. 2 Although no legs are visible in 
the adult larva, they really arise very early in the larval life from 
masses of hypodermis, and grow in the interior of the body, so 
that when the larva is adult the legs exist in a segmented though 
rudimentary condition in the interior of the body. Dewitz's 
study of the wing-development is less complete. 

i. Jahrb. xxiv. 1896, p. 192. - Zeitscltr. iciss. Zool. xxx. 1878, p. 78. 



of adult larva : a, b, c, d, the 
last four segments, c being the 
eleventh body segment, 11 ; 
b bearing two pairs, and c one 
pair, of rudiments. B, Tip of 
abdomen of adult bee : 9, the 
ninth, d, the tenth body seg- 
ment. C, Rudiments in the 
early condition as seen within 
the body : c, first pair ; b, the 
second pair not yet divided into 
two pairs ; b", c', commence- 
ment of external growths from . 
the internal projections. (After 
Dewitz.) 



I O HYMENOPTERA CHAP. 

Four primary divisions of Aculeates are generally recognised, 
viz. Anthophila (Bees), Diploptera (Wasps), Fossores (Solitary 
Wasps), Heterogyna (Ants). Though apparently they are natural, 
it is impossible to define them by characters that are without 
some exceptions, especially in the case of the males. Ashmead 
has recently proposed l to divide the Fossores ; thus making five 
divisions as follows : 

Body with more or less of the hairs on it plumose 1. Anthophila. 

Hairs of body not plumose. 

Pronotum not reaching hack to tegulae . 2. Entomophila 

[ = Fossores part] 
Pronotum reaching hack to tegulae. 

Petiole (articulating segment of abdomen) simple without scales or 

nodes. 
Front wings in repose with a fold making them narrow 

3. Diploptera. 

Front wings not folded 4. Fossores [part]. 

Petiole with a scale or node (an irregular elevation on the upper side) 

5. Heterogyna. 

We shall here follow the usual method of treating all the 
fossorial wasps as forming a single group, uniting Ashmend's 
Entomophila and Fossores, as we think their separation is only 
valid for the purposes of a table ; the Pompilidae placed by the 
American savant in Fossores being as much allied to Entomo- 
phila as they are to the other Fossores with which Ashmead 
associates them. 

Division I. Anthophila or Apidae Bees. 

Some of the hairs of the body 'plumose ; parts of the mouth elon- 
gated, sometimes to a great extent, so as to form a protrusible 
apparatus, usual I;/ tubular with a very flexible tip. Basal 
joint of hind foot elongate. No wingless adult forms ; in 
some cases societies are formed, and then barren females called 
workers exist in, gr<'<// numbers, and carry on the industrial 
operations of the community. Food always derived from the 
vegetable kingdom, or from other Bees. 

There are about 150 genera and 1500 species of bees at 
present known. Some call the division Mellifera instead of 
Anthophila. The term Apidae is used by some authorities to de- 
note all the bees, while others limit this term to one of the families 

1 Proc. ent. Soc. jr<is/i//iit/nn, iii. 1896, p. 334. 



ANTHOPHILA BEES 



I I 



or sub-divisions. The bees are, as a rule, distinguished from 
other Hymenoptera by the hairs, by the great development of the 
mouth parts to form a proboscis (usually, but not correctly, called 
tongue), and by the modification of the hind-legs ; but these 
distinctive characters are in some of the species exhibited in so 
minor a degree of perfection that it is not easy to recognise these 
primitive forms as Anthophila. A few general remarks on the 
three points mentioned will enable the student to better appreciate 
the importance of certain points we shall subsequently deal with. 
The bees are, as a rule, much more covered with hair than 
any other of the Hymenoptera. Saunders l states that he has 
examined the structure of the. 
hairs in all the genera of British 
Aculeata, and that in none but 
the Anthophila do branched and 
plumose hairs occur. The func- 
tion of this kind of hairs is 
unknown; Saunders suggests 1 that 
they may be instrumental in the 
gathering of pollen, but they 
occur in the parasitic bees as 
well as in the males, neither of 
which gather pollen. The variety 
of the positions they occupy on 
the body seems to offer but little 
support to the suggestion. Not 
all the hairs of the bee's body 

are r>lm IOSP some are simule FIG. 5. Hairs of Bees : A, simple hair 

from abdomen of Osmia; B, spiral 

as shown in Fig. 5, A, and this is 
specially the case with the hairs 
that are placed at the edges of 
the dilated plates for carrying 

pollen. In some forms there is an extensive system of simple 
hairs all over the body, and the " feathers " are distributed 
between these ; and we do not see any reason for assuming that 
the feathered are superior to the simple hairs for gathering and 
carrying pollen. Some bees, e.g. Prosopis, Ceratina, have very 
little hair on the body, but nevertheless some plumose hairs 
are always present even though they be very short. 

1 Trans, cnt. Soc. 1878, p. 169. 



1 


f 


\ 




\ 




\ 








\ 




\ 


, 


\ 




\ 


o 




'/ 


\l 

v 

v 

V 

v> 

y 




\ 


/ 


C\ 




\ 






hair from abdomen of Jlegacfiile ; C, 
plumose hair from thorax of Mega- 
chile ; D, from thorax of Andrena 
dorsata ; E, from thorax of Prosopis. 



I 2 



HYMENOPTERA 



CHAP. 



The hind-legs of bees are very largely used in the industrial 
occupations of these indefatigable creatures; one of their chief 
functions in the female being to act as receptacles for carrying 
pollen to the nest : they exhibit, however, considerable diversity. 
The parts most modified are the tibia and the first joint of the 
hind-foot. Pollen is carried by other parts of the body in many 
bees, and even the hind-leg itself is used in different ways for the 

purpose : sometimes the outer face 
of the tibia is highly polished and 
its margins surrounded by hair, in 
which case pollen plates are said to 
exist (Fig. 6, A) ; sometimes the first 
joint of the tarsus is analogous to 
the tibia both in structure and 
function ; in other cases the hind- 
legs are thick and densely covered 
with hair that retains the pollen 
between the separate hairs. In this 
case the pollen is carried home in a 
dry state, while, in the species with 
pollen plates, the pollen is made 
into a mass of a clay-like consist- 
ence. 1 The legs also assist in arrang- 
ing the pollen on the other parts of 
the body. The males do not carry 

Fir,. 6. A. Worker of the honey-bee po ll en am l though their hind-legS 
(Aixsmellijim), with pollen plates - -,-, 

la.i.-n ; B, basal portions of a are also highly modified, yet the 
middle-leg (trochanter with part of modifications do not agree with 

coxa and ot femur) with plumose 

hairs and grains of pollen ; c, one those of the female, and their fiuic- 
hair bearing pollen-grams. 




ftre in ^ probability sexu .,l. 

The parasitic bees also do not carry pollen, and exhibit another 
series of structures. The most interesting case in this series of 
modifications is that found in the genus Apis,Vfhere the hind-leg 
of male, female, and worker are all different (Fig. 25); the limb 
in the worker being highly modified for industrial purposes. 
This case has been frequently referred to, in consequence of the 
difficulty that exists in connection with its heredity, for the 

1 The mode of wetting the pollen is nut clear. Woltl' says it is done 1))- an i-xn- 
dation from the tibia; 11. M idler by ;idmi,\t lire of nectar from tin.' bee's mouth. 
The, latter view is more probably correct. 



BEES PROBOSCIS I 3 



structure exists in neither of the parents. It is, in fact, a case 
of a very special adaptation appearing in the majority of the 
individuals of each generation, though nothing of the sort occurs 
in either parent. 

The proboscis of the bee l is a very complex organ, and in its 
extremely developed forms exhibits a complication of details and 
a delicacy of structure that elicit the admiration of all who study 
it. In the lower bees, how r ever, especially in Prosopis, it exists 
in a comparatively simple form (Fig. 9, B, C), that differs but little 
from what is seen in some Vespidae or Fossores. The upper lip 
and the mandibles do not take any part in the formation of the 
bee's proboscis, which is consequently entirely made up from the 
lower lip and the maxillae, the former of these two organs ex- 
hibiting the greatest modifications. The proboscis is situate on 
the lower part of the head, and in repose is not visible ; a portion, 
and that by no means an inconsiderable one, of its modifications 
being for the purpose of its withdrawal and protection when not 
in use. For this object the under side of the head is provided 
with a very deep groove, in which the whole organ is, in bees 
with a short proboscis, withdrawn ; in the Apidae with a long pro- 
boscis this groove also exists, and the basal part of the proboscis 
is buried in it during repose, while the other parts of the elon- 
gate organ are doubled on the basal part, so that they extend 
backwards under the body, and the front end or tip of the tongue 
is, when in repose, its most posterior part. 

For the extrusion of the proboscis there exists a special 
apparatus that comes into play after the mandibles are unlocked 
and the labrum lifted. This extensive apparatus cannot be satis- 
factorily illustrated by a drawing, as the parts composing it are 
placed in different planes ; but it may lie described by saying 
that the cardo, or basal hinge of the maxilla, changes from an 
oblique to a vertical position, and thrusts the base of the pro- 
boscis out of the groove. The maxillae form the outer sheath of the 
proboscis, the lower lip its medial part (see Figs. 7 and 9 ) ; the 
base of the lower lip is attached to the submentum, which rises 
with the cardo so tha.t labium and maxillae are lifted together : 
the co-operation of these two parts is effected by an angular piece 
called the lorum, in which the base of the submentum rests ; the 

1 In studying the proboscis the student will do well to take a Bombus as an 
example ; its anatomy being more easily deciphered than that of the honey-bee. 



HYMEN OPT ERA 



CHAP. 



submentum is articulated with the mentuui iu such a manner that 
the two can either be placed in planes at a right angle to one 
another, or can be brought into one continuous plane, and by this 
change of plane the basal part of the tongue can also be thrust 
forwards. There is considerable variety in the lengths of these 
parts in different genera, and the lorum varies in shape in accord- 
ance with the length of the subinentuin. The lorum is a 

o 

peculiar piece, and its mechanical adaptations are very remark- 
able ; usually the base of the submentum rests in the angle 
formed by the junction of the two sides of the lorum, but in A'ylu- 
copa, where the submentum is unusually short, this part reposes 




7. 



FIG. 7. Side view of basal portions of proboscis of Bombiis. a, Epipharyugeal sclerites ; 
6, arrow indicating the position of the entrance to pharynx, which is concealed by 
the epipharynx, c ; d, hypopharyugeal sclerites ; e, vacant space between the scales 
of the maxillae through which the nectar comes : /, lobe ; /', stipes ; </, cardo of 
maxilla : h, encephalic pillar on which the cardo swings ; i, angle of junction of lores 
and submentum lorum ; k, mentum ; I, base of labial palp ; m, maxillary palp. 

in a groove on the back of the lorum, this latter having a very 
broad truncated apex instead of an angular one; in the condition 
of repose the apex of the lorum rests in a notch on the middle of 
the back of the oral groove, and in some of the forms with elon- 
gate submentum, this depression is transformed into a deep hole, 
or even a sort of tunnel, so as to permit the complete stowing 
away of the base of the tongue, which would otherwise be pre- 
vented by the long submentum; another function of the lorum 
appears to be that, as it extends, its arms have an outward thrust, 
and so separate the maxillae from the labium. In addition to 
these parts then- are also four elongate, slender sclerites that are 
only brought into view on dissection, and that no doubt assist in 
correlating the movements of the parts of the mouth and hypo- 



BEES PROBOSCIS I 5 



pharynx ; one pair of these strap-like pieces extends backwards 
from the two sides of the base of the epipharynx ; Huxley called 
them selerites of the oesophagus ; a better name would be 
epipharyngeal selerites (Fig. 7, .) : the other pair pass from the 
terminations of the epipharyngeal selerites, along the front face of 
the hypopharynx, down to the mentum, their lower parts being 
concealed by the stipites of the maxillae ; these are the hypo- 
pharyngeal selerites, and we believe it will prove that they play 
a highly important part in deglutition. When the labrum of a 
bee is raised and the proboscis depressed, the epipharynx is seen 
hanging like a curtain from the roof of the head ; this structure 
plays an important part in the act of deglutition. The entrance 
to the pharynx, or commencement of the alimentary canal, is 
placed below the base of the epipharynx. As we are not aware 
of any good delineations of the basal parts of the proboscis we 
give a figure thereof (Fig. 7). The maxillae in the higher bees 
are extremely modified so as to form a sheath, and their palpi 
are minute ; in the lower bees the palpi have the structure usual 
in mandibulate Insects. 

Eeturning to the consideration of the lower lip, we find that 
there is attached to the mentum a pair of elongate organs that 
extend forwards and form a tube or sheath, enclosed by the 
maxillary sheath we have previously mentioned ; these are the 
greatly modified labial palpi, their distal parts still retaining the 
palpar form ; and in the lower bees the labial palpi are, like the 
maxillary, of the form usual in mandibulate Insects. Between 
the labial palps and the central organ of the lip there is attached 
a pair of delicate organs, the paraglossae. 

There remains for consideration the most remarkable part of 
the proboscis, the long, delicate, hairy organ which the bee 
thrusts out from the tip of the shining tube formed by the labial 
palps and the maxillae, described above, and which looks like 
a prolongation of the mentum. This organ is variously called 
ligula, lingua, or tongue. 1 We prefer the first of these names. 

According to Breithaupt and Cheshire the structure of the 
ligula is highly remarkable ; it is a tube (filled with fluid from 
the body cavity), and with a groove underneath caused by a large 
part of the circumference of the tube being invagiiiated ; the 

1 Leuckart proposed the term lingula ; but the word gives rise to the impression that 
it is a mistake for either lingua or ligula. Packard calls the part " hypopharynx." 



i6 



IIVMENOPTERA 



CHAP. 



invaginated part can be thrust out by increase of the pressure of 

the fluid in the tube. A portion 
of the wall of the invaginate part 
is thickened so as to form a chiti- 
nous rod. 

This description vail suffice for 
present purposes, as the other parts 
of the mouth will be readily re- 
cognised by the aid of figure 9, A, 
B, C. In the exquisitely endowed 
South American genus Euylossa (Fig. 
18), the proboscis is somewhat longer 
than the whole of the body, so that 
its tip in repose projects behind 
the body like a sting. 

The correct nomenclature of the 
parts connected with the lower lip 
is not definitely settled, authorities 
not being agreed on several points. 

FIG. 8. -Transverse section of ligula Tlle wll le f the P^OScis is USUally 

of honey-bee, diagram atie. A, called the tongue : this, however, is 
With the long sac invaginate. B. . ., ,. ,. 

evagh.ate : a , chitinous envelope admittedly an erroneous application 

with the bases of the hairs ; h, o f ^1^ term. The terminal deli- 
rod ; i; groove of rod ; d, lumen . . 

due in A to invagiuation of the cate, elongate, flexible organ is by 

rod, iii B to its evagination ; u, gome ca n ec j t he tongue ; but this 
nerve ; tr, trachea. . ... 

again is wrong : the lingua in 

Insects is the hypopharynx ; this part is developed in a peculiar 
manner in bees, but as it is not tongue-like in shape, the term 
lingua is not suitable for it, and should be dismissed altogether 
from the nomenclature of the bee's trophi ; it is used at present 
in two different senses, both of which are erroneous. We see no 
objection to describing the flexible apical portion of the proboscis 
as the ligula. The lorum is probably a special part peculiar to 
the higher bees ; according to Saunders it is not present as a 
specialised part in some of the primitive forms. 1 The application 
of the terms mentuin, submentum and hypoglottis is open to tin- 
same doubts that exist with regard to them in so many other 




1 For figures and descriptions of the proboscides of British bees, refer to E. Saun- 
ders, Jour. Linn. Xoe. xxiii. 1890, pp. 410-432, plates III.-X. : and for details of 
the minute structure ami function to Cheshire, /lees and Bee-keepiny, vol. i. 



BEES PROBOSCIS 



Insects, and we have omitted the term hypogiottis altogether, 
though some may think the mentum entitled to that name. 

The way in which the proboscis of the bee acts has been very 
largely discussed, with 
special reference to the 
question as to whether it is 
a sucking or a licking action. 
It is impossible to consider 
either of these terms as 
applicable. The foundation 
of the action is capillary 
attraction, by which, and by 
slight movements of increase 
and contraction of the 
capacity of various parts, the 
fluid travels to the cavity 
in front of the hypopharynx : 
here the scales of the 
maxillae leave a vacant 
space, (Fig. 7, e) so that a 
cup or cavity is formed, the 
fluid in which is within 
reach of the tip of the depen- 
dent epipharynx (c), which 
hangs down over the front 
of the hypopharynx (and is 
so shaped that its tip covers 
the cup) ; it is between 
these two parts that the 
fluid passes to reach the 
pharynx. It is no doubt 
to slight movements of the 
membranous parts of the 
hypopharynx and of the 
epipharynx that the further progress of the nectar is due, aided 
by contraction and expansion of the pharynx, induced by muscles 
attached to it. It should be recollected that in addition to the 
movements of the head itself, the hypopharynx is constantly 
changing its dimensions slightly by the impulses of the fluid of 
the general body cavity; also that the head changes its position, 
VOL. vi c 




a- 



FIG. 9. A, Proboscis of a "long-tongued " bee, 
Anthophora pilipes ; B, lower, C, upper view 

of proboscis of an " obtuse-tougued " bee 
Prosopis pubescent, u, Labruni ; b, stipes 
r, palpiger ; d, scale : f, lobe ; g, palpus 
h, cardo, of maxilla : i, lorum ; k, submentum 
/, mentum ; m, labial palp ; //, paraglossa 
o, ligula ; p. tip of ligula (with "spoon " at tip 
and some of the hairs more magnified) ; q, 
hypopharyngeal sclerites. 



1 8 HYMENOPTERA 



CHAP. 



and that the proboscis is directed downwards as well as forwards. 
Those who wish to pursue this subject should refer to the works 
of Breithaupt l and Cheshire. 

The other external characters of the Bees call for little re- 
mark. The pronotum is never very large or much prolonged in 
front, and its hind angles never repose on the tegulae as they do 
in the wasps, 2 but extend backwards below the tegulae. The hind 
body is never narrowed at the base into an elongate pedicel, as it 
so frequently is in the Wasps and in the Fossors ; and the pro- 
podeum (the posterior part of the thorax) is more perpendicular 
and rarely so largely developed as it is in the Fossors ; this last 
character will as a rule permit a bee to be recognised at a glance 
from the fossorial Hymenoptera. 

Bees, as every one knows, frequent flowers, and it is usually 
incorrectly said that they extract honey. They really gather 
nectar, swallow it, so that it goes as far as the crop of their ali- 
mentary canal, called in English the honey-sac, and is regurgi- 
tated as honey. Bertrand states that the nectar when gathered 
is almost entirely pure saccharose, and that when regurgitated it 
is found to consist of dextrose and levulose : 3 this change appears 
to be practically the conversion of cane- into grape-sugar. A 
small quantity of the products of the salivary glands is added, 
and this probably causes the change alluded to ; so that honey 
and nectar are by no means synonymous. According to Cheshire 
the glandular matter is added while the nectar is being sucked, 
and is passing over the middle parts of the lower lip, so that 
the nectar may be honey when swallowed by the bee. In 
addition to gathering nectar the female bees are largely occupied 
in collecting pollen, which, mixed with honey, is to serve as 
food for the colony. Many, if not all, bees eat pollen while 
collecting it. The mode in which they accumulate the pollen, 
and the mechanism of its conveyance from hair to hair till it 
reaches the part of the body it must attain in order to be removed 
for packing in the cells, is not fully understood, but it appears 
to be accomplished by complex correlative actions of various parts ; 
the head and the front legs scratch up the pollen, the legs move 
with great rapidity, and the pollen ultimately reaches its desti- 
nation. The workers of the genus Apis, and of some other social 

1 I'.rritli.'iiipt, Arch. Natunjrs. lii. I><1. i. ISSt!, \>. -17. 
2 See Fig. 2fi. p. 71. 3 Bull. Mas. Paris, i. 1895, p. 38. 



BEES HABITS 1,9 



bees, have the basal joint of the hind foot specially adapted to 
deal with pollen (Fig. 25, 2). We have already mentioned the 
modifications of the legs used for its conveyance, and need here 
only add that numerous bees the Dasygastres carry the pollen 
by aid of a special and dense clothing of hairs on the underside 
of the abdomen. 

The buzzing of bees (and other Insects) has been for long a 
subject of controversy : some having maintained that it is parti- 
ally or wholly due to the vibration of parts connected with the 
spiracles, while others have found its cause in the vibrations of 
the wings. According to the observations of Perez and Bellesme, 1 
two distinct sounds are to be distinguished. One, a deep noise, 
is due to the vibration of the wings, and is produced whenever a 
certain rapidity is attained ; the other is an acute sound, and is 
said to be produced by the vibrations of the walls of the thorax, 
to which muscles are attached ; this sound is specially evident 
in Diptera and Hymeuoptera, because the integument is of the 
right consistence for vibration. Both of these observers agree that 
the spiracles are not concerned in the matter. 

The young of bees are invariably reared in cells. These 
(except in the case of the parasitical bees) are constructed by 
the mothers, or by the transformed females called workers. 
The solitary bees store the cells with food, and close up each cell 
after having laid an egg in it, so that in these cases each larva 
consumes a special store previously provided for it. The social 
bees do not close the cells in which the larvae are placed, and 
the workers act as foster-mothers, feeding the young larvae after 
the same fashion as birds feed their nestling young. The food is 
a mixture of honey and pollen, the mixing being effected in various 
ways and proportions according to the species ; the honey seems 
to be particularly suitable to the digestive organs of the young 
larvae, and those bees that make closed cells, place on the outside 
f the mass of food a layer more thickly saturated with honey, 
and this layer the young grub consumes before attacking the 
drier parts of the provisions. The active life of the larva is 
quite short, but after the larva is full-grown it usually passes a 
more or less prolonged period in a state of quiescence before 
assuming the pupal form. The pupa shows the limbs and other 
parts of the perfect Insect in a very distinct manner, and the 
1 C.R. Ac. Paris, Ixxxvii. 1878, pp. 378 and 535. 



o 



20 HYMENOPTERA CHAP. 

development of the imago takes place gradually though quickly. 
Some larvae spin cocoons, others do not. 

A very large number of bees are parasitic in their habits, 
laying an egg, or sometimes more than one, in the cell of a work- 
ing bee of some species other than their own ; in such cases the 
resulting larvae eat and grow more quickly than the progeny of 
the host bee, and so cause it to die of starvation. It has been 
observed that some of these parasitic larvae, after eating all the 
store of food, then devour the larva they have robbed. In 
other cases it is possible that the first care of the parasitic larva, 
after hatching, is to eat the rival egg. 

The taxonomy of bees is in a very unsatisfactory state. The 
earlier Hyinenopterists were divided into two schools, one of 
which proposed to classify the bees according to their habits, 
while the other adopted an arrangement depending on the length 
of the parts of the mouth, the development of the palpi, and the 
form and positions of the organs for carrying pollen. Neither 
of these arrangements was at all satisfactory, and some ento- 
mologists endeavoured to combine them, the result being a 
classification founded partly on habits and partly on certain 
minor structural characters. This course has also proved unsatis- 
factory ; this is especially the case with exotic bees, which have 
been placed in groups that are defined by habits, although very 
little observation has actually been made on this point. 
Efforts have recently been made to establish an improved classifi- 
cation, but as they relate solely to the European bees they are 
insufficient for general purposes. 

The more important of the groups that have been recognised 
are (1) the Obtusilingues, short-tongued bees, with the tip of the 
lingua bifid or broad; (2) Acutilingues, short-tongued bees, with 
acute tip to the tongue ; these two groups being frequently treated 
of as forming the Andrenidae. Coming to the Apidae, or the 
bees with long and folded tongues, there have been distinguished 
(3) Scopulipedes, bees carrying pollen with their feet, and (4) 
I>asygastres, those that carry it under the abdomen; some of the 
p;irasitie and other forms have been separated as (5) Denudatae 
(or ( 'iietilinae) : the l>ombi and the more perfectly social bees 
forming another group, viz. (G) Sociales. A group Andrenoides, 
or Pan urg ides, was also proposed for certain bees considered to 
belong to the Apidae though exhibiting many points of resent- 



i BEES ARCHIAPIDES 21 

blance with the Andrenidae. This arrangement is by no means 
satisfactory, but as the tropical bees have been but little collected, 
and are only very imperfectly known, it is clear that we cannot 
hope for a better classification till collections have been very 
much increased and improved. The arrangement adopted in 
Dalla Torre's recent valuable catalogue of bees 1 recognises no less 
than fourteen primary divisions, but is far from satisfactory. 

The two genera Prosopis and Spliecodcs have been recently 
formed into a special family, AKCHIAPIDAE, by Friese, 2 who, how- 
ever, admits that the association is not a natural one. The 
term should be limited to Prosopis and the genera into which it 
has been, or shortly will be, divided. The primitive nature of 
the members of this genus is exhibited in all the external 
characters that are most distinctive 
of bees; the proboscis (Fig. 9, B, C), is 
quite short, its ligula being very short, 
and instead of being pointed having 
a concave front margin. The body 
is almost bare, though there is some 
very short feathered plumage. The 
hind legs are destitute of modifica- 
tions for industrial purposes. Owing 
to these peculiarities it was for 

long assumed that the species Of FIG. 10. Prosopis signata. Cam- 

PfOSOpis must be parasites. This bridp A, Female; B front of 

head of female ; C, of male. 

is, however, known not to be the 

case so far as many of the species are concerned. They form 
cells lined with a silken membrane in the stems of brambles 
and other plants that are suitable, or in burrows in the 
earth, or in the mortar of walls ; individuals of the same species 
varying much as to the nidus they select. The food they store 
in these cells is much more liquid than usual, and has been 
supposed to be entirely honey, since they have no apparatus for 
carrying pollen. Mr. E, C. L. Perkins has, however, observed 
that they swallow both pollen and nectar, brushing the first- 
named substance to the mouth by aid of the front legs. He 

1 Catalogus Hymcnoptcrorum, Leipzig, 10 vols. 1892-96 ; Bees, vol. x. 

- Zool. Jahrb. Syst. iv. 1891, p. 779. This paper is a most valuable summary 
of what is known as to the habits of European solitary bees, but is less satisfactory 
from a systematic point of view. 




22 IIYMENOPTERA CHAP. 

has ascertained that a few of the very numerous Hawaiian species 
of the genus are really parasitic on their congeners : these parasites 
are destitute of a peculiar arrangement of hairs on the front 
legs of the female, the possession of which, Ly some of the non- 
parasitic forms, enables the bee to sweep the pollen towards its 
mouth. These observations show that the structural peculiarities 
of I'rosojtis are correlative with the habits of forming a peculiar 
lining to the cell, and of gathering pollen by the mouth and 
conveying it by the alimentary canal instead of by external parts 
of the body. Prosopis is a very widely distributed genus, and 
very numerous in species. We have ten in Britain ; several of 
them occur in the grounds of our Museum at Cambridge. 

The species of the genus Colletes are hairy bees of moderate 
size, with a good development of hair on the middle and posterior 
femora for carrying pollen. They have a short, bilobed ligula 
like that of wasps, and therein differ from the Andrenae, which 
they much resemble. With Prosopis they form the group Obtusi- 
lingues of some taxonomists. They have a manner of nesting 
peculiar to themselves ; they dig cylindrical burrows in the 
earth, line them with a sort of slime, that dries to a substance 
like gold-beater's skin, and then by partitions arrange the 
burrow as six to ten separate cells, each of which is filled 
with food that is more liquid than usual in bees. Except in 
regard to the ligula and the nature of the cell-lining, Colletes has 
but little resemblance to Prosopis; but the term Obtusilingurs 
may be applied to Colletes if Prosopis be separated as Archiapidae. 
We have six species of Colletes in Britain. 

Sphecodes is a genus that has been the subject of prolonged 
< inference of opinion. The species are rather small shining 
bees, with a red, or red and black, abdomen, almost with- 
out pollen-collecting apparatus, and with a short but pointed 
ligula. These characters led to the belief that the Insects are 
parasitic, or, as they are sometimes called, cuckoo-bees. But 
evidence could not be obtained of the fact, and as they were seen 
to make burrows it was decided that we have in Spheco</<x 
examples of industrial bees extremely ill endowed for their work. 
Recent observations tend, however, to prove that fyrfiecodes are to 
a large extent parasitic at the expense of bees of the genera 
I ful id us and A ml re mi. Breitenbach has taken S. rubicundus out 
of the brood-cells of Halictus quadricinctus ; and on one of the few 



BEES ANDRENIDES 23 




occasions on which this bee has been found in Britain it was in 
circumstances that left little doubt as to its being a parasite of 
Andrcna niyroaenca. Marchal l has seen S. subquadratus fight 
with Halietas malachurus, 
and kill it previous to taking 
possession of its burrows ; 
and similar observations 
have been made by Ferton. 
As the older observations of 
Smith, Sichel, and Friese 
leave little doubt that Sphe- 
codes are sometimes indus- 
trial bees, it is highly prob- 
able that we have in this 

FIG. 11. Sphecodes gibbus 9- Britain. 
genus the interesting con- 
dition of bees that are sometimes parasitic, at other times 
not ; but so much obscurity still prevails as to the habits of 
Sphecodes that we should do well to delay accepting the theories 
that have been already based on this strange state of matters. 2 
Friese states that in Sphecodes the first traces of collecting- 
apparatus exist ; and, accepting the condition of affairs as being 
that mentioned above, it is by no means clear whether we have 
in Sphecodes bees that are abandoning the parasitic habit or com- 
mencing it ; or, indeed, whether the condition of uncertainty 
may not be a permanent one. It is difficult to decide as to 
what forms are species in Sphecodes owing to the great variation. 
The Hymenopterist Forster considered that 600 specimens sub- 
mitted to him by Sichel represented no less than 140 species, 
though Sichel was convinced that nearly the whole of them 
were one species, S. gibbus. It has recently been found that the 
male sexual organs afford a satisfactory criterion. The position 
of Sphecodes in classification is doubtful. 

The great majority of the species of short-tongued bees found in 
Britain belong to the genera Andrena and Halictus, and with some 
others constitute the ANDRENIDES of many writers. Halictus 
includes our smallest British bees. Their economy escaped the 
earlier observers, but has recently been to some extent unravelled 
by Smith, Fabre, Nicolas, Verhoeff, and others, and proves to be 

1 Bull. Soc. ent. France, 1894, p. cxv. 
- Marchal, Rev. Sci. 15th February 1890, and Ferton, t.c. 19th April. 



24 HYMENOFTKRA CHAP. 

of great interest and variety. Fabre observed H. lineolatus and 
H. sexcinctus 1 under circumstances that enabled him to give them 
continuous attention, whenever requisite, throughout a whole year. 
These bees are to a certain extent social ; they are gregarious ; each 
bee works for its own progeny, but there is collaboration between 
members of a colony, inasmuch as a piece of general work is 
undertaken from which more families than one derive benefit. 
This common work is a gallery, that, ramifying in the earth, 
gives access to various groups of cells, each group the production 
of a single Halictus ; in this way one entrance and one corridor 
serve for several distinct dwellings. The work of excavation is 
carried on at night. The cells are oval, and are covered on the 
interior with a delicate waterproof varnish ; Fabre considers this 
to be a product of the salivary glands, like the membrane we 
noticed when speaking of Colletes. In the south of France both 
sexes of these species are produced from the nests in September, 
and then the males are much more numerous than the females ; 
when the cold weather sets in the males die, but the females 
continue to live on in the cells underground. In the following 
spring the females come out and recommence working at the 
burrows, and also provision the cells for the young ; the new 
generation, consisting entirely of females, appears in July, and 
from these there proceeds a parthenogenetic generation, which 
assumes the perfect form in September, and consists, as we have 
above remarked, in greater part of males. Perez, 2 however, 
considers that Fabre's observations as to the parthenogenetic 
generation were incomplete, and that males might have been 
found a little earlier, and he consequently rejects altogether 
the occurrence of parthenogenesis in Halictus. Nicolas con- 
firms Fabre's observations, so far as the interesting point of the 
work done for common benefit is concerned ; and adds that the 
common corridor being too narrow to permit of two bees passing, 
there is a dilatation or vestibule near the entrance that facilitates 
passage, and also that a sentinel is stationed at this point. 

Smith's observations on Halictus morio in England lead one 
to infer that there is but one generation, the appearance of which 
extends over a very long period. He says, " Early in April the 
females appeared, and continued in numbers iip to the end of 

1 C.R. Ac. Paris, Ixxxix. 1879, p. 1079, and Ann. Set. Nut. ((>), ix. 1879, Xo. 4. 
2 Act. 8uc. Bordeaux, xlviii. 1895, p. 145. 



BEES ANDRENIDES 




June " ; then there was an interval, and in the middle of August 
males began to appear, followed in ten 
or twelve days by females. Hence it is \/ 
probable that in different countries the 
times of appearance and the number of . ; -.\% : --.f^~'V'~r^f 

generations of the same species may vary. /^ 7 | 

Yerhoeff has described the burrows of 
Halictus quadricinctus with some detail. 
The cells, instead of being distributed 
as usual throughout the length of the 
burrow one by one, are accumulated 
into a mass placed in a vault communi- 
cating with the shaft. This shaft is 
continued downwards to a depth of 10 
cm., and forms a retreat for the bees 
when engaged in construction. Several 
advantages are secured by this method, 
especially better ventilation, and pro- 
tection from any water that may enter FIG. 12. Nesting of 
the shaft. The larvae that are present ^T^h ^Tl 
in the brood-chambers at any one 
moment differ much in their ages, a 
fact that throws some doubt 011 the 
supposed parthenogenetic generation. 
No cocoons are formed by these Halictus, 

the polished interior of the cell being a sufficiently refined resting 
place for metamorphosis. Yerhoeff states that many of the 
larvae are destroyed by mouldiness ; this indeed, he considers to 
lie the most deadly of the enemies of Aculeate Hymenoptera. 
The nest of Halictus maculatus has also been briefly described 
by Yerhoeff, and is a very poor construction in comparison with 
that of H. quadricinctus. 

The genus Andrena includes a great number of species, 
Britain possessing about fifty. They may be described in a 
general manner as Insects much resembling the honey-bee 
for which, indeed, they are frequently mistaken but usually a 
little smaller in size. Many of the bees we see in spring, in 
March or April, are of this genus. They live in burrows in the 
ground, preferring sandy places, but frequently selecting a gravel 
path as the locality for their operations ; they nearly always live 



thereto ; n, retreat or con- 
tinuation of the burrow ; w, 
the vaults ; s, the accumula- 
tion of cells. (After Ver- 
hoeff, Verh. Ver. Rheinl. xlviii. 
1891 ; scale not mentioned.) 



26 



HYMENOPTERA 



CHAP. 



in colonies. Great difficulties attend their study on account of 
several points in their economy, such as, that the sexes are 
different, and frequently not found together ; also that there may 
be two generations of a species in one year, these being more or 
less different from one another. Another considerable difficulty 
arises from the fact that these bees are subject to the attacks of 
the parasite Sti/lops, by which their form is more or less altered. 

1 i/ -L ' v 

These Insects feed in the body of the bee in such a way as to 
affect its nutrition without destroying its life ; hence they offer a 
means of making experiments that may throw valuable light on 
obscure physiological questions. Among the effects they produce 
in the condition of the imago bee we may mention the enfeeble- 
ment of the sexual distinction, so that a stylopised male bee 
becomes less different than it usually is from the female, and a 
stylopised female may be ill developed and less different than 
usual from the male. The colours and hair are sometimes altered, 
and distortion of portions of the abdominal region of the bee are 
very common. Further particulars as to these parasites will be 
found at the end of our account of Coleoptera (p. 298). We may 
here remark that these Stylops are not the only parasitic Insects 
that live in the bodies of Andrenidae without killing their hosts, 
or even interrupting their metamorphoses. Mr. E. C. L. Perkins 

recently captured a specimen of Halictus 
rul)icundus,fcom which he, judging from the 
appearance of the example, anticipated that 
a Stylops would emerge ; but instead of this 
a Dipterous Insect of the family Chloropidae 
appeared. Dufour in 1837 called attention 
to a remarkable relation existing between 
Andrena aterrimaand a parasitic Dipterous 
larva. The larva takes up a position in 
the interior of the bee's body so as to be 
partly included in one of the great tracheal 
FIG. 13. Parasitic Dipterous vesicles at the base of the abdomen ; and 
larva in connection with the bee then ma i nta ms the parasite in its 

tracheal system of An- _ r 

Arena aterrima. (After position, and at the same time supplies it 

with air by causing two tracheae to grow 

into its body. Dufour states that he demonstrated the continuity 
of the tracheae of the two organisms, but it is by no means clear 
that the continuity was initially due to the bee's organisation. 




BEES ANDRENIDES 



Dasypoda hirtipes appears to be the most highly endowed of 
the European Andrenides. The 
Insects of the genus Dasypoda 
are very like Andrena, but 
have only two in place of three 
submarginal cells (just beneath 
the stigma) on the front wing. 
The female of D. hirtipes has a 
very dense and elongate pubes- 
cence on the posterior legs, and 




FIG. 14. D. hirtipes 9. Britain. 



carries loads of pollen, each about 
half its own weight, to its nest. 
The habits of this insect have been described by Hermann 
Mtiller. 1 It forms burrows in the ground after the fashion of 
Andrena ; this task is accomplished by excavating with the 
mandibles ; when it has detached a certain quantity of the earth 
it brings this to the surface by moving backwards, and then dis- 
tributes the loose soil over a considerable area. It accomplishes 
this in a most beautiful manner by means of the combined action 
of all the legs, each pair of these limbs performing its share of 
the function in a different manner ; the front legs acting with 
great rapidity making four movements in a second push the 
sand backwards under the body, the bee moving itself at the 
same time in this direction by means of the middle pair of legs ; 
simultaneously, but with a much slower movement, the hind legs 
are stretched and moved outwards, in oar-like fashion, from the 
body, and thus sweep away the earth and distribute it towards 
each side. This being done the bee returns quickly into the hole, 
excavates some more earth, brings it up and distributes it. Each 
operation of excavation takes a minute or two, the distribution 
on the surface only about fifteen seconds. The burrow extends 
to the length of one or two feet, so that a considerable amount of 
earth has to be brought up ; and when the Insect has covered one 
part of the circumference of the mouth of the hole with loose 
earth, it makes another patch, or walk, by the side of the first. 
The main burrow being completed, the Insect then commences 
the formation of brood-chambers in connection with it. Three to 
six such chambers are formed in connection with a burrow ; the 
lower one is first made and is provisioned by the bee : for this 

1 J'crh. Vcr. Eheinland, xli. 1884, p. 1. 



28 HYMENOPTERA CHAP. 

purpose five or six loads of pollen are brought to the cell, each 
load being, as we have already remarked, about half the weight 
of the Insect. This material is then formed into a ball and made 
damp with honey ; then another load of pollen is brought, is 
mixed with honey and added as an outer layer to the ball, which 
is now remodelled and provided on one side with three short feet, 
after which an egg is placed on the top of the mass ; the bin- 
then sets to work to make a second chamber, and uses the 
material resulting from the excavation of this to close completely 
the first chamber. The other chambers are subsequently formed 
in a similar manner, and then the burrow itself is filled up. 
While engaged in ascertaining these facts, Miiller also made 
some observations on the way the bee acts when disturbed 
in its operations, and his observations on this point show 
a very similar instinct to that displayed by Ckalicodoma, 
referred to on a subsequent page. If interrupted while storing a 
chamber the Insect will not attempt to make a fresh one, but 
will carry its stock of provisions to the nest of some other 
individual. The result of this proceeding is a struggle between 
the two bees, from which it is satisfactory to learn that the 
rightful proprietor always comes out victorious. The egg placed 
on the pollen-ball in the chamber hatches in a few days, giving 
birth to a delicate white larva of curved form. This creature 
embraces the pollen-ball so far as its small size will enable it to 
do so, and eats the food layer by layer so as to preserve its 
circular form. The larva when hatched has no anal orifice 
and voids no excrement, so that its food is not polluted ; a 
proper moulting apparently does not take place, for though a 
new delicate skin may be found beneath the old one this latter is 
nut definitely cast off. When the food, which was at first 100 
to 140 times larger than the egg or young larva, is all consumed 
the creature then for the first time voids its refuse. During 
its orowth the larva becomes red and increases in weight from '0025 

O <J 

grains to '26 or '35 grains, but during the subsequent period of 
excretion it diminishes to '09 or '15 grains, and in the course of 
doing so becomes a grub without power of movement, and of a 
white instead of a red colour. After this the larva reposes 
motionless for many months in fact, until the next summer, when 
it throws off the larval skin and appears as a pupa. The larval 
skin thus cast off contrasts greatly with the previous delicate coiuli- 



BEES DENUDATAE 29 



tion of the integument, for this last exuvium is thick and ri<dd. 

o o 

Although it voids no excrement till much later the union of the 
stomach and hind-intestine is accomplished when the larva is 
half-grown. A larva, from which Miiller took away a portion of 
its unconsumed food -store, began directly afterwards to emit 
excrement. The pupa has greater power of movement than 
the resting larva ; when it has completed its metamorphosis 
and become a perfect Insect, it, if it be a female, commences 
almost immediately after its emergence to form burrows by the 
complex and perfect series of actions we have described. 

Parasitic Bees (DENUDATAE). This group of parasitic bees 
includes fourteen European genera, of which six are British. 
They form a group taxonomically most unsatisfactory, the 
members having little in common except the negative characters 
of the absence of pollen-carrying apparatus. Although there 
is a great dearth of information as to the life -histories of 
parasitic bees, yet some highly interesting facts and generalisa- 
tions about their relations with their hosts have already been 

t/ 

obtained. Verhoeff has recently given the following account of 
the relations between the parasitic bee Stelis minuta and its 
host Osmia leucomelana : The Osmia forms cells in blackberry 
stems, provisions them in the usual manner, and deposits an 
egg in each. But the Stelis lays an egg in the store of pro- 
visions before the Osmia does, and thus its egg is placed lower 
down in the mass of food than that of the legitimate owner, 
which is in fact at the top. The Stelis larva emerges from the 
egg somewhat earlier than the Osmia larva does. For a con- 
siderable time the two larvae so disclosed consume together the 
stock of provisions, the Osmia at the upper, the Stelis at the lower, 
end thereof. By the consumption of the provisions the two larvae 
are brought into proximity, and by this time the Stelis larva, being 
about twice the size of the Osmia larva, kills and eats it. Verhoeff 
witnessed the struggle between the two larvae, and states further 
that the operation of eating ' the Osmia larva after it has been 
killed lasts one or two days. He adds that parasitic larvae are 
less numerous than the host larvae, it being well known that 
parasitic bees produce fewer offspring than host bees. Verhoeff 
further states that he has observed similar relations to obtain 
between the larvae of other parasitic bees'and their hosts, but warns 
us against concluding that the facts are analogous in all cases. 



10 



HYMENOPTERA 



CHAP. 




FIG. 15. Nbmada sex-fasriata 9- 

Britain. 



Fabre has made as acquainted with some points in the history of 
another species of the same genus, viz. Stelis nasuta, that show a 
decided departure from the habits of S. minuta. The first-named 
Insect accomplishes the very difficult task of breaking open the cells 
of the mason-bee, Chalicodoma muraria, after they have been sealed 
up, and then, being an Insect of much smaller size than the Chali- 
codoma, places several eggs in one cell of that bee. Friese informs 
us that parasitic bees and their hosts, in a great number of cases, 
not only have in the perfect state the tongue similarly formed, 

but also frequent the same species 
of flower ; thus Colletcs davicsanus 
and its parasite Epeolus variegatus 
both specially affect the flowers of 
Tanacetum vulyare. Some of the 
parasitic bees have a great resem- 
blance to their hosts ; Stelis signata, 
for instance, is said to be so like 
Antliidiiim strigatum that for many 
years it was considered to be a 
species of the genus Anthidium. 
In other cases not the least resemblance exists between the 
parasites and hosts. Thus the species of Nomada that live at 
the expense of species of the genus Andre no, have 110 resemblance 
thereto. Friese further tells us that the Andrena and Nomada 
are on the most friendly terms. Andrena, as is well known, 
forms populous colonies in banks, paths, etc., and in these colonies 
the destroying Nomada flies about unmolested ; indeed, according 
to Friese, it is treated as a welcome guest. He says he has often 
seen, and in several localities, Nomada lathburiana and Andrena 
ovina flying peacefully together. The Nome da would enter a 
burrow, and if it found the Andrena therein, would come out and 
try another burrow ; if when a marauding Nomada was in a 
burrow, and the rightful owner, returning laden with pollen, 
found on entering its home that an uninvited guest was therein, 
the Andrena would go out in order to permit the exit of the 
Nomada, and then would again enter and add the pollen to the 
store. Strange as this may seem at first sight, it is really not 
so, for, as we have before had occasion to observe, there is not the 
slightest reason for believing that host Insects have any idea 
whatever that the parasites or iiiquilines are injurious to their 



i PARASITIC BEES DENUDATAE 31 

race. Why then should they attack the creatures ? Provided the 
parasites do not interfere in any unmannerly way with the hosts 
and their work, there is no reason why the latter should resent 
their presence. The wild bee that seals up its cell when it has 
laid an egg therein, and then leaves it for ever, has no conception 
of the form of its progeny ; never in the history of the race of the 
Andrena has a larva seen a perfect insect and survived thereafter, 
never has a perfect Insect seen a larva. There is no reason what- 
ever for believing that these Insects have the least conception of 
their own metamorphosis, and how then should they have any 
idea of the metamorphosis of the parasite ? If the Andrena found 
in the pollen the egg of a parasitic Nomada, it could of course 
easily remove the egg ; but the Andrena has no conception 
that the presence of the egg ensures the death of its own 
offspring and though the egg be that of an enemy to its race, 
why should it resent the fact ? Is it not clear that the race has 
always maintained itself notwithstanding the enemy ? Nature has 
brought about that both host and parasite should successfully 
co-exist ; and each individual of each species lives, not for itself, 
but for the continuance of the species ; that continuance is pro- 
vided for by the relative fecundities of host and guest. Why 
then should the Andrena feel 
alarm ? If the species of Nomada 
attack the species of Andrena too 
much it brings about the de- 
struction of its own species 
more certainly than that of the 
Andrena. 

Such extremely friendly rela- 
tions do not, however, exist be- 
tween all the parasitic bees and 

FIG. 16. Melecta luctuosa ?. Britain. 

their hosts. Friese says that, so 

far as he has been able to observe, the relations between the two 
are not in general friendly. He states that marauders of the 
genera Melecta and Coelioxys seek to get out of the way when 
they see the pollen-laden host coming home. But he does not 
appear to have noted any other evidence of mistrust between the 
two, and it is somewhat doubtful whether this act can properly 
be interpreted as indicating fear, for bees, as well as other 
animals, when engaged in work find it annoying to be interfered 




32 HYMENOPTERA CHAP. 

with ; it is the interest of the parasite to avoid annoyance and to 
be well-mannered in its approaches. Shuckard, however, says that 
battles ensue between the parasite Melecta and its host Anthopliora, 
when the two bees meet in the burrows of the Anthophora. 1 

We shall have occasion to remark on some of the habits of 
Dioxys cincta when considering the history of the mason -bee 
(Chalicodoma), but one very curious point in its economy must 
here be noticed. The Dioxys, which is a much smaller bee than 
the Chalicodoma, lays an egg in a cell of the latter, and the 
resulting larva frequently has more food in the cell than it can 
consume ; there is, however, another bee, Osmia cyanoxantlia, that 
frequently takes advantage of an unoccupied cell in the nest 
of the Chalicodoma, and establishes its own offspring therein. 
The Dioxys, it seems, cannot, or at any rate does not, distinguish 
whether a cell is occupied by Chalicodoma or by Osmia, and some- 
times lays its egg in the nest of the Osmia, though this bee is 
small, and therefore provides very little food for its young. It 
might be supposed that under these conditions the Dioxys larva 
would be starved to death ; but this is not so ; it has the power 
of accommodating its appetite, or its capacity for metamorphosis, 
to the quantity of food it finds at its disposal, and the egg laid in 
the Osmia cell actually produces a tiny specimen of Dioxys, only 
about half the natural size. Both sexes of these dwarf Dioxys are 
produced, offering another example of the fact that the quantity 
of food ingested during the lifetime of the larva does not influ- 
ence the sex of the resulting imago. 

The highly endowed bees that remain to be considered are 
by some writers united in a group called Apidae, in distinction 
from Andrenidae. For the purposes of this work we shall adopt 
three divisions, Scopulipedes, Dasygastres, Sociales. 

The group SCOPULIPEDES includes such long-tongued, solitary 
bees as are not parasitic, and do not belong to the Dasygastres. 
It is not, however, a natural group, for the carpenter-bees 
(Xylocopa) are very different from Anthopliora. It has 
recently been merged by Friese with Andrenides into a single 
group called Podilegidae. Four British genera, Ceratina, AnfJio- 
pliora, Eucera and Saropoda (including, however, only seven 

1 It is impossible for us here to deal with the question of the origin of tin- para- 
sitic habit in bees. The reader wishing for information as to this may refer to 
Prof. Perez's paper, Act. Sue. Bordeaux, xlvii. 1895. p. 300. 



i SCOPULIPEDES CARPENTER-BEES 33 

species), are referred to the Scopulipedes ; in some forms a con- 
siderable resemblance to the Bonibi is exhibited, indeed the 
female of one of our species of Anthopkora is so very like the 
worker of Uomlns hortorum var. harrisellus, that it would puzzle 
any one to distinguish them by a superficial inspection, the 
colour of the hair on the hind legs being the only obvious differ- 
ence. Anthophora is one of the most extensive and widely 
distributed of the genera of bees. Some of the species make 
burrows in cliffs and form large colonies which are continued for 
many years in the same locality. Friese has published many 
details of the industry and metamorphoses of some of the species 
of this genus ; the most remarkable point he has discovered being 
that A. per sonata at Strasburg takes two years to accomplish the 
life-cycle of one generation. Some of the European species of the 
genus have been found to be very subject to the attacks of para- 
sites. An anomalous beetle, Sitaris, has been found in the nests 
of A. pilipes ; and this same Anthophora is also parasitised by 
another beetle, Meloe, as well as by a bee of the genus Melecta. 

The genus Xylocopa l contains many of the largest and most 
powerful of the bees, and is very widely distributed over the 
earth. In Europe only four or five species have been found, and 
none of them extend far northwards, X. riolacea being the only 
one that comes so far as Paris. They are usually black or blue- 
black in colour, of broad, robust build, with shining integuments 
more or less covered with hair. X. molacea is known as the 
carpenter-bee from its habit of working in dry wood ; it does not 
touch living timber, but will form its nest in all sorts of dried 
wood. It makes a cylindrical hole, and this gives access to three 
or four parallel galleries in which the broad cells are placed ; the 
cells are always isolated by a partition ; the bee forms this by 
cementing together with the products of its salivary glands the 
fragments of wood it cuts out. Its habits have been described 
at length by Reaumur, who alludes to it under the name of 
" abeille perce-bois." This bee hibernates in the imago condition, 
both sexes reappearing in the spring. Possibly there is more than 
one generation in the year, as Reaumur states that specimens 
that were tiny larvae on the 12th of June had by the 2nd of July 
consumed all their stock of provisions ; they then fasted for a few 
days, and on the 7th or 8th of July became pupae, and in the first 

1 Refer to p. 70 iwstea, note, as to a recent discovery about Xylocopa. 
VOL. VI D 



34 



HYMENOPTERA 



CHAP. 



days of August were ready to emerge as perfect Insects. Thus 
the whole cycle of metamorphoses is passed through in about 
eight weeks. This species, though very clever in drilling holes, 
does not hesitate to appropriate old burrows should they be at 
hand. Fabre observed that it was also quite willing to save 
itself labour by forming its cells in hollow reeds of sufficient 
calibre. AVe have figured the larva, and pupa of this species in 
the previous volume (p. 170). 

Xylocopa chloroptera in E. India selects a hollow bamboo for 
its nidus ; it cements together the pieces obtained in clearing 




FIG. 17. Xylocopa (Koptorthosoma), $2). uear.//<nv)///V//r.*ceHs, <J. Sarawak. 

out the bamboo, and uses them as horizontal partitions to separate 
the tube into cells. The species is much infested with a small 
Chalcid of the genus /u-//rtus : 300 specimens of the parasite 
have been reared from a single larva of the bee ; two-thirds of 
the larvae of this bee that Home endeavoured to rear were 
destroyed by the little Chalcid. 

The most beautiful and remarkable of all the bees are 
the species of Euglossa. This genus is peculiar to Tropical 
America, and derives its name from the great length of the 
proboscis, which in some species surpasses that of the body. The 
colours in Et/ylossa proper are violet, purple, golden, and metallic 
green, and two of these are frequently combined in the most har- 
monious manner: the hind tibia is greatly developed and forms 
a plate, the outer surface of which is highly polished, while the 
margins are furnished with rigid hairs. Very little is known as 
to the habits of these bees; they were formerly supposed to be 



DASYGASTRES MASON-BEES 



35 



a 



f 




social ; but this is doubtful, Bates having recorded that E. sur- 
inamensis forms a " solitary nest." Lucas concluded that E. 
cordata is social, 011 the authority of a nest containing 
dozen individuals." No workers 
are known. The species of 
Eulema have a shorter tongue 
than Euglossa, and in form 
and colour a good deal re- 
semble our species of Bonibus 
and Apathus. 

The group DASYGASTRES in- 
cludes seven European genera, 
four being British (Ghelostoma 
being included in Heriacles). 
The ventral surface of the hind 
body is densely set in the 
females with regularly arranged 
hairs, by means of which the 
pollen is carried. In many of 
the Dasygastres (Megacliile, 
e.g.} the labrum is very large, 
and in repose is inflected on 
to the lower side of the head, 
and closely applied to the 
doubled -in tongue, which it - 

FIG. 18. Euglossa cordata, &. Amazons. 
Serves to protect ; the man- A, The Insect with extended proboscis ; 

dibles then lock together out- 
side the labrum, which is thus completely concealed. This group 
includes some of the most interesting of the solitary bees. 

The genus Clicdicodoma is not found in our own country, but 
in the South of France there exist three or four species. Their 
habits have given rise to much discussion, having been described 
by various naturalists, among whom are included Reaumur and 
Fabre. These Insects are called mason-bees, and construct nests of 
very solid masonry. C. muraria is in appearance somewhat inter- 
mediate between a honey-bee and a JSomb-us ; it is densely hairy, 
and the sexes are very different in colour. It is solitary in its 
habits, and usually chooses a large stone as a solid basis for its 
habitation. On this a cell is formed, the material used being 
a kind of cement made by the Insect from the mixture of a 




HYMENOPTERA 



CHAP. 



suitable sort of earth with the material secreted by its own 
salivary glands ; the amount of cement used is reduced 1 >y the 
artifice of building small stones into the walls of the cell ; the 
stones are selected with great care. When a cell about an inch 

in depth has been formed in 
this manner, the bee commences 
to fill it with food, consisting 
of honey and pollen ; a little 
honey is brought and is dis- 
charged into the cell, then some 
pollen is added. This bee, like 
other Dasygastres, carries the 
pollen by means of hairs on 
the under surface of the body ; 
to place this pollen in the cell 
the Insect therefore enters back- 
wards, and then with the pair 
of hind legs brushes and scrapes 
the under surface of the body 
so as to make the pollen fall off 
into the cell ; it then starts for 
Greece, a fresh cargo ; after a few loads 
have been placed in the recep- 
tacle, the Insect mixes the honey and pollen into a paste 
with the mandibles, and again continues its foraging until it 
has about half filled the cell ; then an egg is laid, and the 
apartment is at once closed with cement. This work is all 
accomplished, if the weather be favourable, in about two clays, 
after which the Insect commences the formation of a second 
cell, joined to the first, and so on till eight or nine of these 
receptacles have been constructed ; then comes the final operation 
of adding an additional protection in the shape of a thick layer of 
mortar placed over the whole ; the construction, when thus com- 
pleted, forms a sort of dome of cement about the size of half an 
orange. In this receptacle the larvae pass many months, exposed 
to the extreme heat of summer as well as to the cold of winter. 
The larvae, however, are exposed to numerous other perils ; and we 
have already related (vol. v, p. 540) how Lfitcospis gigas succeeds 
in perforating the masonry and depositing therein an egg, so that 
a Leucospis is reared in the cell instead of a Chalicodoma. 




FlG. 19. ChalicoiliniKi ninri>riii. 
A, Male ; B, female. 



i DASYGASTRES MASON-BEES 37 

This Insect has been the object of some of J. H. Fabre's 
most instructive studies on instinct. 1 Although it is impossible 
for us here to consider in a thorough manner the various points 
he has discussed, yet some of them are of such interest and im- 
portance as to demand something more than a passing allusion. 

We have mentioned that the nest of Clialicodoma is roofed 
with a layer of solid cement in addition to the first covering 
with which the bee seals up each cell. When the metamorphoses 
of the imprisoned larva have been passed through, and the moment 
for its emergence as a perfect Insect has arrived, the prisoner has 
to make its way through the solid wall by which it is encom- 
passed. Usually it finds no difficulty in accomplishing the task 
of breaking through the roof, so that the powers of its mandibles 
must be very great. Reaumur has, however, recorded that a nest 
of this mason-bee was placed under a glass funnel, the orifice of 
which was covered with gauze, and that the Insects when they 
emerged from the nest were unable to make their way through 
the gauze, and consequently perished under the glass cover ; and 
he concluded that such insects are only able to accomplish the 
tasks that naturally fall to their lot. By some fresh experiments 
Fabre, however, has put the facts in a different light. He 
remarks that when the Insects have, in the ordinary course of 
emergence, perforated the walls of their dark prison, they find 
themselves in the daylight, and at liberty to walk a\vay ; when 
they have made their escape from a nest placed under a glass 
cover, they, having no knowledge of glass, find themselves in 
daylight and imprisoned by the glass, which, to their inexperience, 
does not appear to be an obstacle, and they therefore, he thought, 
might perhaps exhaust themselves in vain efforts to pass through 
this invisible obstacle. He therefore took some cocoons contain- 
ing pupae from a nest, placed each one of them in a tube of reed, 
and stopped the ends of the reeds with various substances, in 
one case earth, in another pith, in a third brown paper ; the 
reeds were then so arranged that the Insects in them were in a 
natural position ; in due course all the Insects emerged, none of 
them apparently having found the novel nature of the obstacle a 
serious impediment. Some complete nests were then taken with 
their inmates, and to the exterior of one of them a sheet of 
opaque paper was closely fastened, while to another the same 

1 Souvenirs entomoloyiqucs. 4 vols. Paris, 1879 to 1891. 



38 HYMENOPTERA CHAP. 

sort of paper was applied in the form of a dome, leaving thus a 
considerable space between the true cover of the nest and the 
covering of paper. From the first nest the Insects made their 
escape in the usual manner, thus again proving that paper can 
be easily pierced by them. From the second nest they also 
liberated themselves, but failed to make their way out through 
the dome of paper, and perished beneath it ; thus showing that 
paper added to the natural wall caused them no difficulty, but 
that paper separated therefrom by a space was an insuperable 
obstacle. Professor Perez has pointed out that this is no doubt 
due to the large space offered to the bee, which consequently 
moves about, and does not concentrate its efforts on a single 
spot, as it of course is compelled to du when confined in its 
natural cell. 

The power of the mason -bee to find its nest again when 
removed to a distance from it is another point that was tested 
by Du Hamel and recounted by Reaumur. As regards this 
Fabre has also made some very valuable observations. He marked 
some specimens of the bee, and under cover removed them to 
a distance of four kilometres, and then liberated them ; the 
result proved that the bees easily found their way back again, 
and indeed were so little discomposed by the removal that they 
reached their nests laden with pollen as if they had merely been 
out on an ordinary journey. On one of these occasions he 
observed that a Chalicodoma, on returning, found that another 
bee had during her absence taken possession of her partially 
completed cell, and was unwilling to relinquish it ; whereupon 
a battle between the two took place. The account of this is 
specially interesting, because it would appear that the two com- 
batants did not seek to injure one another, but were merely 
eno-ag-ed in testing, as it were, which was the more serious in its 

C> O O> 

claims to the proprietorship of the cell in dispute. The matter 
ended by the original constructor regaining and retaining posses- 
sion. Fabre says that in the case of Chalicodoma it is quite a 
common thing for an uncompleted cell to be thus appropriated 
by a stranger during the absence of the rightful owner, and that 
after a scene of the kind described above, the latter of the two 
claimants ;il\\;iys regains possession, thus leading one to suppose 
that some sense of rightful ownership exists in these bees; the 
usurper expressing, as it were, by its actions the idea Before I 



i DASYGASTRES MASON-BEES FINDING THE NEST 39 

resign my claims I must require you to go through the exertions 
that will prove you to be really the lawful owner. 

Another experiment was made with forty specimens of Chali- 
codoma pyrenaica, which were removed to a distance of four 
kilometres and then liberated. About twenty of the individuals 
had been somewhat injured by the processes of capturing, mark- 
ing, and transferring, and proved unable to make a proper start. 
The others went off well when released, and in forty minutes the 
arrivals at the nest had already commenced. The next morning 
he was able to ascertain that fifteen at least had found their way 
back, and that it was probable that most of the uninjured bees 
had reached home ; and this although, as Fabre believed, they 
had never before seen the spot where he liberated them. 

These observations on the power of Chalicodoma to regain 
its nest attracted the attention of Charles Darwin, who wrote to 
M. Fabre, and suggested that further observations should be 
made with the view of ascertaining by means of what sense these 
bees were able to accomplish their return. For it must be borne 
in mind that this bee is very different from the domestic bee, 
inasmuch as it enjoys but a brief life in the winged state, and it 
is therefore to be presumed that an individual has no knowledge 
of such comparatively distant localities as those to which Fabre 
transported it. Further observations made by the Frenchman 
have unfortunately failed to throw any light on this point. 
Darwin thought it might possibly be some sensitiveness to 
magnetic conditions that enabled the bees to return home, 
and suggested that they should be tested as to this. Fabre 
accordingly made some minute magnets, and fixed one to each 
bee previous to letting them loose for a return journey. This 
had the effect of completely deranging the bees ; and it was there- 
fore at first thought that the requisite clue was obtained. It 
occurred to the experimenter, however, to try the plan of affixing 
small pieces of straw to the bees instead of magnets, and on this 
being done it was found that the little creatures were just as 
much deranged by the straws as they were by the magnets : thus 
it became evident that no good grounds exist for considering 
that the bees are guided by magnetic influences. 

One of the species * of Chalicodoma observed by Fabre fixes 

1 The " ChaUcodome des galcts" or C. " dcs murailles" of the French writer ; in 
some places he speaks of the species as being C. mumrin, in others as C. parietina. 



40 HYMENOPTERA CHAP. 

its nests to the small boulders brought down and left by the 
Khone on the waste places of its banks. This habit afforded 
Fabre an opportunity of removing the nests during the process 
of construction, and of observing the effect this produced on the 
architects. While a bee that had a nest partially constructed 
was absent, he removed the stone and the nest attached to it from 
one situation to another near at hand and visible from the 
original site. In a few minutes the bee returned and went 
straight to the spot where the nest had been ; finding its home 
absent it hovered for a little while around the place, and then 
alighted on the vacated position, and walked about thereon in 
search of the nest ; being after some time convinced that this 
was no longer there, it took wing, but speedily returned again to 
the place and went through the same operations. This series of 
manoeuvres was several times repeated, the return always being 
made to the exact spot where the nest had been originally located ; 
and although the bee in the course of its journeys would pass 
over the nest at a distance of perhaps only a few inches, it did 
not recognise the object it was in search of. If the nest 
were placed very near to the spot it had been removed from- 
say at a distance of about a yard it might happen that the bee 
would actually come to the stone to which the nest was fixed, 
would visit the nest, would even enter into the cell it had left 
partially completed, would examine circumspectly the boulder, 
but would always leave it, and again return to the spot where 
the nest was originally situated, and, on finding that the nest 
was not there, would take its departure altogether from the 
locality. The home must be, for the bee, in the proper situation, 
or it is not recognised as the desired object. Thus we are con- 
fronted with the strange fact that the very bee that is able to 
return to its nest from a distance of four kilometres can no 
longer recognise it when removed only a yard from the original 
position. This extraordinary condition of the memory of the 
Insect is almost inconceivable by us. That the bee should 
accurately recognise the spot, but that it should not recognise 
the cell it had itself just formed and half-filled with honey-paste, 
seems to us almost incredible ; nevertheless, the fact is quite con- 
sistent with what we shall subsequently relate in the case of the 
solitary wasp BeTiibex. A cross experiment was made by taking 
away the stone with the attached nest of the bee while the latter 



i DASYGASTRES MASON-BEES INSTINCT 41 

was absent, and putting in its place the nest of another indi- 
vidual in about the same stage of construction ; this nest was 
at once adopted by the bee, which indeed was apparently in no 
way deranged by the fact that the edifice was the work of another. 
A further experiment was made by transposing the positions of 
two nests that were very near together, so that each bee when 
returning might be supposed to have a free choice as to which 
nest it would go to. Unhesitatingly each bee selected the nest 
that, though not its own, was in the position where its own had 
been. This series of experiments seems to prove that the Chalci- 
doma has very little sense as to what is its own property, but, on 
the other hand, has a most keen appreciation of locality. As, 
however, it might be supposed that the bees were deceived by the 
similarity between the substituted nests, Fabre transposed two 
nests that were extremely different, one consisting of many cells, 
the other of a single incomplete cell ; it was, of course, a necessary 
condition of this experiment that each of the two nests, however 
different in other respects, should possess one cell each in similar 
stages of construction ; and when that was the case each bee 
cheerfully adopted the nest that, though very different to its 
own, was in the right place. This transposition of nests can be 
rapidly repeated, and thus the same bee may be made to go on 
working at two different nests. 

Suppose, however, that another sort of change be made. Let 
a nest, consisting of a cell that is in an early stage of construc- 
tion, be taken away, and let there be substituted for it a cell 
built and partially stored with food. It might be supposed that 
the bee would gladly welcome this change, for the adoption of 
the substituted cell would save it a great deal of work. Not so, 
however ; the bee in such a case will take to the substituted cell, 
but will go on building at it although it is already of the full 
height, and will continue building at it until the cell is made as 
much as a third more than the regulation height. In fact the 
bee, being in the building stage of its operations, goes on build- 
ing, although in so doing it is carrying on a useless, if not an 
injurious, work. A similar state ensues when the Insect ceases 
to build and begins to bring provisions to the nest ; although a 
substituted cell may contain a sufficient store of food, the bee goes 
on adding to this, though it is wasting its labours in so doing. 
It should be noted that though the bee must go through the 



42 HYMENOPTERA CHAP. 

appropriate stages of its labours whether the result of so doing 
be beneficial or injurious, yet it is nevertheless to some extent 
controlled by the circumstances, for it does not in such cases 
complete what should have been the full measure of its own 
individual work ; it does not, for instance, raise the cell to twice 
the natural height, but stops building when the cell is about 
one-third larger than usual, as if at that stage the absurdity of 
the situation became manifest to it. 

Fabre's experiments with the Chalicodoma are so extremely 
instructive as regards the nature of instinct in some of the 
highest Insects, that we must briefly allude to some other of his 
observations even at the risk of wearying the reader who feels 
but little interest in the subject of Insect intelligence. 

Having discovered that a mason-bee that was engaged in the 
process of construction would go on building to an useless or 
even injurious extent, Fabre tried another experiment to ascer- 
tain whether a bee that was engaged in the process of provision- 
ing the nest, would do so in conditions that rendered its work 
futile. Taking away a nest with completely built cell that a bee 
was storing with food, he substituted for it one in which the cell 
was only commenced, and therefore incapable of containing food ; 
when the bee with its store of provisions reached this should-be 
receptacle it appeared to be very perplexed, tested the im- 
perfect cell with its antennae, left the spot and returned again ; 
repeating this several times it finally went to the cell of 
some stranger to deposit its treasure. In other cases the bee 
broke open a completed cell, and having done so went on bringing 
provisions to it ; although it was already fully provisioned and an 
egg laid therein : finally, the little creature having completed 
the bringing of this superfluous tale of provisions, deposited 
a second egg, and again sealed up the cell. But in no case 
does the bee go back from the provisioning stage to the build- 
ing stage until the cycle for one cell of building, provisioning, 
and egg-laying is completed : but when this is the case, the 
building of a fresh cell may be again undertaken. This is a 
good example of the kind of consecutive necessity that seems 
to be one of the chief features of the instinct of these industrious 
little animals. Another equally striking illustration of these 
peculiarities of instinct is offered by interfering with the act of 
putting the provisions into the cell. It will be recollected that 



i DASYGASTRES- MASON-BEES PARASITES 43 

when the bee brings provisions to add to the stock, it carries 
both honey and pollen ; in order to deliver these it begins by 
entering head first into the cell and disgorging the honey, then 
emerging it turns round, enters backwards and scrapes off the 
pollen from its body. If after the honey has been discharged, 
the bee be interfered with and gently removed to a slight dis- 
tance with a straw, it returns to complete its task, but instead of 
going on with the actions at the point at which the interruption 
took place, it begins the series over again, going in at any rate 
partially head first, although it has no honey to discharge, and 
having performed this useless ceremony it then emerges, turns 
round and adds the pollen. This illustration is in some respects 
the reverse of what might have been expected, for the Insect 
here does not continue the act at the interrupted point, but begins 
the series of actions afresh. 

It would be reasonable to suppose that an Insect that takes 
the pains to provide for the safety of its progeny by constructing 
a complex edifice of cement, secures thereby the advantage of 
protection for its young. But this is far from being the case. 
Notwithstanding the cement and the thick dome of mortar, the 
Chalicodoma is extremely subject to the attacks of parasites. 
The work performed by the creature in constructing its mass of 
masonry is truly astounding ; Fabre calculated from measure- 
ments he made that for the construction and provisioning of a 
single cell, the goings and comings of the bee amounted to 15 
kilometres, and it makes for each nest sometimes as many as 
fifteen cells. Notwithstanding all this labour, it would appear 
that no real safety for the larvae is obtained by the work. Some 
sixteen possibly more other species of Insects get their living 
off this industrious creature. Another bee, Stelis nasuta, breaks 
open the cells after they have been completely closed and places 
its o\vn eggs in them, and then again closes the cells with 
mortar. The larvae of this Stelis develop more rapidly than do 
those of the Chalicodoma, so that the result of this shameless 
proceeding is that the young one of the legitimate proprietor 
as we human beings think it is starved to death, or is possibly 
eaten up as a dessert by the Stelis larvae, after they have 
appropriated all the pudding. 

Another bee, Dioxys cincta, is even more audacious ; it flies 
about in a careless manner among the Chalicodoma at their 



44 HYMENOPTERA CHAP. 

work, and they do not seem to object to its presence unless it 
interferes with them in too unmannerly a fashion, when they 
brush it aside. The Dioxys, when the proprietor leaves the cell, 
will enter it and taste the contents ; after having taken a few 
mouthfuls the impudent creature then deposits an egg in the 
cell, and, it is pretty certain, places it at or near the bottom of 
the mass of pollen, so that it is not conspicuously evident to the 
Chalicodoma when the bee again returns to add to or complete 
the stock of provisions. Afterwards the constructor deposits its 
own egg in the cell and closes it. The final result is much the 
same as in the case of the Stelis, that is to say, the Chalicodoma 
has provided food for an usurper ; but it appears probable that 
the consummation is reached in a somewhat different manner, 
namely, by the Dioxi/s larva eating the egg of the Chali- 
codoma, instead of slaughtering the larva. Two of the Hymenop- 
tera Parasitica are very destructive to the Chalicodoma, viz. 
Leiicospis gigas and Mo nodontomcrus nitidus ; the habits of which 
-ve have already discussed (vol. v. p. 54:!) under Chalcididae. 
Lainpert has given a list of the Insects attacking the mason-bee 
or found in its nests ; altogether it would appear that about 
sixteen species have been recognised, most of which destroy the 
bee larva, though some possibly destroy tlje bee's destroyers, and 
two or three perhaps merely devour dead examples of the bee, or 
take the food from cells, the inhabitants of which have been 
destroyed by some untoward event. This author thinks that 
one half of the bees' progeny are made away with by these 
destroyers, while Fabre places the destruction in the South of 
France at a still higher ratio, telling us that in one nest of nine 
cells, the inhabitants of three were destroyed by the Dipterous 
Insect, Anthrax trifasciata, of two by Leucospis, of two by Stelis, and 
of one by the smaller Chalcid ; there being thus only a single 
example of the bee that had not succumbed to one or other of 
the enemies. He has sometimes examined a large number of 
nests without finding a single one that had not been attacked by 
one or other of the parasites, and more often than not several of 
the marauders had attacked the nest. 

It is said by Lampert and others that there is a passage in 
Pliny relating to one of the mason-bees, that the Roman author 
had noticed in the act of carrying off stones to build into its 
nest ; being unacquainted with the special habits of the bee, he 



DASYGASTRES CARDER-BEE 



45 



seems to have supposed that the insect was carrying the stone 
as ballast to keep itself from being blown away. 

The bees of the genus Anthidium are known to possess the 
habit of making nests of wool or cotton, that they obtain from 
plants growing at hand. We 
have one species of this genus 
of bees in Britain ; it some- 
times may be seen at work in 
the grounds of our Museum 
at Cambridge : it is referred 
to by Gilbert White, who 
says of it, in his History 
of Selborne : " There is a sort 
of wild bee frequenting the 
garden-campion for the sake 
of its tomentum, which prob- 
ably it turns to some purpose 
in the business of nidincation. 
It is very pleasant to see with 
what address it strips off the 
pubes, running from the top 
to the bottom of a branch, 
and shaving it bare with the 
dexterity of a hoop -shaver. 
When it has got a bundle, 
almost as large as itself, it flies 
away, holding it secure between its chin and its fore legs." 
The species of this genus are remarkable as forming a con- 
spicuous exception to the rule that in bees the female is 
larger than the male. The species of Anthidium do not form 
burrows for themselves, but either take advantage of suitable 
cavities formed by other Insects in wood, or take possession of 
deserted nests of other bees or even empty snail-shells. The 
workers in cotton, of which our British species A. manic atum is 
one, line the selected receptacle with a beautiful network of 
cotton or wool, and inside this place a finer layer of the material, 
to which is added some sort of cement that prevents the honied 
mass stored by the bees in this receptacle from passing out of it. 
A. diadema, one of the species that form nests in hollow steins, 
has been specially observed by Fabre ; it will take the cotton for 




FIG. 20. Anthidium manicatum, Carder- 
bee. A, Male ; B, female. 



46 IIYMENOPTERA CHAP. 

its work from any suitable plant growing near its nest, and does not 
confine itself to any particular natural order of plants, or even to 
those that are indigenous to the South of France. When it has 
brought a ball of cotton to the nest, the bee spreads out and 
arranges the material with its front legs and mandibles, and 
presses it down with its forehead on to the cotton previously 
deposited ; in this way a tube of cotton is constructed inside the 
reed ; when withdrawn, the tube proved to be composed of about 
ten distinct cells arranged in linear fashion, and connected firmly 
together by means of the outer layer of cotton ; the transverse 
divisions between the chambers are also formed of cotton, and 
each chamber is stored with a mixture of honey and pollen. 
The series of chambers does not extend quite to the end of the 
reed, and in the unoccupied space the Insect accumulates small 
stones, little pieces of earth, fragments of wood or other similar 
small objects, so as to form a sort of barricade in the vestibule, 
and then closes the tube by a barrier of coarser cotton taken 
frequently from some other plant, the mullein by preference. 
This barricade would appear to be an ingenious attempt to keep 
out parasites, but if so, it is a failure, at any rate as against 
Leticospis, which insinuates its eggs through the sides, and 
frequently destroys to the last one the inhabitants of the 
fortress. Fabre states that these Anthidium., as well as J\fcf/cJ/ i/c, 
will continue to construct cells when they have no eggs to place 
in them ; in such a case it would appear from his remarks that 
the cells are made in due form and the extremity of the reed 
closed, but no provisions are stored in the chambers. 

The larva of the Anthidium forms a most singular cocoon. 
We have already noticed the difficulty that arises, in the case 
of these Hymenopterous larvae shut up in small chambers, as 
to the disposal of the matters resulting from the incomplete 
assimilation of the aliment ingested. To allow the once-used 
food to mingle with that still remaining unconsumed would be 
not only disagreeable but possibly fatal to the life of the larva. 
Hence some species retain the whole of the excrement until 
the food is entirely consumed, it being, according to Adlerz, stored 
in a special pouch at the end of the stomach ; other Hymen- 
optera, amongst which we may mention the species of Osmici, 
place the excreta in a vacant space. The Anthidium adopts, 
however, a most remarkable system: about the middle of its 



i DASYGASTRES ANTHIDIUM 47 

larval life it commences the expulsion of " frass " in the shape of 
small pellets, which it fastens together with silk, as the} 7 are 
voided, and suspends round the walls of the chamber. This 
curious arrangement not only results in keeping the embarrassing 
material from contact with the food and with the larva itself, 
but serves, when the growth of the latter is accomplished, as the 
outline or foundations of the cocoon in which the metamorphosis 
is completed. This cocoon is of a very elaborate character ; it 
has, so says Fabre, a beautiful appearance, and is provided with 
a very peculiar structure in the form of a small conical pro- 
tuberance at one extremity pierced by a canal. This canal is 
formed with great care by the larva, which from time to time 
places its head in the orifice in process of construction, and 
stretches the calibre by opening the mandibles. The object of 
this peculiarity in the fabrication of the elaborate cocoon is not 
clear, but Fabre inclines to the opinion that it is for respiratory 
purposes. 

Other species of this genus use resin in place of cotton as their 
working material. Among these are Anthidium septemdentatum and 
A. bcllicosum. The former species chooses an old snail-shell as its 
nidus, and constructs in it near the top a barrier of resin, so as to 
shut off the part where the whorl is too small ; then beneath the 
shelter of this barrier it accumulates a store of honey-pollen, de- 
posits an egg, and completes the cell by another transverse barrier of 
resin ; two such cells are usually constructed in one snail-shell, and 
below them is placed a barricade of small miscellaneous articles, 
similar to what we have described in speaking of the cotton- 
working species of the genus. This bee completes its metamor- 
phosis, and is ready to leave the cell in early spring. Its con- 
gener, A. liellicosum, has the same habits, with the exception 
that it works later in the year, and is thus exposed to a great 
danger, that very frequently proves fatal to it. This bee does 
not completely occupy the snail- shell with its cells, but leaves 
the lower and larger portion of the shell vacant. Now, there, is 
another bee, a species of Osmia, that is also fond of snail-shells 
as a nesting-place, and that affects the same localities as the 
A. septemdentatum; very often the Osmia selects for its nest 
the vacant part of a shell, the other part of which is occupied by 
the Anthidium ; the result of this is that when the metamorphoses 
are completed, the latter bee is unable to effect its escape, and 



4 8 



HYMENOPTERA 



CHAP. 



thus perishes in the cell. Fabre further states with regard to 
these interesting bees, that no structural differences of the feet 
or mandibles can be detected between the workers in cotton and 
the workers in resin ; and he also says that in the case where 
two cells are constructed in one snail-shell, a male individual is 
produced from the cell of the greater capacity, and a female from 
the other. 

Osmia is one of the most important of the genera of bees 
found in Europe, and is remarkable for the diversity of instinct 
displayed in the formation of the nests of the various species. 
As a rule they avail themselves for nidification of hollow 
places already existing ; choosing excavations in wood, in the 

mortar of walls, and even in 
sandbanks ; in several cases 
the same species is found to 
be able to adapt itself to 
more than one kind of these 
very different substances. This 
variety of habit will render 
it impossible for us to do 
justice to this interesting 
genus within the space at 




FIG. 21. Osmia tricornis, ?. Algeria. 



our disposal, and we must 
content ourselves with a con- 
sideration of one or two of the more instructive of the traits 
of Ositiia life. 0. tridentata forms its nest in the steins of 
brambles, of which it excavates the pith ; its mode of working 
and some other details of its life have been well depicted by 
Fabre. The Insect having selected a suitable bramble-stalk with 
a cut extremity, forms a cylindrical burrow in the pith thereof, 
extending the tunnel as far as will be required to allow the 
construction of ten or more "cells placed one after the other in 
the axis of the cylinder ; the bee does not at first clear out quite 
all the pith, but merely forms a tunnel through it, and then 
commences the construction of the first cell, which is placed at 
the end of the tunnel that is most remote from the entrance. 
This cavity is to be of oval form, and the Insect therefore cuts 
away more of the pith so as to make an oval space, but somewhat 
truncate, as it were, at each end, the plane of truncation at the 
proximal extremity being of course an orifice. The first cell 



i DASYGASTRES OSMIA 49 

thus made is stored with pollen and honey, and an egg is 
deposited. Then a barrier has to lie constructed to close this 
chamber ; the material used for the barrier is the pith of the 
stem, and the Insect cuts the material required for the purpose 
from the walls of the second chamber ; the excavation of the 
second chamber is, in fact, made to furnish the material for clos- 
ing up the first cell. In this way a chain of cells is constructed, 
their number being sometimes as many as fifteen. The mode 
in which the bees, when the transformations of the larvae and 
pupae have been completed, escape from the chain of cells, has 
been the subject of much discussion, and errors have arisen from 
inference being allowed to take the place of observation. Thus 
Dufour, who noted this same mode of construction and arrange- 
ment in another Hymenopteron (Odyncrus nidulator}, perceived 
that there was only one orifice of exit, and also that the Insect 
that was placed at the greatest distance from this was the 
one that, being the oldest of the series, might be expected to 
be the first ready to emerge ; and as the other cocoons would 
necessarily be in the way of its getting out, he concluded that 
the egg that \vas last laid produced the first Insect ready for 
emergence. Fabre tested this by some ingenious experiments, 
and found that this was not the case, but that the Insects became 
ready to leave their place of imprisonment without any reference 
to the order in which the eggs were laid, and he further noticed 
some very curious facts with reference to the mode of emergence 
of Osmia tridentata. Each Insect, when it desires to leave the 
bramble stem, tears open the cocoon in which it is enclosed, and also 
bites through the barrier placed by the mother between it and 
the Insect that is next it, and that separates it from the orifice 
of exit. Of course, if it happen to be the outside one of the 
series it can then escape at once ; but if it should lie one farther 
down in the Indian file it will not touch the cocoon beyond, but 
waits patiently, possibly for days ; if it then still find itself con- 
fined it endeavours to escape by squeezing past the cocoon that 
intervenes between it and liberty, and by biting away the material 
at the sides so as to enlarge the passage ; it may succeed in doing 
this, and so get out, but if it fail to make a side passage it will 
not touch the cocoons that are in its way. In the ordinary 
course of events, supposing all to go well with the family, all 
the cocoons produce their inmates in a state for emergence within 

VOL. VI E 



50 HYMENOPTERA CHAP. 

a week or two, and so all get out. Frequently, however, the 
emergence is prevented by something having gone wrong with 
one of the outer Insects, in which case all beyond it perish unless 
they are strong enough to bite a hole through the sides of the 
bramble-stem. Thus it appears that whether a particular Osmia 
shall be able to emerge or not depends on two things (1) whether 
all goes well with all the other Insects between it and the orifice, 
and (2) whether the Insect can bite a lateral hole or not; this 
latter point also largely depends on the thickness of the outer 
part of the stem of the bramble. Fabre's experiments on 
these points have been repeated, and his results confirmed by 
Nicolas. 

The fact that an Osmia would itself perish rather than attack 
the cocoon of its brother or sister is certainly very remarkable, 
and it induced Fabre to make some further experiments. He 
took some cocoons containing dead specimens of Osmia, and placed 
them in the road of an Osmia ready for exit, and found that in 
such case the bee made its way out by demolishing without any 
scruple the cocoons and dead larvae that intervened between it 
and liberty. He then took some other reeds, and blocked the 
way of exit with cocoons containing living larvae, but of another 
species of Hymenoptera. Solcnius vagus and Osmia detrita were 
the species experimented on in this case, and he found that the 
Osmia destroyed the cocoon and living larvae of the Solenius, 
and so made its way out. Thus it appears that Osmia will 
respect the life of its own species, and die rather than destroy it, 
but has no similar respect for the life of another species. 

Some of Fabre's most instructive chapters are devoted to the 
habits and instincts of various species of the genus Osmia. It 
is impossible here to find space even to summarise them, still 
more impossible to do them justice ; but we have selected the 
history just recounted, because it is rare to find in the insect 
world instances of such self-sacrifice by an individual for one of 
the same generation. It would be quite improper to generalise 
from this case, however, and conclude that such respect for its 
own species is common even amongst the Osmia. Fabre, indeed, 
relates a case that offers a sad contrast to the scene of self- 
sacrifice and respect for the rights of others that we have roughly 
portrayed. He was able to induce a colony of Osmia tricornis 
(another species of the genus, be it noted) to establish itself and 



i DASYGASTRES LEAF-CUTTING BEES 5 I 

work in a series of glass tubes that he placed on a table in his 
laboratory. He marked various individuals, so that he was able 
to recognise them and note the progress of their industrial works. 
Quite a large number of specimens thus established themselves 
and concluded their work before his very eyes. Some individuals, 
however, when they had completed the formation of a series of 
cells in a glass tube or in a reed, had still not entirely completed 
their tale of work. It would be supposed that in such a case 
the individual would commence the formation of another series 
of cells in an unoccupied tube. This was not, however, the case. 
The bee preferred tearing open one or more cells already completed 
in some cases, even by itself scattering the contents, and de- 
vouring the egg ; then again provisioning the cell, it would deposit 
a fresh egg, and close the chamber. These brief remarks will 
perhaps suffice to give some idea of the variety of instinct and 
habit that prevails in this very interesting genus. Friese observes 
that the variety of habits in this genus is accompanied as a rule 
by paucity of individuals of a species, so that in central Europe 
a collector must be prepared to give some twenty years or so of 
attention to the genus before he can consider he has obtained all 
the species of Osmia that inhabit his district. 

As a prelude to the remarks we are about to make on the 
leaf-cutting bees of the genus Meyachile it is well to state that 
the bee, the habits of which were described by Eeaumur under 
the name of " 1'abeille tapissiere," and that uses portions of the 
leaves of the scarlet poppy to line its nest, is now assigned to 
the genus Osmia, although Latreille, in the interval that has 
elapsed since the publication of Reaumur's work, founded the 
genus Antliocopa for the bee in question. Megacliile is one of 
the most important of the genera of the Dasygastres, being 
found in most parts of the world, even in the Sandwich 
Islands ; it consists of bees averaging about the size of 
the honey-bee (though some are considerably larger, others 
smaller), and having the labrum largely developed ; this organ is 
capable of complete inflection to the under side of the head, and 
when in the condition of repose it is thus infolded, it underlaps 
and protects the larger part of the lower lip ; the mandibles 
close over the infolded labrum, so that, when the Insect is at rest, 
this appears to be altogether absent. These bees are called 
leaf-cutters, from their habit of forming the cells for their nest 



HYMENOPTERA 



CHAP. 



out of pieces of the leaves of plants. We have several species in 
Britain ; they are very like the common honey-bee in general 
appearance, though rather more robustly formed. These Insects, 
like the Osmiae, avail themselves of existing hollow places as 
receptacles in which to place their nests. M. alljocincta frequently 
takes possession of a deserted worm-burrow in the ground. The 
burrow being longer than necessary the bee commences by cutting 
off the more distant part by means of a barricade of foliage ; this 
being done, it proceeds to form a series of cells, each shaped like 
a thimble with a lid at the open end (Fig. 22, A). The body of 
the thimble is formed of large oval pieces of leaf, the lid of 
smaller round pieces ; the fragments are cut with great skill from 
the leaves of growing 1 plants by the Insect, which seems to have 
an idea of the form and size of the piece of foliage necessary for 
each particular stage of its work. 

Home has given particulars as to the nest of MegacTiile a utltm- 
cina (fascieulata), an East Indian species. 1 The material employed 

was either the leaves of the Indian 
pulse or of the rose. Long pieces 
are cut by the Insect from the 
leaf, and with these a cell is formed ; 
a circular piece is next cut, and 
with this a lid is made for the 
receptacle. The cells are about 
the size and shape of a common 
thimble ; in one specimen that 
Home examined no less than 
thirty-two pieces of leaf disposed 
in seven layers w T ere used for one 
cell, in addition to three pieces for 
the round top. The cells are 
carefully prepared, and some kind 
of matter of a gummy nature is 
believed to be used to keep in 

FIG. 22. Nidification o( leaf -cutting 

bee, Megachile anthracina. A, ..m- place the pieces forming the ill- 
cell separated with lid open; the terior * The ceR , d 
larva (a) reposing on the foocl ; B, 

p;ut of a string of the cells. (After end to end, as shown in Fig. 22, B ; 

five to seven cells form a series, 
and four or six series are believed to be constructed by one pair 

1 Trims. Zool. ,SW. London, vii. 1870, p. 178. 




i BEES DASYGASTRES SOCIALES 5 3 

of this bee, the mass being located in a hollow in masonry or 
some similar position. Each cell when completed is half filled 
with pollen in the usual manner, and an egg is then laid in it. 
This bee is much infested by parasites, and is eaten by the Grey 
Hornbill (Meniceros Mcornis*). 

Megachile lanata is one of the Hymenoptera that in East 
India enter houses to build their own habitations. According 
to Home both sexes take part in the work of construction, and 
the spots chosen are frequently of a very odd nature. The 
material used is some kind of clay, and the natural situation 
may be considered to be the interior of a hollow tube, such as 
the stem of a bamboo ; but the barrel of a gun, and the hollow 
in the back of a book that has been left lying open, have been 
occasionally selected by the Insect as suitable. Smith states 
that the individuals developed in the lower part of a tubular 
series of this species were females, " which sex takes longer to 
develop, and thus an exit is not required for them so soon as for 
the occupants of the upper cells which are males." J/. proxima, a 
species almost exactly similar in appearance to M. lanata, makes 
its cells of leaf-cuttings, however, and places them in soft soil. 

Eabre states that M. albocincta, which commences the 
formation of its nest in a worm -burrow by means of a barricade, 
frequently makes the barricade, but no nest ; sometimes it will 
indeed make the barricade more than twice the proper size, and 
thus completely fill up the worm burrow. Fabre considers that 
these eccentric proceedings are due to individuals that have already 
formed proper nests elsewhere, and that after completing these 
have still some strength remaining, which they use up in this 
fruitless manner. 

The Social bees (SOCIALES) include, so far as is yet known, 
only a very small number of genera, and are so diverse, both in 
habits and structure, that the propriety of associating them in 
one group is more than doubtful ; the genera are Bomlms (Fig. 
.'!.">1, vol. v.), with its commensal genus or section, Ps//A///-//.s 
(Fig. 23); Melipona (Fig. 24), in which Trigona and Tetragona 
may at present be included, and Apis (Fig. 6) ; this latter genus 
comprising the various honey-bees that are more or less com- 
pletely domesticated in different parts of the world. 

In the genus Boml'us the phenomena connected with the 
social life are more similar to what we find among wasps 



54 HYMENOPTERA CHAP. 

than to what they are in the genus Apis. The societies come to 
an end at the close of the season, a few females live through the 
winter, and each of these starts a new colony in the following 
spring. Males, females and workers exist, but the latter are 
not distinguished by any good characters from the females, and 
are, in fact, nothing but more or less imperfect forms thereof; 
whereas in Apis the workers are distinguished by structural 
characters not found in either of the true sexes. 

Hotter has given a description of the commencement of a 
society of Bonibus lapidarius. 1 A large female, at the end of May, 
collected together a small mass of moss, then made an expedition 
and returned laden with pollen ; under cover of the moss a cell 
was formed of wax taken from the hind-body and mixed with 
the pollen the bee had brought in ; this cell was fastened to a 
piece of wood ; when completed it formed a subspherical recep- 
tacle, the outer wall of which consisted of wax, and whose interior 
w r as lined with honey-saturated pollen ; then several eggs were 
laid in this receptacle, and it was entirely closed. Hotter took 
the completed cell away to use it for museum purposes, and the 
following day the poor bee that had formed it died. From 
observations made on Bomb us agronun he was able to describe 
the subsequent operations ; these are somewhat as follows : The 
tirst cell being constructed, stored, and closed, the industrious 
architect, clinging to the cell, takes a few days' rest, and after 
this interval commences the formation of a second cell ; this is 
placed by the side of the first, to which it is connected by a 
mixture of wax and pollen ; the second cell being completed a 
third may be formed ; but the labours of the constructor about 
this time are augmented by the hatching of the eggs deposited 
a few days previously ; for the young larvae, having soon disposed 
of the small quantity of food in the interior of the waxen cell, 
require feeding. This operation is carried on by forming a small 
opening in the upper part of the cell, through which the bee 
conveys food to the interior by ejecting it from her mouth 
through the hole ; whether the food is conveyed directly to 
the mouths of the larvae or not, Hotter was unable to observe ; 
it being much more difficult to approach this royal founder 
without disturbing her than it is the worker-bees that carry on 
similar occupations at a subsequent period in the history of the 
1 Mt. Ver. Steiermark, xxxi. 1882, p. 69. 



SOCIALES BUMBLE-BEES 5 5 



society. The larvae in the first cell, as they increase in size, 
apparently distend the cell in an irregular manner, so that it 
becomes a knobbed and rugged, truffle-like mass. The same 
thing happens with the other cells formed by the queen. Each 
of these larval masses contains, it should be noticed, sister-larvae 
all of one age ; when full grown they pupate in the mass, and 
it is worthy of remark that although all the eggs in one larval 
mass were laid at the same time, yet the larvae do not all pupate 
simultaneously, neither do all the perfect Insects appear at 
once, even if all are of one sex. The pupation takes place 
in a cocoon that each larva forms for itself of excessively 
fine silk. The first broods hatched are formed chiefly, if not 
entirely, of workers, but small females may be produced before 
the end of the season. Huber and Schmiedeknecht state that 
though the queen provides the \vorker-cells with food before the 
eggs are placed therein, yet no food is put in the cells in which 
males and females are produced. The queen, at the time of 
pupation of the larvae, scrapes away the wax by which the 
cocoons are covered, thus facilitating the escape of the per- 
fect Insect, and, it may also be, aiding the access of air to the 
pupa. The colony at first grows very slowly, as the queen can, 
unaided, feed only a small number of larvae. But after she 
receives the assistance of the first batch of workers much more 
rapid progress is made, the queen greatly restricting her labours, 
and occupying herself with the laying of eggs; a process that 
now proceeds more and more rapidly, the queen in some cases 
scarcely ever leaving the nest, and in others even becoming 
incapable of flight. The females produced during the inter- 
mediate period of the colony are smaller than the mother, but 
supplement her in the process of egg - laying, as also do the 
workers to a greater or less extent. The conditions that deter- 
mine the egg-laying powers of these small females and workers 
are apparently unknown, but it is ascertained that these powers 
vary greatly in different cases, so that if the true queen die the 
continuation of the colony is sometimes effectively carried on by 
these her former subordinates. In other cases, however, the 
reverse happens, and none of the inhabitants may be capable of 
producing eggs : in this event two conditions may be present ; 
either larvae may exist in the nest, or they may be absent. In 
the former case the workers provide them with food, and the 



56 HYMENOPTERA CHAP. 

colony may thus still be continued ; but in the latter case, 
there being no profitable occupation for the bees to follow, 
they spend the greater part of the time sitting at home in the 
nest. 

Supposing all to go well with the colony it increases very 
greatly, but its prosperity is checked in the autumn ; at this 
period large numbers of males are produced as well as new 
queens, and thereafter the colony comes to an end, only a few 
fertilised females surviving the winter, each one to commence for 
herself a new colony in the ensuing spring. 

The interior of the nest of a bumble-bee (Boinbus) frequently 
presents a very irregular appearance ; this is largely owing to the 
fact that these bees do not use the cells as cradles twice, but form 
others as they may be required, on the old remains. The cells, 
moreover, are of different sizes, those that produce workers being 
the smallest, those that cradle females being the largest, while 
those in which males are reared are intermediate in size. 
Although the old cells are not used a second time for rearing 
brood they are nevertheless frequently adapted to the purposes 
of receptacles for pollen and for honey, and for these objects they 
may be increased in size and altered in form. 

It may be gathered from various records that the period 
required to complete the development of the individual Bombus 
about midsummer is four weeks from the deposition of the egg 
to the emergence of the perfect Insect, but exact details and 
information as to whether this period varies with the sex of the 
Insect developed are not to be found. The records do not 
afford any reason for supposing that such distinction will be 
found to exist : the size of the cells appears the only correlation, 
suggested by the facts yet known, between the sex of the in- 
dividual and the circumstances of development. 

The colonies of Bombus vary greatly in prosperity, if W 7 e take 
as the test of this the number of individuals produced in a 
colony. They never, however, attain anything at all approach- 
ing to the vast number of individuals that compose a large colony 
of wasps, or that exist in the crowded societies of the more 
perfectly social bees. A populous colony of a subterranean 
Bombus may attain the number of oOO or 400 individuals. 
Those that dwell on the surface are as a rule much less populous, 
as they are less protected, so that changes of weather are more 



i SOCIALES NESTS OF BOMBUS 57 

prejudicial to them. According to Smith, the average number 
of a colony of B. muscorum in the autumn in this country is 
about 120 viz. 25 females, 36 males, 59 workers. No mode 
of increasing the nests in a systematic manner exists in this 
genus ; they do not place the cells in stories as the wasps do ; 
and this is the case notwithstanding the fact that a cell is not 
twice used for the rearing of young. When the ground-space 
available for cell-building is filled the Bombus begins another 
series of cells on the ruins of the first one. From this reason 
old nests have a very irregular appearance, and this condition of 
seeming disorder is greatly increased by the very different sizes 
of the cells themselves. We have already alluded to some of 
these cells, more particularly to those of different capacities to 
suit the sexes of the individuals to be reared in them. In 
addition to these there are honey - tubs, pollen-tubs, and 
the cells of the Psithyrus (Fig. 23), the parasitic but friendly 
inmates of the Bombus- nests. A nest of Bombus, exhibiting 
the various pots projecting from the remains of empty and 
partially destroyed cells, presents, as may well be imagined, a very 
curious appearance. Some of the old cells apparently are partly 
destroyed for the sake of the material they are composed of. 
Others are formed into honey- tubs, of a make-shift nature. It 
must be recollected that, as a colony increases, stores of pro- 
visions become absolutely necessary, otherwise in bad weather 
the larvae could not be fed. In good weather, and when flowers 
abound, these bees collect and store honey in abundance ; in 
addition to placing it in the empty pupa-cells, they also form 
for it special receptacles ; these are delicate cells made entirely 
of wax filled with honey, and are always left open for the benefit 
of the community. The existence of these honey - tubs in 
1 tumble-bees' nests has become known to our country urchins, 
whose love for honey and for the sport of bee-baiting leads to 
wholesale destruction of the nests. According to Hoffer, special 
tubs for the storing of pollen are sometimes formed ; these are 
much taller than the other cells. The Psithyrus that live in the 
nests with the Bombus are generally somewhat larger than the 
latter, and consequently their cells may be distinguished in the 
nests by their larger size. A bumble-bees' nest, composed of all 
these heterogenous chambers rising out of the ruins of former 
layers of cells, presents a scene of such apparent disorder that 



58 HYMENOPTERA CHAP. 

many have declared that the bumble-bees do not know how to 
build. 

Although the species of Bombus are not comparable with the 
hive-bee in respect of the perfection and intelligent nature of 
their work, jet they are very industrious Insects, and the con- 
struction of the dwelling-places of the subterranean species is 
said to be carried out in some cases with considerable skill, a 
dome of wax being formed as a sort of roof over the brood cells. 
Some work even at night. Tea has recorded the capture of a 
species in Upper Burmah working by moonlight, and the same 
industry may be observed in this country if there be sufficient 
heat as well as light. Godart, about 200 years ago, stated that 
a trumpeter-bee is kept in some nests to rouse the denizens to 
work in the morning : this has been treated as a fable by 
subsequent writers, but is confirmed in a circumstantial manner 
by Hoffer, who observed the performance in a nest of B. ruderatus 
in his laboratory. On the trumpeter being taken away its office 
was the following morning filled by another individual. The 
trumpeting was done as early as three or four o'clock in the 
morning, and it is by no means impossible that the earliness of 
the hour may have had something to do with the fact that for 
200 years no one confirmed the old naturalist's observation. 

One of the most curious facts in connection with Bomlnis is 
the excessive variation that many of the species display in the 
colour of the beautiful hair with which they are so abundantly 
provided. There is not only usually a difference between the 
sexes in this respect, but also extreme variation within the 
limits of the same sex, more especially in the case of the males 
and workers ; there is also an astonishing difference in the size 
of individuals. These variations are carried to such an extent 
that it is almost impossible to discriminate all the varieties of a 
species by inspection of the superficial characters. The struc- 
tures peculiar to the male, as well as the sting of the female, 
enable the species to be determined with tolerable certainty. 
Cholodkovsky, 1 on whose authority this statement as to the sting 
is n uide, has not examined it iu the workers, so that we do not 
know whether it is as invariable in them as he states it to be 
in queens of the same species. According to Handlirsch,' each 

1 Zont. An;, vii. 1884, p. 312. 
2 SJ3. Gcs. Wien. xxxviii. 1888, p. 34. 



SOCIALES PSITHYRUS 59 



species of Bombus has the capacity of variation, and many of the 
varieties are found in one nest, that is, among the offspring of a 
single pair of the species, but many of the variations are restricted 
to certain localities. Some of the forms can be considered as 
actual (" fertige ") species, intermediate forms not being found, and 
even the characters by which species are recognised being some- 
what modified. As examples of this he mentions Bombus silvarum 
and B. arenicula, B.pratorum and B. scrim sJtiranus. In other cases, 
however, the varieties are not so discontinuous, intermediate forms 
being numerous ; this condition is more common than the one we 
have previously described ; B. terrestris, B. horturum, B. lapidarius 
and B. pomorum are examples of these variable species. The 
variation runs to a considerable extent in parallel lines in the 
different species, there being a dark and a light form of each ; also 
each species that has a white termination to the body appears in 
a form with a red termination, and vice versd. In the Caucasus 
many species that have everywhere else yellow bands possess 
them white ; and in Corsica there are species that are entirely 
black, with a red termination to the body, though in continental 
Europe the same species exhibit yellow bands and a white ter- 
mination to the body. AVith so much variation it will be readily 
believed that much remains to be done in the study of this 
fascinating genus. It is rich in species in the Northern hemi- 
sphere, but poor in the Southern one, and in both the Ethiopian 
and Australian regions it is thought to be entirely wanting. 

The species of the genus Psithyrus (Apatkus of many authors) 
inhabit the nests of Bombus; although less numerous than the 
species of the latter genus, they also are widely distributed. They 
are so like Bombus in appearance that they were not distinguished 
from them by the earlier entomologists ; and what is still more 
remarkable, each species of Psithyrus resembles the Bombus with 
which it usually lives. There appear, however, to be occasional 
exceptions to this rule, Smith having seen one of the yellow- 
banded Psithyrus in the nest of a red-tailed Bombus. Psitliyrus 
is chiefly distinguished from Bombus by the absence of certain 
characters that fit the latter Insects for their industrial life ; the 
hind tibiae have no smooth space for the conveyance of pollen, 
and, so far as is known, there are only two sexes, males and per- 
fect females. The Bombus and Psithyrus live together on the 
best terms, and it appears probable that the latter do the former 



6o 



HYMENOPTERA 



CHAP. 



no harm beyond appropriating a portion of their food supplies. 
Schmiedeknecht says they are commensals, not parasites ; Lut it 
must be admitted that singularly few descriptions of the habits 
and life-histories of these interesting Insects have been recorded. 

Hoffer has, however, 
made a few direct 
observations which 
confirm, and at the 
same time make 
UK ire definite, the 
vague ideas that 
have been generally 
prevalent among 
He 
took 




entomologists. 



FIG. 23. Psitliyrus restalix, Britain. A. Female, x :] ; 
B. outer side of hind lee. 



found and 
home a nest 
bus variabilis, which 
contained also a 
female of Psitliyrus 

ccinipestris, so that he was able to make observations on the two. 
The PsitJiyrus was much less industrious than the Bombus, and 
only left the nest somewhat before noon, returning home again 
towards evening ; after about a month this specimen became still 
more inactive, and passed entire days in the nest, occupying itself 
in consuming the stores of honey of its hosts, of which very large 
quantities were absorbed, the Psitliyrus being much larger than the 
host-bee. The cells in which the young of the Psithyrus are hatched 
are very much larger than those of the Bombus, and, it may therefore 
be presumed, are formed by the Psitliyrus itself, for it can scarcely 
be supposed that the Bv minis carries its complaisance so far as 
to construct a cell specially adapted to the superior stature of its 
uninvited boarder. When a Psitkyrus has been for some time a 
regular inhabitant of a nest, the Bombus take its return home from 
time to time as a matter of course, displaying no emotion what- 
ever at its entry. Occasionally Hoffer tried the introduction of 
a Psitliyrus to a nest that had not previously had one as an in- 
mate. The new arrival caused a great hubbub among the Bombus, 
which rushed to it as if to attack it, but did not do so, and the 
alarm soon subsided, the Psitliyrus taking up the position in 
the nest usually affected by the individuals of the species. On 



i SOCIALES STINGLESS BEES 6 I 

introducing a female Psitliyrus to a nest of Bombus in which a 
Psitliyrus was already present as an established guest, the latter 
asserted its rights and drove away the new comer. Hoffer also 
tried the experiment of placing a Psitliyrus campestris in the nest 
of Bombus lapidarius a species to which it was a stranger ; not- 
withstanding its haste to fly away, it was at once attacked by 
the Bombus, who pulled it about but did not attempt to sting it. 

When Psitliyrus is present in a nest of Bombus it apparently 
affects the inhabitants only by diminishing their stores of food to 
so great an extent that the colony remains small instead of largely 
increasing in numbers. Although Bombus variabilis, when left 
to itself, increases the number of individuals in a colony to 200 
or more, Hoffer found in a nest in which Psithyrus was present, 
that on the 1st of September the assemblage consisted only of a 
queen Bombus and fifteen workers, together with eighteen speci- 
mens of the Psitliyrus, eight of these being females. 

The nests of Bombus are destroyed by several animals, probably 
for the sake of the honey contained in the pots ; various kinds 
of small mammals, such as mice, the weasel, and even the fox, 
are known to destroy them ; and quite a fauna of Insects may be 
found in them ; the relations of these to their hosts are very little 
known, but some undoubtedly destroy the bees' larvae, as in the 
case of Meloe, Mutilla and Conops. Birds do not as a rule attack 
these bees, though the bee-eater, M<:r<>p* "jiii'stcr, has been known 
to feed on them very heavily. 

The genera of social bees known as Melipona, Trigona or Tetra- 
gona, may, according to recent authorities, be all included in one 
genus, Melipona. Some of these Insects are amongst the smallest 
of bees, so that one, or more, species go by the name of " Mosquito- 
bees." The species appear to be numerous, and occur in most of the 
tropical parts of the continents of the world, but unfortunately 
very little is known as to their life-histories or economics ; they 
are said to form communities consisting at times of a countless 
number of individuals ; but it has not been thoroughly ascer- 
tained whether these are the produce of a single queen, as in 
the case of the hive-bee, or whether there may be more than one 
egg-producer in each community. The late F. Smith thought 
the former of these alternatives would prove to lie correct. 
These mosquito-bees are frequently spoken of as stingless bees, 
but this is not quite correct, for although they do not sting, 



62 



HYMENOPTERA 



CHAP. 



von Ihering 1 says that all the essential elements of the sting 
are present, the pointed or penetrating part of the apparatus 
being stunted. 

It would serve no useful purpose to attempt to construct the 
social history of these stingless bees from the numerous brief 
scattered accounts in entomological literature, for they refer to 
different species ; it is, however, positively stated by Smith on the 
authority of Peckolt ~ that Trigona mosquito sends off swarms after- 
the manner of the hive-bee in this country, and that after search- 
ing six hives only one royal female could be found in each. 

The nests of many of these little bees are rich in honey, and 
they have a host of enemies from man and monkeys downwards : 

and as they do not defend 
themselves by stinging, it might 
be supposed they would have 
but a poor time of it. From 
the accounts that have been 
published we may, however, 
gather that they are rich in 
devices for the protection of 
their nests, and for the exclu- 
sion of intruders. Bates has 
given some particulars as to 
Mi-lipona, interrupts (fasci- 
culat(i) ; it is about one-third 
shorter than the hive-bee, and its colonies are composed of an im- 
mense number of individuals. The workers are usually occupied 
in gathering pollen ; but they also collect clay in a similar manner, 
and convey it to the nest, where it is used for building a wall to 
complete the fortification of the nest, which is placed either in a 
suitable bank, or in a trunk of a tree; in either situation it is 
completely built in with clay. A nest which Bates saw opened 
contained about two quarts of pleasantly-tasted liquid honey. 
Forty-five species of these little bees were found in different 
parts of the Amazons Valley, the largest kind being half an inch 
in length, the smallest very minute, not more than one-twelfth 
of an inch. These little creatures are thus masons as well 
as workers in wax and resin, and they are also gatherers of 
nectar, pollen, and resin. 

1 Hut. Xtn-hr. xii. 1866, p. 177. ' Tr. cut. Xoc. London, 1868, p. 133. 




FIG. 24. Mclipima sp. 9. Amazons. 



i SOCIALES STINGLESS BEES 63 

According to Gosse, one of these bees is well known in Jamaica, 
where they are called " Angelitos," in consquence of their not sting- 
ing people. He observed a nest of this bee in a tree, and found 
it to be much infested by black ants anxious to obtain entrance 
to it ; three bees, however, stood sentinel in the entrance, so as to 
completely block it and keep out intruders, but the middle bee 
moved on one side out of the way directly one of its fellows 
wished to come in or out of the nest. The honey accumulated 
by this species is kept in clusters of cups about the size of a 
pigeon's egg, at the bottom of the hive and away from the brood- 
cells. The queen or mother-bee is lighter in colour than the 
others, and has the hind body twice the length of theirs. 

Hoc-kings 1 has given us some details as to the natural history 
of two of these bees that inhabit Australia, where they are called 
" Karbi " and " Kootchar," the first being, it is supposed, Trigona 
carbonaria, Smith: it is usually about three -sixteenths of an 
inch in length, the queen, when fully developed, being nearly 
twice that length. The comb is built in a most peculiar form, 
being, it is said, in the shape of a spiral staircase, and tapering 
towards the ends : honey -pots and pollen are constructed for 
the storage of food. The comb is encased in wax, and outside it 
a labyrinth of waxen passages is formed. The entrance to the 
colony is guarded by a line of bees who inspect every one that 
arrives, and it is surprising to see how soon a stranger is dis- 
covered and pounced upon before it has time even to alight ; the 
intruder, when caught, is held by several bees, who put it on the 
rack by holding and stretching out its limbs to their full extent, 
retaining it in this position for as long as an hour, by which time 
the unfortunate prisoner is usually dead. These bees, as well as 
'many other allied species, fight desperately with their mandibles, 
and are apparently of a very fierce disposition. The other 
species, called " Kootchar," is said to produce a very large number 
of drones, and the habits and dispositions of the bees differ con- 
siderably from those of the " Karbi " : the entrance to their hive is 
guarded by a pipe of propolis (a sort of resinous wax) about an inch 
in length, having an exceedingly sticky outer edge, and it is by this 
pipe alone that access to the interior can be gained. At night 
the entrance is closed by numerous minute globules of semi-fluid 
gum placed against it, thus forming a thin wall full of air-holes. 

1 Tr. cnt. Soc. London, 1884, p. 149. 



64 HYMENOPTERA 



CHAP. 



The colonies of " Kootchar " can be united by taking away a queen 
and then packing her brood-nest, bees and all, against that of 
the colony it is to be joined to. This cannot be done with the 
' Karbi." The account given by Mr. Hockings contains a great 
many other interesting details, and there can be no doubt that 
a full account of the natural history of these Insects would be 
very instructive. 

Fritz Miiller has recorded a singular case bearing on the 
instinct of these social Insects : he says that a nest of a small 
Triyona was built in a hollow tree, and that as a consequence of 
the irregularity of the hole the bees were obliged to give a very 
irregular shape to their combs of honey. These bees were 
captured and put in a spacious box (presumably together with 
the irregular comb, but this he unfortunately does not mention) : 
after a year, " when perhaps not a single bee survived of those 
which had come from the canella tree," they still continued to 
build irregular combs, though quite regular combs were built by 
several other communities of the same species that he had kept. 
These bees, he also tells us, do not use pure wax for the construc- 
tion of their combs, but mix it with resin or gum that gives it a 
peculiar odour and appearance. He captured two communities 
of a common Melipona, one of which had the combs made of dark 
reddish brown, the other of pale yellowish brown, wax, and in 
captivity in a distant locality each of the tw r o communities 
continued to form its comb in the same way, thus showing the 
continuity that prevails in these cases as long as circumstances 
permit. Miiller thinks this due to imitation, but it seems at 
least as probable that it is due to perception of the properties of 
the nest. The nest has a certain colour that the worker-bee 
matches. 

Several species of the Melipona, and Triijoim were imported 
from Brazil to France, and kept there for some time in captivity 
by M. Drory. Girard has published 1 some details as to these 
colonies, and is of opinion that some of them indicate an in- 
telligence or instinct superior to that of the honey-bee. The 
<|iir-rn-bce of M. scutell<irix seems to display more intelligence 
than the corresponding sex of A. mellifica. The mode of feeding 
the larvae apparently differs from that of A. mellifica, a provision 
of pollen being first placed in the cell, then some honey ; when 

1 .hni. Soc. enl. France (5), iv. 187-4, p. 567. 



i SOCIALES HONEY-BEE 65 

sufficient food for the whole consumption of a larva is accumulated 
the queen deposits an egg in the cell, which is at once completely 
closed by the worker. The interior of the abode of these bees is 
quite dark, only a very small orifice being left, and in this a sen- 
tinel is constantly on the alert. The same writer states that 
Tr'njona crassipcs has the very peculiar habit of always locating 
its brood-comb in the nest of a species of Termes. 

The honey-bee, Apis mellijiccc (Fig. 6), is considered the highest 
form attained by the Anthophilous division of the Hymenoptera. 
The differentiation of the three forms, male, female, and worker, 
is here carried to a greater degree of perfection than in the other 
bees. The drones are the males ; the individuals we see gather- 
ing honey are always workers, neither the male nor the female in 
this species taking any part in procuring food for themselves or 
for the colony. In addition to this the colonies formed may be 
described as permanent : they do not come to an end at the close 
of one season, and provision is made for the formation of a new 
colony while the old one still persists, by means of a peculiar pro- 
cess called swarming. The life-history of Apis mellifica and its 
anatomy and physiology have been discussed in a whole library 
of works, and we need only notice the chief features. When a 
swarm of bees leaves a hive it consists of the queen-bee or 
female, and a number of workers, these latter being, in fact, the 
surplus population that has been produced in the hive. The 
swarm is not a nuptial flight, as is often supposed, but an act 
of emigration. When this swarm has been housed, the bees 
commence operations in their new quarters, by secreting wax ; 
they are enabled to do this by having consumed much saccharine 
food ; the wax is produced by means of glands in the hind-body 
over the inner faces of the ventral plates of the abdominal rings, 
and it makes its appearance there, after passing from the interior 
of the body through some peculiar membranes on the ventral 
segments, in the form of thin projecting plates. These the bee 
takes off with an apparatus on the hind pair of legs and applies, 
after working up with the mandibles, to form the cells in 
which young ones are to be reared and food stored. A large 
number of bees working in common thus produce the regular 
and beautiful structure known as the comb ; the queen afterwards 
lays an egg in each cell, and as these soon hatch, great labour is 
thrown on the workers, which have then to feed the young ; this 
VOL. VI F 



66 HYMENOPTERA CHAP. 

they do by eating honey and pollen, which, being formed into a 
sort of pap by a portion of their digestive organs, is then re- 
gurgitated and given to the young, a quantity of it being placed 
in the cell, so that the larva is bathed by it, and possibly may 
absorb the food by the skin as well as the mouth. When the 
colony is in good progress and young bees emerge, these act as 
nurses, the older ones cease to prepare food and act as foragers, 
bringing in honey and pollen which are each stored in separate 
cells. The larva in the cell increases its size and sheds a very 
delicate skin several times ; when the larva has reached its 
full size no more food is supplied, but the worker-bees seal up 
the cell by means of a cover formed of pollen and wax, in such a 
manner as to be pervious to air : sealed up in the cell the larva 
spins a cocoon for itself, remains therein for a little time as a 
larva, then changes to a pupa, and thereafter bites its way out 
through the cover of the cell, and appears for the first time as 
a new being in the form of a worker-bee ; the whole process of 
development from the egg-state to the perfect condition of the 
worker-bee occupies about three weeks. 

When the denizens of a hive are about to produce another 
queen, one or more royal cells are formed ; these are much 
larger than the ordinary worker-cells, and of a quite different 
form. In this cell is placed an egg, not differing in any respect 
from the egg that, if placed in an ordinary cell, produces a 
worker ; when the egg has produced a larva this is tended with 
great care and fed throughout its life with royal jelly. This 
food appears to be the same as that supplied to an ordinary 
worker-larva when it is first hatched ; but there is this differ- 
ence, that whereas the worker-larva is weaned, and supplied, 
after the first period of its existence, with food consisting largely 
of honey, pollen and water, the queen-larva is supplied with the 
pap or royal jelly until it is full grown. Some difference of 
opinion exists as to this royal jelly, some thinking that it is a 
different substance from what the workers are fed with ; and it 
is by no means improbable that there may be some difference in 
the secretion of the glands that furnish a part of the material 
composing the pap. The queen is produced more rapidly than 
workers are, about sixteen days being occupied in the process of 
her development. Only one queen is allowed in a hive at a 
time ; so that when several queen-cells are formed, and queen- 



i SOCIALES HONEY-BEE 6/ 



larvae nurtured in them, the first one that is developed into a 
perfect queen goes round and stings the royal nymphs to death 
while they are still in their cells. The production of drones is 
supposed to depend chiefly on the nature of the egg laid by the 
queen ; it being considered that an unfertilised egg is deposited 
for this purpose. There is still some doubt on this point, how- 
ever. Though there is no doubt that drones are produced in 
great numbers from unfertilised eggs, yet there is not evidence 
that they cannot also be produced from fertilised eggs. 1 The 
drone-cells are somewhat larger than the ordinary worker-cells, 
but this is probably not of much import, and it is said that the 
larvae intended to produce drones receive a greater proportion of 
pap than worker-larvae do : about twenty-four days are required 
to produce a drone from the egg. 

From this sketch it will be seen that the production of the 
worker (or third sex, as it is improperly called, the workers 
being really females atrophied in some points and specially 
developed in others) is dependent on the social life, in so far at 
any rate as the special feeding is concerned. There is good 
reason for supposing that A. mellifica has been kept in a state of 
domestication or captivity for an enormous period of time ; and 
this condition has probably led to an increase of its natural 
peculiarities, or perhaps we should say to a change in them to 
suit a life of confinement. This is certainly the case in regard 
to swarming, for this process takes place with comparative 
irregularity in Apis mclUfica in a wild condition. The killing of 
superfluous queens is also probably a phenomenon of captivity, 
for it varies even now in accordance with the numbers of the 
colony. It is interesting to notice that in confinement when a 
swarm goes from the hive it is the old queen that accompanies 
it, and this swarm as a rule settles clown near the old hive, so 
that the queen-bee being already fertilised, the new swarm and 
its subsequent increase are nothing but a division of the old 
hive, the total products of the two having but a single father 
and mother. When a second swarm goes off from a hive it is 
accompanied by a young queen, who frequently, perhaps, in the 
majority of cases, is unfertilised ; this swarm is apt to fly for 
long distances, so that the probability of cross-fertilisation is 

1 See Perez, Act. Soc. Bordeaux, xxxiii. 1880, p. Ixv. ; and Cameron, Tr. Soc. 
Glasgow, 11. s. ii. 1889, }'. 194. 



68 IIVMENOPTERA 



CHAP. 



greatly increased, as the fertilisation of the young new queen is 
effected during a solitary flight she makes after the colony has 
settled down. But in a state of nature the colonies do not send 
off swarms every year or once a -year, but increase to an enormous 
extent, going for years without swarming, and then when their 
home is really filled up send off, it may he presumed, a number 
of swarms in one year. Thus the phenomena of bee-life in a 
wild condition differ considerably from those we see in. artifi- 
cial confinement. And this difference is probably greatly accen- 
tuated by the action of parasites, the proportions of which to their 
guests are in a state of nature liable to become very great ; as 
we have seen to be the case in Jjombus. 

Under these circumstances it is not a matter for surprise 
when we find that the honey-bee has formed distinct races 
analogous to those that exist in the case of the domesticated 
vertebrate animals. The knowledge of these races is, however, 
at present very little advanced, and is complicated by the fact 
that only imperfect information exists as to the true species 
of the genus Apis. There is a bee very like our common honey- 
bee found in southern Europe called A. ligusiica ; this is 
certainly a variety of A. mellifica, and the same remark applies 
to a bee found in Egypt, and called A.fasciata. This gives the 
honey-bee a very wide distribution, extending possibly over the 
whole of the palaearctic region : besides this, the species has been 
introduced into various other parts of the world. 

According to Karsch the honey-bee shows in Germany several 
varieties, all of which belong to the northern form, which may 
be spoken of as the A. domestica of Eay ; the A. ligustica and A. 
fasciata form as we have said distinct races, and it is a remark- 
able fact that these races remain distinct even when imported 
into other climates; though for how long a period of time this 
remains true there is very little evidence to show. The northern 
form, A. domestica, is now found in very widely separated parts 
of the world, in some of which it is wild ; Smith mentions it 
as occurring in the West India islands, throughout the North 
American continent as far south as Mexico, even in Central and 
Southern Africa, and in Australia and New Zealand. The var. 
/it/i/stica has been found also at the Cape of Good Hope. The 
other species known of the genus Apis all belong to the Old 
World, so that there is very little doubt that A. mdlijica is also 



SOCIALES HONEY-BEE 



6 9 





C-: 



a true native of the eastern hemisphere, and its original home 
may possibly have been not far from the shores of the eastern 
portion of the Mediterranean sea. Seven or eight other species 
of Apis are known, all but one of which occur in Asia, ex- 
tending as far as Timor and Celebes. The exceptional one, A. 
adansonii, occurs in tropical Africa and 
in Madagascar. Gerstaecker thought 
these species might be reduced to four, 
but Smith's statement that the males 
and even the workers show good dis- 
tinctive characters seems to be correct. 
Very little is known as to the honey- 
bees of China and Japan. 

The queen-bee greatly resembles the 
worker, but, has the hind body more 
elongated ; she can, however, always be 
distinguished from the worker by the 
absence of the beautiful transverse, 
comb-like series of hairs on the inner 
side of the first joint of the hind foot, 
the planta, as it is called by the bee- 
keeper : she has also no wax plates and 
differs in important anatomical peculi- 
arities. The male bee or drone is very 
different, being of much broader, more 
robust build, and with very large eyes 
that quite meet in the middle of the 
upper part of the head : he also has the 
hind leg differently shaped. The form of 

this limb enables the male of A. mellifica FIG. 25. Portions of hind-feet, 
to be distinguished from the correspond- 
ing sex of allied species of the genus. 

We are indebted to Home for some 
particulars as to the habits of A. dorsata, 
an allied East Indian species. He informs 
us that these bees greatly disfigure buildings, such as the Taj Mahal 
at Agra, by attaching their pendent combs to the marble arches, 
and are so pertinacious that it is almost useless to destroy the 
nests. This bee is said to be so savage in its disposition that it 
cannot be domesticated ; it attacks the sparingly clad Hindoos 







1, of male, 2, of worker, 3, 
<il queen, of the honey-bee; 
series on the left, outer 
faces ; on the right, inner 
faces. , Tip of tibia : b, 
first joint ; c, second joint 
of tarsus. 



70 HYMENOPTERA 



CHAP. 1 



with great ferocity when they disturb its nest. Notwithstand- 
ing its inclination and power to defend its societies this Insect 
appears to be destroyed wholesale. Colonel Eamsay failed to 
establish hives of it, because the Insects were eaten up by lizards. 
The crested honey-buzzard carries off large portions of the comb, 
and devours it on a branch of some tree near by, quite regardless 
of the stings of the bees; while the fondness of bears for the 
honey of the " Dingar," as this species is called, is well known. 



XOTE TO P. 33 : It has just been discovered that a most remarkable 
symbiosis, with structural modification of the bee, exists between the females 
of Xylocopa, of the Oriental sub-genus Koptorthosoma, and certain Acarids. 
A special chamber, with a small orifice for entry, exists in the abdomen of 
the bee, and in this the Acari are lodged. See Perkins, Ent. Mag. xxxv. 
1899, p. 37. 



NOTE TO P. 80 : referring to the habits of social wasps in warm countries. 
The anticipation we ventured to indulge in is shown to be correct by the 
recent observations of Yon Ihering. 1 He states that social wasps in Brazil 
may be divided into two great groups by their habits, viz. 1. Summer com- 
munities, lasting for one year, and founded annually by fertilised females 
that have hibernated example, Polistes ; 2. Perennial communities, founded 
by swarms after the fashion of bee colonies examples, Polybia, Chartergus. 

1 Ann. Not. Hist. (6), xix. 1897, p. 136. 



XOTE TO VOL. V. PP. 545, 546 : The development of Encyrtus fuscicollis 
has now been studied by Marchal, who has discovered the existence of 
embryonic dissociation. The chain of embryos and the epithelial tube in 
which they are placed, are formed as follows : the Encyrtus deposits an egg 
in the interior of the egg of the Hyponomeuta. This does not kill the egg 
of the Lepidopteron, but becomes included in the resulting caterpillar. The 
amnion of the Chalcid egg lengthens, and forms the epithelial tube ; while 
the cells within it become dissociated in such a way as to give rise to a 
chain of embryos, instead of a single embryo. O.K. Ac. Paris, cxxvi. 1898 V 
p. 662, and translation in Ann. Nat. Hist. (7), ii. 1898, p. 28. 



CHAPTER II 

HYMEXOPTERA ACULEATA COXTIXUE1> - DIVISION II. DIPLOPTERA 
OR WASPS - EUMENIDAE, SOLITARY TRUE WASPS - YESPIDAE, 
SOCIAL WASPS - MASARIDAE 

Division II. Diploptera Wasps. 

Anterior wings longitudinally plicate in repose; the pronotum 
extending back, so as to form 
on each side an angle reposing 
on the tea id c i .; the based seg- 
ments of the hind, bodi/ not 
bearing nodes or scales ; the 
hind tarsi formed for simple 
ti-<ill-ing. The species either 
or social in their 




77.. ... . FIG. 26. Upper aspect of pronotum 

habits; some existing rn three and mesonotum of a wasp, - 

forms, males, females, and nes coarctata. a, Angle of prono- 
-. turn ; b, tegula ; c, base of wing ; 

WOT/CeTS. (^ iriesonotum. 

THIS division of Hymenoptera includes the true wasps, but not 
the fossorial wasps. The name applied to it lias been suggested 
by the fact that the front wings become doubled in the long direc- 
t it in when at rest, so as to make them appear narrower than in 
must other Aculeata (Fig. 27). This character is unimportant 
in function so far as we know. 1 and it is not quite constant in 
the division, since some of the Masaridae do not exhibit it. The 
character reappears outside the Diploptera in the genus Zei(cc>-- 

a member of the Chalcididae in the parasitic series of Hymen- 
optera the species of which greatly resemble wasps in coloration. 
A better character is that furnished by the well-marked angle. 

1 Janet lias suggested that the folding is done to keep the delicate hind-margins 
of the wings from being frayed. 



HYMENOPTERA 



CHAP. 



formed by the pronotum on the dorsal part (Fig. 26). By a 
glance at this part a Diplopterous Insect can always be readily 
distinguished. 

Three families are at present distinguished in the Diploptera, 
viz. Eumeiiidae, Vespidae and Masaridae. We anticipate that 
Eumenidae and Vespidae will ultimately be found to constitute 
but one family. 

Fam. 1. Eumenidae Solitary True Wasps. 

Clau's of the feet toothed or Infid ; middle tibiae with only one 
xjn/r at tip. Social assemblages are not formed, mid tin re is 
a n y/-<//7,r/--rr/.s'/V, the duties of nest-construction, etc., Icing 
performed solely l>y the female. 

The Eumenidae, or solitary wasps, are very little noticed by 
the ordinary observer, but they are nevertheless 'more numerous 

than the social Vespidae, about 800 
species being known. In Britain we 
have sixteen species of the solitary, as 
against seven of the social wasps. 
The Eumenidae exhibit a considerable 
diversity in form and structure ; some 
of them have the pedicel at the base 
of the abdomen very elongate, while 
in others this is so short as to be 
imperceptible in the ordinary position 
of the body. A repetition of similar 
differences of form occurs in the social 

wasps, so that notwithstanding the 
FIG. 27. Eumenes flavouicta 9- TO- i i -^ ii i 

Burma. The Wigs on the left difference in habits there seenw to be 
in the position of repose, to 110 satisfactory way of distinguishing 

the members of the two families ex- 
cept by the structure of the claws and tibial spurs. 

Fabre has sketched the habits of a species of Eumenes, 
probably E. pomiformis. This Eumenes constructs with clay a 
small vase-like earthenware vessel, in the walls of which small 
stones are embedded (like Fig. 28, B). This it fills with food 
for the young. The food consists of caterpillars to the number 
of fourteen or sixteen for each nest. These caterpillars art- 
believed to be stung by the parent-wasp (as is the case in the 




II 



DIPLOPTERA WASPS EUMENIDAE 



73 



fossorial Hymenoptera), but complete evidence of this does not 
seem to be extant, and if it be so, the stinging does not 
completely deprive the caterpillars of the capacity of movement, 
for they possess the power of using their mandibles and of 
making strokes, or kicking with the posterior part of the body. 
It is clear that if the delicate egg of the Eumenes or the deli- 
cate larva that issues from it were placed in the midst of a 
mass of this kind, it would probably 
suffer destruction ; therefore, to 
prevent this, the egg is not placed 
among the caterpillars, but is sus- 
pended from the dome covering 
the nest by a delicate thread 
rivalling in fineness the web of the 
spider, and being above the mass 
of food it is safe. When the 
young larva leaves the egg it still 
makes use of the shell as its habit- 
ation, and eats its first meals 
from the vantage-point of this 
suspension; although the mass of FIG. 28. Nidification of solitary wasps: 




the food grows less by consumption, secticm tlnou " h ne g> *' f 

o J_ in'i-its reniformis ; B, of Eumenes 

the little larva is Still enabled to arbiistonnn. , The suspended egg 

reach it by the fact that the egg- ,',. ^,"^ t ? 
shell splits up to a sort of ribbon, 

and thus adds to the length of the suspensory thread, of which it 
is the terminal portion. Finally the heap of caterpillars shrinks 
so much that it cannot be reached by the larva even with the 
aid of the augmented length of the suspensory thread ; by this 
time, however, the little creature has so much increased in size 
and strength that it is able to take its place amongst the food 
without danger of being crushed by the mass, and it afterwards 
completes its metamorphosis in the usual manner. 

It is known that other species of Eumenes construct vase- 
like nests ; E. unguiculata, however, according to an imperfect 
account given by Perris, makes with earth a closed nest of 
irregular shape, containing three cells in one mass. The saliva 
of these builders has the power of acting as a cement, and of 
forming with the clay a very impenetrable material. One 
species, E. coarctata, L. of this genus occurs in Britain. The clay 



74 



HYMENOPTERA 



CHAP. 




to wood : B, 



nests (Fig. 29) of this Insect are often attached to the twigs of 
shrubs, while those of the two species previously mentioned are 

usually placed on objects that offer a 
large surface for fixing the foundations 
to, such as walls. According to Goureau 
the larva of this species forms in one 
corner of its little abode, separated by a 
partition, a sort of dust-heap in which 
it accumulates the various debris re- 
sulting from the consumption of its 
stores. 

Eumenes ccniica, according to Home, 
constructs in Hindostan clay-nests with 
very delicate walls. This species pro- 
visions its nest with ten or Uvelve green 
de- caterpillars ; on one occasion this ob- 
^rver took' from one cell eight green 
tion of the cell. (After caterpillars and one black. It is much 

Andre.) ., , 

attacked by parasites owing, it is 

thought, to the delicacy of the walls of the cells, which are 
easily pierced ; from one group of five cells two specimens only 
of the Eumenes were reared. 

()<li/nerus, with numerous sub-genera, the names of which 
are often used as those of distinct genera, includes the larger 
part of the solitary wasps ; it is very widely distributed over the 
earth, and is represented by many peculiar species even in the 
isolated Archipelago of Hawaii ; in Britain we have about fifteen 
species of the genus. The Odynerus are less accomplished 
architects than the species of Eumenes, and usually play the 
more humble parts of adapters and repairers ; they live either in 
holes in walls, or in posts or other woodwork, or in burrows in 
the earth, or in stems of plants. Several species of the sub- 
genus H<>/i/<ij>/rs have the remarkable habit of constructing 
burrows in sandy ground, and forming at their entry a curvate, 
freely projecting tube placed at right angles to the main bur- 
row, and formed of the grains of sand brought out by the 
Insect during excavation and cemented together. The habits of 
one such species were described by Eeaumur, of another by 
Dufour ; and recently Fabre has added to the accounts of these 
naturalists some important information drawn from his own 



II 



WASPS EUMENIDAE 



75 



observations on 0. reniformis. This Insect provisions its cell 
with small caterpillars to the number of twenty or upwards (Fig. 
28, A.) The egg is deposited before the nest is stocked with food ; 
it is suspended in such a manner that the suspensory thread 
allows the egg to reach well down towards the bottom of the 
cell. The caterpillars placed as food in the nest are all curled 
up, each forming a ring approximately adapted to the calibre of 
the cell. Fabre believes these caterpillars to be partly stupefied 
by stinging, but the act has not been observed either by himself, 
Beaumur, or Dufour. The first caterpillar is eaten by the wasp- 
larva from its point of suspension ; after this first meal has been 
made the larva is supposed to undergo a change of skin ; it then 




FIG. 30. Odynerus antilope ?. Britain. 

abandons the assistance of the suspensory thread, taking up a 
position in the vacant chamber at the end of the cell and draw- 
ing the caterpillars to itself one by one. This arrangement 
permits the caterpillars to be consumed in the order in which 
they were placed in the cell, so that the one that is weakest on 
account of its longer period of starvation is first devoured. 
Fa,bre thinks all the above points are essential to the successful 
development of this wasp-larva, the suspension protecting the egg 
and the young larva from destruction by pressure or movement 
of the caterpillars, while the position of the larva when it leaves 
the thread and takes its place on the floor of the cell ensures its 
consuming the food in the order of introduction ; besides this the 
caterpillars used are of a proper size and of a species the 



76 IIVMENOPTERA 



CHAP. 



individuals of which have the habit of rolling themselves up in a 
ring ; while, as the calibre of the tube is but small, they are 
unable to straighten themselves and move about, so that their 
consumption in proper order is assured. Some interesting 
points in the habits of an allied species, 0. (Pteroclieilus) spinipes 
have been observed by Verhoeff; the facts as regards the con- 
struction and provisioning of the cell are almost the same as in 
0. reniformis. The species of Odynerus are very subject to the 
attacks of parasites, and are, it is well known, destroyed to an 
enormous extent by Chrysididae. Verhoeff says that the wasp 
in question supplied food much infested by entoparasites ; further, 
that a fly, Argi/romoeba sinuata, takes advantage of the habit of 
the Odyne.rtis of leaving its nest open during the process of pro- 
visioning, and deposits also an egg in the nest ; the Odyncrus 
seems, however, to have no power of discovering the fact, or more 
probably has no knowledge of its meaning, and so concludes the 
work of closing the cell in the usual way ; the egg of the 
Ariiyromoelxi hatches, and the maggot produced feeds on the 
caterpillars the wasp intended for its own offspring. Yerhoeff 
observed that the egg of the wasp-larva is destroyed, but he does 
not know whether this was done by the mother Argyromoeba or 
by the larva hatched from her egg. Fabre's observations on 
allied species of Diptera render it, however, highly probable that 
the destruction is effected by the young fly-larva and not by 
the mother-fly. 

Mr. E, 0. L. Perkins once observed several individuals of our 
British 0. callosus forming their nests in a clay bank, and pro- 
visioning them with larvae, nearly all of which were parasitised, 
and that to such an extent as to be evident both to the eye and 
the touch. In a few days after the wasps' eggs were laid, swarms 
of the minute parasites emerged and left no food for the Odynci'n*. 
Curiously, as it would seem, certain of the parasitised and stored- 
up larvae attempted (as parasitised larvae not infrequently do), 
to pupate. From which, as Mr. Perkins remarks, we may infer 
that (owing to distortion) the act of paralysing by the wasp had 
been ineffectual. Mr. Perkins has also observed that some of the 
numerous species of Hawaiian Odynerus make a single mud-cell, 
very like the pot of an Eumenes, but cylindrical instead of 
spherical. This little vessel. is often placed in a, leaf that a 
spider curls up ; young molluscs of the genus Achatinella also 



ii WASPS EUMENIDAE 77 

avail themselves of this shelter, so that a curious colony is formed, 
consisting of the Odynerus in its pot, of masses of the young 
spiders, and of the little molluscs. 

Home has recorded that the East Indian 0, puncttim is fond 
of availing itself of holes in door-posts where large screws have 
heen ; after the hole has been filled with provisions, the orifice 
is covered over level with the surface of the wood so that it 
eludes human observation. It is nevertheless discovered by an 
Ichneumon-fly which pierces the covering with its ovipositor and 
deposits an egg within. 

The genus Abispa is peculiar to Australia and includes some 
very fine solitary wasps, having somewhat the appearance of very 
large Odynerus : these Insects construct a beautiful nest with a 
projecting funnel-shaped entrance, and of so large a size that it 
might pass for the habitation of a colony of social wasps ; it 
appears, however, that this large nest is really formed by a single 
female. 

The species of the genus Rhygchium are also of insecticide 
habits, and appear to prefer the stems of pithy plants as the 
nidus for the development of the generation that is to follow 
them. Lichtenstein says that a female of the European R. 
oculatum forms fifteen to twenty cells in such a situation, and 
destroys 150 to 200 caterpillars, and he suggests that, as it is 
easy to encourage these wasps to nest in a suitable spot, we should 
utilise them to free our gardens from caterpillars, as we do cats 
to clear the mice from our apartments. 

The East Indian R. carnaticum seems to have very similar 
habits to its European congener, adapting for its use the hollow 
stems of bamboos. Home has recorded a case in which a female 
of this species took possession of a stem in which a bee, Megachile 
lanata, had already constructed two cells ; it first formed a parti- 
tion of mud over the spot occupied by the bee, this partition being 
similar to that which it makes use of for separating the spaces 
intended for its own young. This species stores caterpillars for 
the benefit of its larvae, and this is also the case with another 
Eastern species, E. nitidulum. This latter Insect, however, does 
not nidificate in the stems of plants, but constructs clay cells 
similar to those of Uumenes, and fixes them firmly to wood. 
Rhygchium Innineum is said by Sir Bicharcl Owen to obliterate 
hieroglyphic inscriptions in Egypt by its habit of building mud 



78 11YMF,\OPTKRA CIIAI-. 



nests amongst them. An individual ol' this wasp was found 
by Pr. Birch when unrolling a mummy "Then 1 being every 
reason to believe that the In see I hail remained in the position in 
which it was found ever sinee tin 1 last rites were paid to the 
aneieiit Egyptian." 

Fam. 2. Vespidae Social Wasps. 

Claws of the feet simple, neither toothed nor bijid, middle tibiae 
with tico spurs (ft the tip. Insects living in societies, for/// - 
inif a common dicellitii/ of a papery or card-like material ; 
each generation consists of males and females and of workers 
imperfect females that assist the reproductive female by 
carrying on the -industrial occupations. 

The anterior wing possesses four submarginal eells, as in the 
Eumenidae. The attention of entomologists has been more 
directed to the habits and architecture than to the taxonomy of 
these Insects, so that the external structure of the Insects them- 
selves has not been so minutely or extensively scrutinised as is 
desirable; de Saussure, the most important authority, buses his 
classification of the Insects themselves on the nature of the nests 
they form. These habitations consist of an envelope, protecting 
cells similar in form to the comb of the honey-bee, but there is 
this important difference between the two, that while the bee 
forms its comb of wax that it secretes, the wasps make use of 
paper or card that they form from fragments of vegetable tissue, 
more, particularly woodv fibre amalgamated by means of 

IV O J 

cement secreted by glands ; the vegetable fragments are obtained 
by means of the mandibles, the front legs playing a much less 
important part in the economy of the Vespidte than they do in 
that of the bees and fossorial Hymenoptera, 

In most of the nests of Yespida? the comb is placed in stages 
or stories one above the other, and separated by an intervening 
space, but in many cases there is only one mass of comb. It is the 
rule that, when the eells of the comb are only partially formed, eggs 
are deposited in them, and that the larva resulting from the egg 
is fed and tended by the mother, or by her assistants, the workers ; 
as the larvae grow, the cells are increased in correspondence with 
the size of the larva : the subsequent metamorphosis to pupa and 
imauo taking place in the cells after thev have been entirely 

O -L v v 



n 



SOCIAL WASI'S YKSPIDAK 



79 



a 



cjo-e<l. Th'- food supplied is of a v./ried nature according to 
til-- species, l-'-in-j either ;minjal or vegetable, or both. 

Although the nests of the social wasps are very elaborate con- 
structions, y:t they n-rve the purposes of the Insects for only a 
sinide season. This, is '-)-t.;i.iu]y t:be case in our own country. 
H' I-- e;i<-h nest i.-: e<,ii,]j|i.|jcp<| l.y a single female or queen; she 

irst pei Toi ju- unaided all the duties for the inauguration of 

FIG. 31. Section of the 
subterranean nest of the 
common wasp, TV/." 
gerrnanica, in posit 
(After Janet.) , One 
of the chambers of an 
ant's nest, Losing fiavus, 
placed above the wasps' 
nest ; b, root to which 
the first attachment of 
the nest was made ; c, 
secondary attachments ; 
d, the first-made attach- 
ment ; e, a flint within 
the envelopes of the 
nest ; /,the chief suspen- 
sory pillar of the second 
layer of comb ; y, lateral 
galleries ; h, one of the 
secondary pillars of 
suspension between two 
layers of comb ; i, the 
layers of wasp - paper 
forming the envelope of 
nest ; j, vacant space 
round the nest ; k. flints 
that fell to the bottom 
during the work of ex- 
cavation ; I, numerous 
larvae* of any, Pegr/myia 

placed vertically in ground beneath the nest ; m 1 to m 1 , the layers of comb, 
in in- th': cells are indicated , in m s (above the main figure; the arrangement of the 
three cells forming the commencement of the new layer of comb, m 7 , is shown ; n, 
galler irom surface ; o, burrow of a mole ; x>> interval of 90 mm. between top 

of uest and surface ; ^> height of tne nest, 163 mm. 

the '-olouy; she lays the foundation of the cells, deposits the 

_ .- in them, feeds the young, arid thus rears a brood of work 
that ;it on'-'- ;i-.-ist her, and for the future relieve her of a con- 
-i'l'-mljle portion of her former occupations; the nest is by them 
;nl<l'"l to and increased, till the cold weather of the autumn is at 
luind: at tliis time many males and females are produced; the 
cold weather either destroys the inhabitants of the nest, or re- 
duces their vitality so that it is impossible for them to pursue 
successfully the avocations necessary for their subsistence, and 




SO HYMENOPTERA 



CHAP. 



they succumb to adversity. The young females, however, hiber- 
nate, and each one that lives through the winter is the potential 
founder of a new nest in the way we have already described. It 
might be supposed that in tropical countries where no cold 
season occurs the phenomena would be different, that the colonies 
would be permanent, and that the nests would be inhabited until 
they were worn out. De Saussure, however, informs us that this 
is not the case, but that in the tropics also the colonies die off 
annually. " The nests are abandoned," he says, " without it being- 
possible to discover the reason, for apparently neither diminution 
of temperature nor scarcity of food cause them (the Insects) to 
suffer. One is tempted to suppose that the death of the Insects 
is the result of a physiological necessity." 

Nests of Social Wasps. In Europe wasps' nests disappear 
very soon after they are deserted. As it would appear from de 
Saussure's conclusions that in the tropics as well as in the temperate 
regions the rule is that the colonies endure only a portion of one 
year, and that a new nest is commenced by a single founder once 
in twelve months, it is a somewhat remarkable fact that some 
tropical wasp-nests are much more durable than the lives of the 
inhabitants require, so that solidly constructed nests are often 
found hanging to the trees long after they have been deserted, 
and are sometimes overgrown with moss. Cuming has recorded 
the fact that he found in. South America an old wasp-nest that had 
been taken possession of by swallows. "\Ve do not assign, how- 
ever, much importance to the views of de Saussure, because we 
may anticipate that enquiry will reveal much variety in the 
habits of tropical and sub-tropical wasps. It is known that 
species exist that store up honey, after the fashion of bees, and 
von Ihering has recently shown l that in Brazil, species of several 
genera form new colonies by swarming, after the manner of bees. 
So that it is possible that certain colonies may remain for a long 
period in the same nest. 

Much more variety exists in wasps' nests than would be sup- 
posed probable ; those formed by some of the tropical species of 
Vespidae are enveloped in so solid and beautifully constructed an 
envelope of papier-mache, that they resist with complete success 
the torrential rains of the tropics ; while some of those found in 
our own country are made of extremely soft and delicate paper, 

1 ZuoL ^/f~. xix. 1896, p. 449. See also note, antca, p. 70. 



II 



VESPIDAE WASPS' NESTS 



8l 



which is probably chiefly glandular products. Our British 
Yespidae number only eight species, all belonging to the one 
genus Vespa, and yet they exhibit three different modes of 
nidification. Vespa vulgaris, V. germanica and V. rvfa form 
subterranean nests, while V. arborea, V. sylvestris and V. norvcyica 
suspend their habitations from the branches of trees, bushes, or 
strong annual plants. Vespa crabro, the hornet, usually adopts 
an intermediate course, forming its nest above ground, but in a 
spot where it is protected and concealed. 
The favourite habitat of this formidable 
Insect is the interior of an old tree, but 
the hornet will sometimes avail itself of 
the protection of a thatched roof. Both 
it and other arboreal species are said, 
however, to occasionally make subter- 
ranean nests. It is ascertained that 
V. austriaca, the eighth species, is an 
inquiline. 

De Saussure, 1 the monographer of the 
social wasps, classifies them according to 
the architecture of their nests. He estab- 
lishes three groups: (1) Stelocyttares, in 
which the layers of comb are not con- 
nected with the envelope, but are sup- 
ported by pillars made by the wasps (Fig. 
31); (2) Poecilocyttares, an unsatisfac- 
tory group of which the chief character- 
istics appear to be that the nest is always 
covered by an envelope, and the comb is 

supported by an object such as the branch 

P -, i i v-i i FIG. 32. Nest of (?) PolyUa 

ot a tree, round, or on, which the envelope sp _ The enve]o pe partly 

is placed (Fig. 32) ; (3) Phragmocyttares, cut P e " ; . entrance. 

. . , . , (After de Saussure.) 

in winch the layers ot comb are supported, 

in part or entirely, by the envelope of the nest, communication 
being effected by a hole in each layer of the comb (Fig. 33). 
de Saussure's classification is far from satisfactory. There are 
many social wasps that construct nests destitute of any proper 
envelope ; as an example of this, we may mention the species of 

1 Monographic des gutpes societies, Geneva, 1853-1858, pp. cc. and 356, plates 
i.-xxxvii. 

VOL. VI G 




O 



82 HVMEXOPTERA 



CHAP. 



the abundant genus Polistes ; these Insects make hexagonal cells, 
of paper -like material, forming an irregular comb, or mass, 
attached to bushes by a stalk near its centre ; these nests are 
placed so that the mouths of the open cells look downwards. 
The species of Ischnogaster (Fig. 34) make layers of comb, con- 
nected by a pedicel, but without any envelope ; these Insects 
form a section of Stelocyttares called Gymuodomes. 

Most of the nests of the Poecilocyttares have only a single 
layer of cornb. The wasps of the genera Synoeca and Polybia 
have the habit of spreading a layer of cells on a leaf, or on the 
bark of a tree, and of covering this with an envelope that is 
pierced by a single orifice only, but that does not rest on the 
cells, and so allows circulation of the Insects between the cells 
and the envelope. This appears to be the arrangement in a 
nest of Synoeca cyanea preserved in the British Museum ; in 
this construction a large layer of cells is moulded on the branch 
of a tree, whose contour, for a length of two or three feet, it con- 
sequently follows ; while outside the mass there is placed a con- 
tinuous envelope, leaving a considerable distance between it and 
the cells. 

It would be impossible in the space at our disposal to give a 
satisfactory account of all the forms of wasp-nests, and we must 
therefore refer the student to de Saussure's work, confining 
ourselves to a brief notice of some specially interesting forms. 
The habitation of the Brazilian Polylna (Myrapetra) scutellaris 
is a very solid, closed structure, covered externally with rough 
knobs or angular projections. Although of very large size- 
it may be upwards of two feet in length it is suspended from a 
branch, and has but one orifice ; the arrangement of the combs 
in the interior is that of the Phragrnocyttares, they being 
firmly attached to the outer envelope, and so placed as to form a 
curved surface, the convexity of which is downwards : the number 
of wasps in a well-developed nest of this kind must be very great. 
This species is said to be a honey-gathering wasp. 

One of the best known of the South American wasps' nests is the 
construction (Fig. 33) of Chartergus chartarius ; these nests are so 
regularly shaped, and formed of papier-mache so compact and solid, 
as to look like stone : this edifice is attached in a very firm manner 
to the branch of a tree, and has a single portal of entry beneath ; 
its interior arrangement is much like that of Myrapetra scutellaris. 



II 



VESPIDAE WASPS' XESTS 




A very remarkable wasp's nest is preserved in the British 
Museum of Xatural History ; it is considered to be the work of 
M~"/ttez"./ni" ili inidiata Sauss. an Eumenid 
wasp ; it is a large mass of cells encircling 
the branch of a tree, which therefore pro- 
jects somewhat after the manner of an 
axle through the middle : the cells are 
very numerous, and are quite as regular 
as those of the most perfect of the combs 
of bees : the mass is covered with a very 
thick layer of paper, the nest having 
somewhat the external appearance of half 
a cocoa-nut of twice the usual size. 

Apoica pallida, a South American 
Insect, forms a nest in a somewhat similar 
manner to Polistes, but it is covered on 
its outer aspect by a beautiful paper FIG. 33. Section of nest of 

i . ,, ,1 in i Chartergus chartarius. 

skill, SO that the nest looks Somewhat South America, o. Entrance. 

like a toadstool of large size attached (After de Saussure.) 
to the branch of a tree. 

The nests of the Insects of the genus Polylna which we 
have already mentioned as located by de Saussure in his unsatis- 
factory group Poecilocyttares usually have somewhat the form 
and size of pears or apples suspended to twigs of trees or bushes ; 
these little habitations consist of masses of cells, wrapped in 
wasp-paper, in which there are one or more orifices for ingress and 
egress. Smith says that the combs in the nest of P. pygmaea 
are of the most exquisite construction, and that it i^s by no 
means an uncommon circumstance to find the outer envelope of 
the nest ornamented with patches of delicate hexagonal .tracery. 
This nest is about the size of an orange. 

We have already noticed the variety of nests formed by our 
British species of the genus Vespa ; in other parts of the world 
the edifices formed by species of Vespa attain a very large size. 
V. crabroniformis in China, and V. velutina in India, make nests 
several feet or even yards in length, inhabited by an enormous 
number of individuals ; they are apparently constructed of a 
material like brittle paper, and are arranged much like the 
nests of our British hornet, V. cralro. Vespa orientalis mixes a 
considerable quantity of earth with the paper it uses for its 



84 HYMENOPTERA 



CHAP. 



constructive efforts. In the British Museum collection there is 
a nest said to be that of the Japanese hornet, V. japonica. This 
is completely covered by a paper envelope, and has apparently" 
only a single small orifice for ingress and egress. In the same 
collection there is a nest from Bahia (believed to be that of 
a social wasp, though of what species is unknown), the outer 
wall of which is apparently formed entirely of earth, and is a 
quarter or half an inch thick : the comb inside appears also to 
be formed of clay, the whole forming an elaborate construction 
in pottery. One is tempted to believe it may prove to be the 
production of a social Eumenid. 

Habits of Social Wasps. We have already briefly noticed 
the way in which a colony of wasps is founded, but some further 
particulars as to the mode in which the society is increased and 
developed may be mentioned. The queen-wasp makes at first only 
a very small group of three or four incomplete cells ; each cell is at 
first circular, or nearly so, and moreover is of smaller diameter than 
it will afterwards be. In each of the first three or four incomplete 
cells an egg is laid, and more cells are commenced ; but as the 
eggs soon hatch and produce larvae that grow rapidly, the labours 
of the queen-wasp are chiefly directed to feeding the young. At 
first she supplies them with saccharine matter, which she pro- 
cures from flowers or fruits, but soon gives them a stronger diet 
of insect meat. This is procured by chasing living Insects of 
various kinds. Some species of wasps prefer particular kinds of 
Insects, and the hornet is said to be very fond of the honey-bee, but 
as a rule Diptera are the prey selected. When an Insect has been 
secured, the hard and innutritions parts are bitten off, and the 
succulent parts, more especially the thorax which contains chiefly 
muscular tissue, are reduced to a pulp by means of the mandibles; 
this is offered to the larvae, which are said to stretch out their 
heads to the mother to receive the food, after the manner of nest- 
ling birds. When a larva is full grown it spins a cocoon in the 
cell and changes to a pupa. It is said by some entomologists 
that the queen-wasp closes the cell for the purpose of the larval 
metamorphosis; but -this is contradicted by others, and is prob- 
ably erroneous. In about a month, or a little less, from the 
time of deposition of the egg, the perfect Insect is ready for issue, 
and almost immediately after leaving its cell it assists in the 
work that is going on for the development of the society. The 



II 



VESPIDAE SOCIAL WASPS HABITS 85 



Insects produced at this early period of the colony are exclusively 
workers, i.e. imperfect females. They relieve the queen of 
the task of supplying the larvae with food, and she henceforth 
remains within the nest, being, it is said, herself fed by her 
workers ; the society now rapidly increases in numbers, and fresh 
combs are formed, the tipper layer being always the oldest. 
About the month of August, cells of larger size than those that 
have previously been constructed are formed, and in these males 
and perfect females are produced ; in a few weeks after this the 
colony languishes and becomes extinct. When it is no longer 
possible for the enfeebled wasps to carry out their tasks of feeding 
the brood, they drag the larvae out of the cells and destroy them. 
An uncertain number of queen-wasps seek protected nooks in 
which to pass the winter, and each of these queens may be the 
founder of a nest in the ensuing spring. It should be remarked 
that de Saussure states that all the intermediate grades between 
perfect and imperfect females exist, and Marchal's recent observa- 
tions confirm this. There is in fact no line of demarcation 
between worker and queen in the wasps as there is in the honey- 
bee. Von Siebold long since drew attention to the existence of 
parthenogenesis in certain species of wasps, and it appears prob- 
able that it is of common occurrence. 

Our knowledge of the social life of European wasps has 
recently been much increased by the observations of tw T o French 
naturalists, P. Marchal and C. Janet. The latter has given 
an elaborate history of a nest of the hornet, showing the rate 
and variations of increase in numbers. His observations on this 
and other species indicate that warmth is of the utmost im- 
portance to wasps ; the Insects themselves create a consider- 
able amount of heat, so that the temperature of their abodes is 
much greater than that of the air. He considers that in Europe 
an elevated temperature is essential for the development of the 
individual, 1 and that the chief object of the various wrappers of 
paper with which the Insects surround their nests is to keep 
up this high temperature. These envelopes give a great deal 
of trouble to the Insects, for they have to be repeatedly 

1 Hence probably the great difference in the abundance of wasps in different 
years : if a period of cold weather occur during the early stages of formation of a 
wasp family, operations are suspended and growth delayed ; or death may even 
put an end to the nascent colony. 



86 HYMEXOPTERA 



CHAT- 



destroyed and reformed, as the combs they contain increase in 
size. Marchal's observations l relate chiefly to the production of 
the sexes and worker-forms, in the subterranean species, J~cspa 
germanica and I", rulgaris. The layers of comb include cells of 
two sizes. The upper layers, which are the first formed, consist 
of small cells only : the lower combs are constructed (at Paris) 
early in August, and consist of larger cells from which males 
and large females are reared. The males are, however, reared 
also in large numbers in the small cells. If the queen be re- 
moved, the workers become fertile, and produce paithenogenetically 
many eggs, but all of the male sex. He entertains no doubt 
that even when the queen is in full vigour the workers produce 
males if there is an abundant food supply. 

The social wasps at present known number 500 or 600 species. 
Pollutes is a very extensive genus, and it has also a very wide 
geographical distribution : some of the species and those found in 
widely-distant parts of the world are remarkable on account of 
their excessive variation in colour, and it is worthy of note that the 
extreme forms have been more than once taken from the same nest. 

Xext to Polistes, l^espa is the most numerous in species, 
about 150 being known, and it is to this genus that all our 
British social wasps belong. Xo Insects are .better known 
in our islands than these wasps, owing to the great numbers 
of individuals that occur in certain seasons, as well as to 
their frequently entering our habitations and partaking of our 
food, and to the terror that is occasioned by their supposed 
ferocity and desire to sting. This last feature is a complete 
mistake ; wasps never sting unless they are roused to do so by 
attacks, or by considerable interference with their work. The 
only real danger arises from the fact that a wasp may be occa- 
sionally taken into the mouth with fruit, or may be handled 
unawares. "\Vhen they are flying about they are perfectly 
harmless unless attacked or irritated, and even if they settle 
on the person no danger of their stinging exists unless move- 
ment is made. Sichel correctly states that a person may 
station himself close to a wasp's nest and 'remain there without 
any risk at all, provided that he makes no movement ; indeed, it 
is more than probable that if no movement, or if only gentle 

1 CE. Ac. Paris, cxvii. 1S93, p. 584 ; op. cit. cxxi. 1895, p. 731 ; Arch. Zool. 
exper. (3) iv. 1896, pp. 1-100. 



II 



VESPIDAE SOCIAL WASPS 



movement, be made, the wasps are unaware of the presence of an 
intruder. It is, however, well ascertained that if they are 
molested at their work, more especially when they are actually 
engaged in the duties of the nest, they are then extremely vin- 
dictive, and follow for a considerable distance those who have 
irritated them. The East Indian V. velutina is specially fierce 
when aroused, and is said by Home to have followed a party 



/* 



r-< vj-:^- ' 

- 




FK;. 34. Ischnoyaster mdlyi. Java. A, Female imago (the line at the side shows its 
length) ; B, nest, C, maxilla ; D, labiuni ; E. mandible (tip downwards). The nest 
is probably upside down, although shown here as by de Saussure. 

through dense jungle for miles, and on some occasions to have 
stung animals, and even human beings, to death. 

This vindictiveness is, however, only an exceptional mood due 
to some interference with the colony. Even the hornet, not- 
withstanding its threatening appearance, is harmless unL 
unduly provoked ; its nests and their inhabitants can be kept in 
domesticity, exhibited to strangers, even moved from place to 
place, yet the hornets will not take offence if due gentleness be 
observe 1. It is said that wasps will rear the progeny of a neigh- 
bour in circumstances where this assistance is necessurv. Hess 

w 

has related a case in which a queen-hornet had commenced a nest, 
and was killed by an accident, leaving young brood in the comb 



88 



HYMENOPTERA 



CHAP. 



unprovided for : as a result many of the helpless grubs died, and 
others were in a state of starvation, when a strange queen-hornet 
appeared, associated itself with the comb, and, adopting the orphan 
brood, nourished them and brought them to their full size. 

We have already alluded to the fact that, so far as external 
structure is concerned, there is no great difference between the 
social and the solitary wasps. Both, too, run through analogous 
series of forms and colours, and the genus Isclmoy aster (Fig. 34) 
seems to connect the two groups by both its structure and mode 
of life. The social habits are in many species only inferred, and 
with greater knowledge will probably prove fallacious as a guide 
to classification ; indeed we have already said that in the genus 
Vespa perhaps the most perfectly social of all the wasps there 

is one species that has no worker, 
and that lives, it is supposed, as 
a parasite, in the nests of its 
congeners. For this species, V. 
austriaca, it has been proposed to 
create a separate genus, Pseudo- 
vespa, on account of this peculiarity 
of habit, although no structural 
character has been detected that 
could distinguish it. De Saussure 
has stated his conviction that 
workers do not exist in some of 
the exotic genera, so that it appears 
highly probable that with the pro- 
gress of knowledge the present divi- 
sion between social and solitary 
wasps will prove untenable. 

Remains of Insects referred to 
the genera Polistes and Vcspa 
have been found in tertiary strata 




FIG. 35. Masaris vespiformis. A, male 
B, female. Egypt. (Ar'ter Schaum.) 



in various parts of Europe and in North America. 

Fam. 3. Masaridae. 

Anterior winy with two complete sub-marginal cells. Antennae 
tixindly incrassate or clubbed at the extremity. Claws dis- 
tinctly or oljsoletely dentate. 

This is a group of fifty or sixty species with but few genera, 



MASARIDAE 



8 9 



and most of its components appear to. be Insects of the greatest 
rarity. In their appearance the Insects of this Family differ con- 
siderably from the other Diploptera, and as the wings are only 
imperfectly, or not at all, plicate, it must be admitted that the 
systematic affinities of the group require reconsideration. The 
pronotal structure is, however, completely that of Diploptera. 
The typical form of the Family, Masaris vespiformis, though 
described a hundred years since, is a species of such ex- 
treme rarity, and its sexes are so different, that entomologists 
have only recently been able to agree about it. It has been 
found in Egypt and Algeria. The genera Ceramius, Jugurthia, 
< t >"<trtenia and Coelonites are also members of the Mediterranean 
fauna, while Paragia is Australian, and Trimeria South American. 
Several species of the genus Masaris inhabit Xorth America, and 
Cresson has recently described another 
Masarid genus from the same country, 
under the name of Et'j>i/i'i/i/i</. 

The little that is known of their 
natural history is almost limited to an 
account given by Giraud of the habits of 
Ceramics lusitanicus, of which species he 
found a colony near Briangon. The Insect 
makes nests in the earth ; they are entered 
by means of a chimney - like passage 
analogous to what is formed by certain 
Odynerus ; the gallery when completed is 
about six centimetres long, and at its 
extremity is an earthen cell in which the 
larva lives ; this is fed by the mother, 
who brings to it from time to time a 
supply of a paste, described as being 
somewhat like dried honey. The growth 
of the larva is believed to be rapid. 

Some fragmentary observations made 
by Lichtenstein on Coelonites abbreviatus have also been recorded. 
This species, near Montpellier, constructs earthen cells ; they are 
not, however, subterranean, but are placed side by side on the dry 
stems of plants (Fig. 36); these cells are stored with a material 
similar to that supplied by Ceramius lusitanicus to its young. 




FIG. 36. Cells constructed 
by Coelonites abbreviatus. 
(After Andre.) 



CHAPTER III 

HYMENOPTERA ACULEATA COXTIXUE1) DIVISION III. FOSSORES OR 

FOSSORIAL SOLITARY WASPS FAMILY SCOLIIDAE OR SUBTER- 
RANEAN FOSSORS FAMILY POMPILIDAE OR RUNNERS FAMILY 

SPHEGIDAE OR PERFECT-STINGERS 

Division III. Fossores. 

Aculeate Hymenoptera, in which the abdomen, thouyh very diverse 
inform, does not bear prominences on the upper aspect of the 
basal segments ; front iving without longitudinal fold along 
the middle ; hairs of body nut plumose. Only two forum 
(male and female) of each specie*. 

FOSSORIAL Hymenoptera are distinguished from other Aculeates 
at present only by negative characters, i.e. they are Aculeates, 
but are not ants, bees or wasps. According to their habits they 
fall into four, by no means sharply distinguished, groups (1) 
those that form no special receptacles for their young, but are 
either of parasitic or sub-parasitic habits, or take advantage of 
the abodes of other Insects, holes, etc. ; (2) constructors of cells 
of clay formed into pottery by the saliva of the Insect, and by 
drying ; (3) excavators of burrows in the ground ; (4) makers of 
tunnels in wood or stems of plants. Several species make use 
of both of the last two methods. The habits are carnivorous ; 
the structures formed are not for the benefit of the makers, but 
are constructed and stored with food for the next generation. 
Their remarkable habits attracted some attention even 2000 
years or more ago, and were to some extent observed by Aristotle. 
The great variety in the habits of the species, the extreme 
industry, skill, and self-denial they display in carrying out their 
voluntary labours, render them one of the most instructive groups 
of the animal kingdom. There are no social or gregarious 



CHAP. Ill 



FOSSORIAL SOLITARY WASPS 



forms, they are true individualists, and their lives and instincts 
offer many subjects for reflection. Unlike the social Insects they 
can learn nothing whatever from either example or precept. 
The skill of each individual is prompted by no imitation. The 
life is short, the later stages of the individual life are totally 
different from the earlier : the individuals of one generation 
only in rare cases see even the commencement of the life of the 
next ; the progeny, for the benefit of which they labour with 




FIG. 37. Scelifthron nigripes 9 (Sub-Fam. Sphegides). Amazons, x \. 

unsurpassable skill and industry, being unknown to them. Were 
such a solicitude displayed by ourselves we should connect it 
with a high sense of duty, and poets and moralists would vie in 
its laudation. But having dubbed ourselves the higher animals, 
we ascribe the eagerness of the solitary wasp to impulse or 
instinct, and we exterminate their numerous species from the face 
of the earth for ever, without even seeking to make a prior ac- 
quaintance with them. Meanwhile our economists and moralists 
devote their volumes to admiration of the progress of the civilisa- 
tion that effects this destruction and tolerates this negligence. 



92 HYMENOPTERA CHAP. 

It should be noted that in the solitary as in the social Insects 
the males take no part whatever in these industrial occupations,- 
and apparently are even unaware of them. It is remarkable 
that, notwithstanding this, the sexual differences are in the 
majority less than is usual in Insects. It is true that the various 
forms of Scoliidae exhibit sexual distinctions which, in the case 
of Thynnides and Mutillid.es are carried to an extreme degree, 
but these are precisely the forms in which skill and ingenuity are 
comparatively absent, the habits being rather of the parasitic 
than of the industrial kind, while the structure is what is 
usually called degraded (i.e. wingless). The great difference 
between the habits of the sexes, coupled with the fact that there 
is little or no difference in their appearance, has given rise to a 
curious Chinese tradition with regard to these Insects, dating 
back to Confucius at least. 1 The habit of stinging and storing 
caterpillars in a cell, from which a fly similar to itself afterwards 
proceeds having been noticed, it was supposed to be the male that 
performed these operations ; and that when burying the cater- 
pillars he addressed to them a spell, the burden of which is 
' mimic me." In obedience the caterpillars produce the wasp, 
which is called to this day " Jiga," that is in English " mimic 
me." The idea was probably to the effect that the male, not 
being able to produce eggs, used charmed caterpillars to continue 
the species. 

Summary of the Prey of Fossores. 

Group of Fossores. F< M id or Occurrence. 

Fam. Scoliidae. 

Sub-Fain. Mutillides . . As parasites on Hymenoptera Aculeata. 
., Thynnides . . (?) Parasites on Lepidopterous pupae. 
., Scoliides . . . Larvae of Coleoptera [(?) spiders in the case of 

Elis 4-ofr/fr(]. 

., Rhopalosomides. Unknown. 
Sapygides . . The provisions stored by bees. Caterpillars 

(teste Smith). 

p p .,. , (Spiders. Rarely Orthoptera (Gryllidae and 

{ Blattidae, teste Bingham) or Coleoptera. 
/ Orthoptera (especially Locustidae), larvae 

Fain. Sphegidae. ) of Lepidoptera, Spiders [(?) same species 

Sub-Fain. Sphegides . . \ (Sceliphron madraspatanum and Sphex 

), both spiders and caterpillars]. 



Kumagusu Minakata, in Nature, 1. 1894, p. 30. 



Ill 



FOSSORIAL WASPS 



93 



Group of FossoreG. 

Fam. Sphegidae. 

Sub-Fam. Ampulicides . 
Larrides . 



Trypoxylonides , 
Astatides . 



Food or Occurrence. 

Orthoptera (Blattidae only). 

Orthoptera of various divisions. Aculeate 

Hymenoptera, in the case of Palarus. 

[Spiders stolen from nests of Pelopaeus by 

Larrada.] 

Spiders, caterpillars, Aphidae. 
Astata loops uses Pentatomid bugs, cockroaches, 

and even Aculeate Hymeiioptera (Oxylelus, 

teste Smith). 
Diptera and Cicada. 
Diptera, Homoptera (Gorytes mystaceus takes 

Aplirophora out of its " cuckoo-spit "). 
Aculeate Hymenoptera (Philantlius). Hard 

beetles, viz. Curculionidae, Buprestidae, 

Chrysomelidae (Cerceris). 
Small Homoptera, even Aphidae. Diptera 

(Tipulidae) in Hawaii. 
Diptera, Aphidae [? the same species of wasps 

both of these]. Other small Homoptera. 

Ants (in the case of Fertonius). Parasitic 

optera (in the case of Lindenius). 

Great diversity of opinion exists as to the classification of the 
Fossores. This arises chiefly from the incomplete state of the 
collections studied, and from the fact that the larger part of the 
works published are limited to local faunae. Opinions as to the 
families vary ; some admitting only three or four, others upwards 
of twenty. After consideration of the various views, the writer 
thinks it best to admit at present only three families, which 
speaking broadly, correspond with habits, viz. (1) Scoliidae, 
subterranean stingers; (2) Pompilidae, runners; (3) Sphegidae, 
stingers above ground. 



Bembecides 
Nyssonides . 

Philanthides 



Mimesides 



Crabronides 



1. Scoliidae. Pronotum and tegulae in contact. Abdomen with the 

plane of the ventral surface interrupted by a chink between the 
first and second segments. Numerous wingless forms. 

2. Pompilidae. Pronotum and tegulae in contact. Abdomen with the 

plane of the ventral surface not interrupted by a chink. Legs 
very long. No wingless forms. 

3. Sphegidae. Pronotum and tegulae not in contact. No wingless 

forms. 



We shall treat as sub-families those divisions of Scoliidae and 
Sphegidae considered by many as families. 



94 HYMENOPTERA CHAP. 

Fam. 1. Scoiiidae. 

The members of this family, so far as is known, display less 
perfect instincts than the Sphegidae and Pompilidae, and do not 
construct cells or form burrows. Information as to the habits is 
almost confined to European forms. AVe adopt five sub-families. 

Sub-Fam. 1. Mutillides.--77/r stiles of the pronotum, reach the 
ti'ijiilae : the female is destitute of wings and ocelli, 
frequently h.aviny I In 1 parts of f/ir thorax so closely soldered 
that the divisions between them are obliterated : the males are 
winged, furnished with ocelli, and having the thoracic divisions 
distinct ; intermediate tibiae with tiuo apical spurs. Front 
i ring with, two or three suh-i>mryi iml fells. The larvae live 
parasitically at the expense, of other Hymenoptera Aculeata. 

The Mutillides have some resemblance to ants, though, as 
they are usually covered with hair, and there is never any node 
at the base of the abdomen, they are readily distinguished from 
the Formicidae. The great difference between the sexes is their 
most striking character. Their system of coloration is often 
very remarkable, the velvet-like pubescence clothing their bodies 
being variegated with patches of sharply contrasted vivid colour : 
in other cases the contrast of colour is due to bare, ivory-like 
spaces. They have the faculty of stridulating, the position and 
nature of the organ for the purpose being the same ;is in ants. 

Very little exact information exists as to the habits and life- 
histories of the species. Christ and Drewsen, forty or fifty years 
ago, recorded that M. euro'paea lives in the nests of bees of the 
genus Jiombus, and Hoffer has since made some observations on the 
natural history of the same species in South East Europe, where 
this Mitt ill a is found in the nests of ten or eleven species of 
Bombus, being most abundant in those of B. ayrorum, and B. 
variabilis ; occasionally more individuals of Mnfilln than of bees 
may be found in a nest. He supposes that the egg of the 
Ma tHlu, is placed in the young larva of the Bombux, and hatches 
in about three days; the larva feeds inside the bee-larva, and 
when growth is completed a cocoon is spun in the interior of 
the pupa -case of the bee. When the perfect Insects emerge, 
the males leave the nest very speedily, but the females remain 
for some time feeding on the bees' honey. Females are usually 
produced in greater numbers than males. This account leaves 



Ill 



SCOLIIDAE MUTILLIDES 



95 



much to be desired. From the observations of Radoszkowsky 
it is clear that other species of Mutillides are by no means 
confined to the nests of Bovibus but live at the expense of 
Aculeate Hymenoptera of various groups. This naturalist asserts 
that the basal abdominal segment of the parasite resembles in 
form that of the species on which it preys. 

The apterous condition of the females of Mutillides and 
Thynnides is very anomalous in the Fossors ; this sex being in 
the other families distinguished for activity and intelligence. 
The difference between the sexes is also highly remarkable. The 
males differ from the females by the possession of wings and by the 
structural characters we have mentioned, and also in a most striking 




8 



FIG. 38. Mutilla stridula. Europe. A, Male ; B, female. 

manner in both colour and form ; Burmeister, indeed, says that in 
South America the metropolis of Mutillides there is not a single 
species in which the males and females are alike in appearance ; this 
difference becomes in some cases so extreme that the two sexes of 
one species have been described as Insects of different families. 

Upwards of one thousand species are assigned to the genus 
Mutilla, which is distributed over the larger part of the world ; 
there is so much difference in these species as to the nervuration 
of the wings in the males, that several genera would be formed 
for them were it not that no corresponding distinctions can be 
detected in the females. Three or four species of Mutilla are 
described as being apterous in the male as well as in the 
ft -male sex ; they are very rare, and little is known about them. 
Only three species of Mutillides occur in Britain, and they are 
but rarely seen, except by those who are acquainted with their 



Q6 



HYMENOPTERA 



CHAP. 



habits. The African and East Indian genus, Apterogyna, includes 
some extremely peculiar Hymenoptera ; the males have the wing 
nervuratioii very much reduced, and the females are very ant-like. 
owing to the deep constriction behind the first abdominal ring. 

Sub-Fam. 2. Thynnides. Males and females very different in 

form; the male winged, the front winy with three, or on/// 

two, sub-marginal cells ; the female wingless and with ///> 

thorax divided into three sub-equal 




FIG. 39. Methoca ichneumonides. A, Male ; B, female. 

Britain. 



The Thynnides are by some entomologists not separated from 

theMutillides; but 
the distinction in 
the structure of 
the thorax of the 
females is very 
striking. In the 
Thynnides the 
nervuratioii of the 
wing appears 
always to extend 
to the outer 
margin, and in the 
Muti Hides not to 
do so. This family 
is represented in 
Britain by a single very rare Insect, Methoca ichneumonides : to 
the unskilled observer the female would appear to be without 
doubt an ant. This Insect is by some considered as the type 
of a family distinct from the Thynnides proper. Thynnides are 
numerous in Australia. Very little is really known as to their 
habits, though it has been stated that they are parasitic on 
Lepidoptera, Bakewell having obtained specimens from sub- 
terranean cocoons of that Order. Those who are interested in 
differences between the sexes of one species should examine the 
extraordinary examples of that phenomenon presented by the 
Thynnides ; the dissimilarity throughout the group which is 
now of considerable extent being so extreme that no ento- 
mologist would from simple inspection believe the two sexes 
to have any connection ; but the fact that they are so con- 
nected has been demonstrated beyond doubt. In very few 



Ill 



SCOLIIDAE THYNNIDES SCOLIIDES 



97 



cases, however, have the sexes been matched, so that at present 
males are no doubt standing in the lists of Hymenoptera as one 
species and their females as other species. 

Sub-Fam. 3. Scoliides. Pronotum reaching back to the tegulae ; 
legs stout; intermediate tibiae until one apical spur; both 
sexes winged ; the nervures not extending to the posterior 
(i.e. distal} margin. 

This group includes some of the largest and most powerful of 
the Aculeate Hymenoptera. Its members are usually hairy 
Insects with thick legs, 
the colour being black, 
more or less variegated 
with bands or spots of 
red or yellow ; the hind 
body is elongate, has 
only a very short pedi- 
cel, and in the male 
is usually termin- 
ated by three project- 
ing spines. The pro- 
notum is of variable 
dimensions, but its front 
angles are always co- 
adapted with the points of insertion of the front wings. The 
nervuratioii of the front wings is confined to the basal part, the 
extensive apical or outer area possessing no nervures. There is 
frequently a great difference in the size of the two sexes of the same 
species, the female being very much larger than the other sex. The 
larvae, so far as is known, devour those of Lamellicorn Coleoptera. 

Fabre has investigated the habits of some of the species of 
Scoliides found in France, and has informed us that their means 
of subsistence consists of larvae of the larger Lamellicorn beetles, 
Cetonia, Oryctes, Anoxia, and Euclilora ; these beetles belong to 
very different divisions of the Lamellicornia, but they have in 
common the fact that their larvae are of subterranean habits, 
living in the earth or in accumulations of debris in which there 
is a large proportion of vegetable matter or roots. The female 
Scolia penetrates into the ground in order to find the Lamellicorn 
larvae necessary as food for its progeny. Scolia bifasciata 

VOL. VI H 




FIG. 40. Scolia haemorohoidalis 9- Europe. 



98 HYMENOPTERA CHAP. 

attacks the larvae of several species of Cetonia, and S. (Colpa) 
interrupta chooses the larvae of the chafers Anoxia villosa and 
A. matutinalis. The mother Scolia enters the ground in August 
or September, and having found a suitable larva stings it and 
deposits an egg on the ventral surface of the prey ; the paralysed 
larva is left where it was found, no attempt being made to place 
it in a special receptacle. The egg is placed on the ventral 
surface, well behind the feet, under a mass of matter in the 
alimentary canal. Shortly after being hatched the young- 
destroyer penetrates with its head the skin of the victim, and in 
this position commences to feed ; it is necessary that it should 
obtain its food without killing the Cetonia larva, for it cannot 
prosper on decaying food, so that if the Cetonia larva die the 
Scolia larva likewise perishes ; the latter, accordingly, does 
not withdraw its head from the interior of the victim, but 
remains always in the same position, as it grows larger extending 
its head forwards into the front part of the interior of its victim ; 
the internal organs of the latter are consumed in a systematic 
order so as to delay bringing about its death till the last moment, 
and thus all the interior of the Cetonia larva is appropriated till 
nothing remains but an empty skin. By a series of experiments, 
Fabre showed how essential it is that this apparently revolting 
operation should be carried on with all details strictly en regie. 
If the head of the Scolia larva be taken out from the victim and 
applied to another part of the body of the Cetonia, the result is 
that it cannot eat ; even if it be replaced in the original situa- 
tion, after being taken away, it frequently happens that the 
Cetonia larva dies, its death involving also that of the destroyer. 
It is necessary, too, that the victim should be paralysed, for if an 
intact Cetonia larva be taken and bound down in such a' position 
that it cannot move, and if a small orifice in its skin be made in 
the proper spot and a young Srolia larva be placed on it, the 
little parasite will avail itself of the opportunity and commence 
to feed on the larva provided for it, but the latter will speedily 
die, and the Scolia necessarily perishes with it. Thus both the 
paralysis of the victim and the special mode of eating are essential 
to the life of the Scolia. The operation of stinging the larva, so 
as to produce the necessary paralysis, or rather insensibility, is a 
difficult one, and requires great skill and patience. The Cetonia 
larva is of large size, and must be pierced in one particular spot ; 



in SCOLIIDAE SCOLIIDES SAPYGIDES 99 

in order to reach this the Scolia mounts on its victim, and is 
frequently dislodged by its struggles ; sooner or later, however, 
the proper position is obtained by the wasp, and the larva is 
then stung in the exact spot necessary to allow the sting (and the 
poison introduced by it) to reach the most important of the 
nervous ganglia that control the movements of the body, this spot 
being, in the case of the Cetonia, the line of demarcation between 
the pro- and meso-thorax, on the middle line of the ventral 
surface of the body. The Scolia gives but one sting to the 
victim, and this it will not administer until it can do so exactly 
in the proper place. This practice of devouring the victim 
slowly, without killing it till all is eaten, is very widely spread in 
the Hymenoptera, and it is satisfactory to find that we may infer 
from Fabre's observations that it is not so horrible as it 
would at first appear ; for it is probable that the stinging 
prevents decomposition of the victim, not by reason, as some have 
supposed, of the poison injected by the wasp having an antiseptic 
effect, but rather by means of destroying sensibility, so that the 
creature does not die from the pain, as it is believed it did in 
certain cases where Fabre induced the young Scolia larva to feed 
on a victim that had not been stung. We may here remark 
that very little exact information exists as to the operation of 
stinging. Fabre attaches great importance to the sting- being 
inflicted on a nerve -ganglion. Whether a sting that did not 
reach this part might not have a sufficient effect appears, how- 
ever, doubtful. 1 

A remarkable form of Scoliides, with wings of smaller size 
than usual and deeply divided, has been described by Saunders 
under the name Pseudomeria graeca. Still more remarkable is 
Komarovia victoriosa found in Central Asia ; in this Insect the 
male retains the appearance of a slender, pallid Scolia, but the 
female differs totally in form, and has the peculiar wings so re- 
duced in size as to be useless for flight. 

Sub-Fam. 4. Sapygldes. Closely allied to the Scoliides, but j>os- 
sessing slender legs and antennae ; also the first abdominal 
segment is less disconnected from the second, so that the outline 

1 As this work is passing through the press we receive a book by Mr. and Mrs. 
Peckhani on The Instincts and Habits of the Solitary Wasps, Madison, 1898. 
They are of opinion that, in the case of some species, it does not matter much 
whether the victim is or is not killed by the stinging. 



100 



HYMENOPTERA 



CHAT. 




FIG. 41. $cij>i/yn -l-j/i'/tctata ?, 
Britain. 



is less interrupted ; the eyes are deeply emarginate ; the hind 
body is not spinose at the apex. 

The economy of Sapygn, the only genus, has been the subject 

of difference of opinion. The views 
of Latreille and others that these 
species are parasitic upon bees is 
confirmed by the observations of 
Fabre, from which it appears that 
S. 5 -punctata lives in the burrows 
of species of the bee-genus Osmia, 
consuming the store of provisions, 
consisting of honey-paste, that the 
bee has laid up for its young. Ac- 
cording to the same distinguished 
observer, the Sapyga larva exhibits 

h\ -pel-metamorphosis (i.e. two consecutive forms), and in its young 

state destroys the egg of the bee ; but his observations on this point 

are incomplete and need 

repetition. AVe have two 

species of Sapyga in 

Britain ; they differ in 

, * 

colour, and the sexes of 
S. 5 -punctata also differ 
in this respect ; the 
a bdomen , spot t e d with 
white in both sexes, is 
in the female variegate 
with red. Smith found 
our British Sapyga o- 
punctata carrying cater- 
pillars. 



"CR- 




Sub-Fam. 5. Rhopalo- 
somides. --Antennae 

elongate, spinigerous ; FIG. 42. Rhopalosoma poeyi. A, female imago ; 

ocelli very prominent ; B < IVoilt of " Cuba " (After Westwood -) 
tarsi of peculiar structure, their claws bifid. 

This sub-family has recently been proposed by Ashmead J for 

1 P. cut. Soc. Washington, iii. 1896, p. 303. 



in FOSSORES 101 

an extremely rare American Insect that had previously been placed 
by Cresson among parasitic Hymenoptera. Westwood classed 
RJiopalosoma among Diploptera, saying of it " animal quoad 
affmitates excrucians." We reproduce Westwood's figure, but not 
being acquainted with the Insect we can express no opinion as to 
whether it is allied to the Scoliidae or to the Sphegidae. The 
habits are, we believe, quite unknown. 

Fam. 2. Pompilidae. 

Pronotum at the sides reaching the tegulae ; hind body never 
definitely pedicellate, though the first segment is sometimes 
elongate and conical; hind legs long ; eyes elliptic inform, 
not emarginate. 

The Pompilidae are perhaps the most extensive and important 
of the groups of Fossores, and are distributed over all the lands of 
the globe, with the exception of some islands and of the inclement 
arctic regions. The sting of the Pompilidae, unlike that of most 
of the Fossores, inflicts a burning and painful wound; the creatures 
sometimes attain a length of two or three inches, and a sting from 
one of these giants may have serious results. Although there is 
considerable variety in the external form of the members of the 
group, the characters given above will enable a Pompilid to be 
recognised with approximate certainty. The elongation of the 
hind legs includes all the parts, so that while the femur extends 
nearly as far back as the extremity of the body in dried 
examples at any rate the tibiae and the long tarsi extend far 
beyond it ; thus these Insects have great powers of running ; they 
are indeed remarkable for extreme activity and vivacity. They 
may frequently be seen running rapidly on the surface of the 
ground, with quivering wings and vibrating antennae, and 
are probably then employed in the search for prey, or some other 
of the operations connected with providing a store of food for 
their young. Spiders appear to be their special, if not their only, 
prey. Several authors have recorded details as to the various 
ways in which the prey is attacked. Fabre has observed the 
habits of several species, and we select his account of the modus 
operandi of species of the genera Pompilus and Calicurgus, in 
their attacks on poisonous spiders that inhabit holes in the 
ground or in walls. The wasp goes to the mouth of the spider's 
burrow, and the latter then dashes to the entry, apparently 



102 



HYMENOPTERA 



CHAP. 



enraged at the audacity of its persecutor. The Calicurgus will 
not actually enter a burrow when there is a spider in it, 
because if it did so the spider would speedily dispose of the 
aggressor by the aid of its poisonous fangs. The Calicurgus, 
therefore, has recourse to strategy with the object of getting 
the spider out of its nest ; the wasp seizes its redoubtable 

foe by one foot and pulls ; 
probably it fails to extract 
the spider, and in that case 
rapidly passes to another 
burrow to repeat its tactics ; 
sooner or later a spider is 
in some moment of inatten- 
tion or incapacity dragged 
from its stronghold, and, 
being then comparatively 
helpless, feels itself at a 
disadvantage and offers but a 
feeble resistance to the wasp, 
which now pounces on its 
body and immediately in- 
flicts a sting between the 
fangs of the foe, and thus 
at once paralyses these dangerous weapons ; thereafter it stings 
the body of the spider near to the junction of the abdomen and 
cephalothorax, and so produces complete inactivity. Having 
secured its prey, the wasp then seeks a suitable hole in which 
to deposit it ; probably an empty burrow of a spider is selected 
for the purpose, and it ma.y be at a height of several feet in a 
wall ; the Hymenopteron, walking backwards, drags its heavy 
prey up the wall to bring it to the den. When this is accom- 
plished an egg is deposited on the spider, and the wasp goes 
in search of a fragment or two of mortar, with winch the mouth 
of the burrow is finally blocked. Fabre's accounts refer to the 
habits of several species, and give a good insight into some points 
of the instincts of both the spider and the wasp. It seems that a 
sense of superiority is produced in one or other of the foes, accord- 
ing as it feels itself in suitable conditions ; so that though a spider 
out of its burrow and on the ground is speedily vanquished by the 
1'oiupilid, yet if the two be confined together in a vase, both are 




FIG. 43. Calicurgus hyalinatus ?. Britain. 



ill POMPILIDAE 103 

shy and inclined to adopt defensive or even evasive tactics, the 
result probably being that the wasp will be killed by the spider 
during the night, that being the period in which the attacking 
powers of the spider are more usually brought into play. 

It seems to be the habit of some Pompilus to procure a victim 
before they have secured a place for its reception ; and Fabre took 
advantage of this fact, and made very interesting observations on 
some points of the instinct of these wasps. Having found a 
Pompilus that, after having caught a spider and paralysed it, 
was engaged in making a retreat for its reception, he abstracted 
the booty, which was deposited at the top of a small tuft of 
vegetation near to where the Pompilus was at work. In this 
case the burrow in course of preparation was subterranean, and 
was formed by the Pompilus itself, which therefore could not, 
while it was engaged underground, see what took place near it. 
It is the habit of the wasp to leave its work of excavation from 
time to time, and to visit the prey as if to assure itself of the 
safety of this object, and to enjoy the satisfaction of touching it 
with the mouth and palping it. Desirous of testing the wasp's 
memory of locality, Fabre took the opportunity, while the Insect 
was working at the formation of its burrow, of removing, as we 
have said, the booty from the place where it had been deposited, 
and putting it in another spot some half-yard off. In a short 
time the Pompilus suspended work and went straight to the spot 
where it had deposited its property, and finding this absent, 
entered on a series of marches, counter-marches, and circles round 
the spot where it had left the prey, as if quite sure that 
this was really the place where the desired object ought to be. 
At last convinced that the paralysed prey was no longer where 
it had been placed, the Pompilus made investigations at a greater 
distance and soon discovered the spider. Fabre recounts that its 
movements then appeared to indicate astonishment at the change 
of position that it thus ascertained to have occurred. The wasp, 
however, soon satisfied itself that this was really the very 
object it was seeking, and seizing the spider by the leg slightly 
altered its position by placing it on the summit of a small tuft 
of vegetation ; this latter proceeding being apparently always 
carried out by this species of Pompilus. Then it returned to its 
excavation, and Fabre again removed the spider to a third spot ; 
the wasp when it next rested from its work made its way 



104 HYMENOPTERA CHAP. 

immediately to the second spot, where it had last left the spider, 
thus showing that it possessed an accurate memory for locality ; 
the wasp was very much surprised at the absence of the valued 
prize and persisted in seeking it in the immediate vicinity with- 
out once returning to the place where it had been first located. 
Fabre repeated this manoeuvre five times, and the Pompilus 
invariably returned at once to the spot where it had last left its 
prey. The acute memory for localities displayed by this Insect 
seems to be more or less general throughout the Aculeate 
Hymenoptera, and is of very great importance to them. The 
power of finding the object appears to depend on sight, for when 
Fabre, after removing the spider to a fresh spot, made a slight 
depression in the ground, placed the spider in it and covered it 
over with a leaf, the wasp did not find it. At the same time, the 
Insect's sight must be a very different sense from our own, for the 
wasp, when seeking its lost booty, frequently passed within a couple 
of inches of it without perceiving it, though it was not concealed. 

Belt gives an example of the habits of the Mexican Pompilus 
2^olistoides. He noticed it, when hunting for spiders, make a dart 
at a web in the centre of which a spider was stationed ; by this 
movement the creature was frightened and fell to the ground, 
where it was seized by the wasp and stung. The Pompilus then 
dragged its prisoner up a tree and afterwards Hew off with it, 
the burden being probably too heavy for conveyance to the nest 
without the vantage of an elevation to start from. 

Several modifications adopted by Pompilidae in their mode of 
stinging their spider- victims have been recorded by Ferton ; these 
we cannot allude to in detail, but will nevertheless mention that 
one species stings the body of its spider-prey at random, and 
that in other cases it would appear that the paralysis of the 
spider is evanescent. In short, there are various degrees of 
perfection in the details of the art of stinging. 

The most remarkable of the forms of Pompilidae are the 
numerous species of Pe^)sis, a genus peculiar to America, whence 
upwards of 200 species are already known. 1 Some of them 
;iltain a length of two inches or more, and are able to conquer 
the largest spiders ; even the formidable Mygale avicularis 
succumbs to their agility and skill. Some of these Pepsis have 
beautifully coloured wings ; according to Cameron, this may be 

1 Monograph by Lucas, Berlin ent. Zcitsclir. xxxix. 1894. 



in POMPILIDAE IO5 

due to scales. P. formosus, Say, is called in Texas the tarantula- 
killer ; according to Buckley, its mode of attack on the huge 
spider is different from that made use of by its European ally. 
When it discovers a tarantula it flies " in circles in the air, 
around its victim. The spider, as if knowing its fate, stands up 
and makes a show of fighting, but the resistance is very feeble 
and of no avail. The spider's foe soon discovers a favourable 
moment and darts upon the tarantula, whom it wounds with its 
sting, and again commences flying in circles." The natural 
retreat of this huge spider, Mygale hentzii, is in holes in the 
ground, and this account does not inform us whether the spider 
allows itself to be overcome when in its nest, or is only attacked 
when out of its retreat. 

The genus Mygnimia includes a very large number of species, 
and has a wider geographical distribution than Pcpsis, being 
found in the tropical regions of both the Old and New Worlds, 
some of them rivalling in size and ferocity the larger specimens 
of the genus Pcpsis. In the Insects of this genus there is usually 
a more or less distinct small space of more pallid colour on the 
middle of each front wing. Parcqiompilus is a curious genus 
consisting of Insects of a great variety of peculiar coloration, 
and having the wings short, so as to be of little vise for flight. 
P. gravesii is an inhabitant of Chili. 

Agenia carbonaria and A. liyalipennis are small and feeble 
Insects inhabiting the south of Europe. A. carbonaria extends to 
the south of England. They construct, as nests for their offspring, 
small earthenware vessels, differing in form according to the 
species, those of A. liycdipennis being vase-like in shape, while 
those of A. carbonaria are contracted near the mouth, something 
after the fashion of a wide-mouthed bottle. The Insect is able 
by some means Fabre thinks by the use of saliva to varnish 
the interior of the vessel so that it will not absorb water ; the 
outside of the cells is, however, not so protected, and speedily 
crumbles away when exposed to the action of water ; hence the 
vessel is placed in a protected situation, such as in a tree-stump, 
or a hole in a wall, or even in an empty snail-shell under a heap 
of stones. The cells are stored with spiders that have been 
paralysed by stinging and that serve as food for the larva of the 
Agenia. The larva of A. carbonaria has been described, and 
some particulars as to its habits have been given by Verhoeff. 



106 HYMENOPTERA 



CHAP. 



It has been stated that this wasp does not paralyse its prey 
by stinging, but substitutes a process of biting to prevent the 
spider from hurting the larva that is to feed on it ; and Verhoeff 's 
observations seem to show that the legs of the spider are broken 
by some proceeding of the kind. The Agenia larva is of peculiar 
shape, the head not being innexed, while the pleurae of each 
segment, from the second onwards, are prominent, so as to give 
the outline of the body a scalloped appearance. This larva is 
much infested by an Ichneumon that devours, it appears, not 
only the larva itself, but also the spider that was destined to be 
food for the larva. Verhoeff seems to have found some evidence 
that Pompilus sericeus may also be a parasite on the Agenia. 

The construction of earthenware cells, instead of the burrows 
usual in Pompilidae, by the species of this genus is one of the 
cases alluded to in our introductory remarks as to allied Fossores 
exhibiting different habits. Mr. Pride has recently sent us from 
Brazil similar earthen vessels constructed by some Pompilid. 

The habits of Pompilids of the genus Ceropales are analogous to 
those of the parasitic bees. Perez has recently given us information 
as to a very curious form of parasitism in this genus ; he says that 
when a Pompilus has obtained a spider as provision for its young, 
it is pursued by a Ceropales, which lays an egg un the spider, thus 
as it were substituting in advance its own young for that of the 
Pompilus. Information as to the subsequent course of events in 
this case is not at present forthcoming. In another case a 
Ceropales was observed to oviposit on the spider, not while this 
is being carried in, but subsequently by entering the nest for the 
purpose ; a habit quite similar to that of some parasitic bees. 
Ferton has recently made the unexpected discovery that some 
Pompilus act as robbers ; one individual taking away by force the 
spider that another has captured and is carrying off. 

Lichtenstein described a Pompilid larva, that he afterwards 
ascertained to be Calicurgus lii/nli nutus, as possessing the extra- 
ordinary habit of feeding as an external parasite fixed to the 
dorsal surface of a spider ; thus repeating, it would appear, the 
liabits of some of the Ichnemonidae, though the perfect Insect 
(Fig. 143) does not differ in structure from its congeners. 
Emery has given an account of some Pompilids that do not 
bury their prey, but after stinging it and depositing an egg, 
simply leave the spider on the spot, 



in FOSSORES SPHEGIDAE SPHEGIDES IO/ 

Buller has described the habits of a Pompilid in New Zealand ; 
his account is interesting because it shows a remarkable similarity 
in the proceedings of this antipodean wasp to those of its con- 
geners on our own side of the world. The species is not 
scientifically named, but it appears that it is known in New 
Zealand as " the Mason-bee." It forms a nest of yellow clay 
consisting apparently of about eight cells, each of which is filled 
with one or more spiders in a paralysed condition. The figure 
given of the larva of this Insect by Buller shows it to possess a 
peculiarly formed head. 

It is pleasing to find that Pompilidae do not make use of cruel 
methods when others will serve their purpose. We are informed 
that a large Australian Pompilid Priocnemis bicolor may find 
a Cicada sucking sap from a hole it has pierced in a tree. The 
Priocnemis has not the art of making the puncture necessary to 
procure sap, so the wasp seizes the Cicada, and shakes it till 
it leaves its hold and flies away, when the Priocnemis takes its 
place and sips the sap. It is added that the wasp never hurts 
the Cicada. 

Fam. 3. Sphegidae. 

Pronotum free from the tegulae ; when the stigmatic lobes extend as 
far back as the wing -insertion, they are placed below it and 
separated l>y a space from it. 

This large assemblage of Fossores is the one about which the 
greatest difference of opinion prevails. It is based entirely on 
the prothoracic characters mentioned above, and cannot be looked 
on as natural. We shall, however, follow Kohl x in treating for 
the present as only one family the divisions considered by many 
as distinct families. They are ten in number. 

Sub-Fam. 1. Sphegides. Hind body with a slender pedicel of 
variable length ; two spurs on the middle tibia. The pro- 
podeum usually horizontally elongate? 

This group includes a great number of species, about 200 of 
which are referred to the genus Sphex. 

1 "Die Gattungen der Sphegiden, " Ann. Hofmus. Wicn. xi. 1896, pp. 233-596. 
Seven plates. 

2 We will take this opportunity of correcting an error in the explanation of Fig. 
333 of the preceding volume, showing the propodeum, etc. of Sphex chrysis. f points 
to a division of the mesonoturn, not of the metanotum, as there stated. 



IO8 HVMENOPTERA 



CHAP. 



The habits of one species of this genus have been fully de- 
scribed by Fabre ; he assigns to the species the name of S. flavi- 
pennis, but Kohl considers that it is more probably S. maxillosus. . 
This Insect forms its nests, in the South of France, in the ground, 
excavating a main shaft with which are connected cells intended 
for the reception of the provisions for the young. The entrance 
to the burrow is formed by piercing a hole in the side of a very 
slight elevation of the soil. Thus the entrance to the construc- 
tion consists of a horizontal gallery, playing the part of a vesti- 
bule, and this is used by the Sphcx as a place of retreat and 
shelter for itself; at the end of the vestibule, which may be two 
or three inches long, the excavation takes an abrupt turn down- 
wards, extending in this manner another two or three inches, 
and terminating in an oval cell the larger diameter of which is 
situate in a horizontal plane. When this first cell has been com- 
pleted, stored with food, and an egg laid in it, the entrance to 
it is blocked up, and another similar cell is formed on one side ; 
a third and sometimes a fourth are afterwards made and pro- 
visioned, then the Insect commences anew, and a fresh tunnel is 
formed ; ten such constructions being the number usually prepared 
by each wasp. The Insect works with extreme energy, and as 
the period of its constructive activity endures only about a month, 
it can give but two or three days to the construction and pro- 
visioning of each of its ten subterranean works. The provisions, 
according to Fabre, consist of a large species of field-cricket, of 
which three or four individuals are placed in each cell. Kohl 
states, however, that in Eastern Europe an Insect that he 
considers to be the same species as Fabre's Spliex, makes use of 
locusts as provisions, and he thinks that the habit may vary 
according to the locality or to the species of Orthoptera that 
may be available in the neighbourhood. However that may 
be, it is clear from Fabre's account that this part of the 
^l>]iexs duties do not give rise to much difficulty. The cricket, 
having been caught, is paralysed so that it may not by its 
movements destroy the young larva for whose benefit it is 
destined. The Sphex then carries it to the burrow to store it in 
one of the cells; before entering the cell the Insect is in the 
habit of depositing its prey on the ground, then of turning round, 
entering the burrow backwards, seizing as it does so the cricket 
by the antennae, and so dragging it into the cell, itself going back- 



. in SPHEGIDAE SPHEGIDES IOQ 

wards. The habit of depositing its prey on the ground enabled 
Fabre to observe the process of stinging ; this he did by himself 
capturing a cricket, and when the wasp had momentarily quitted 
its prey, substituting the sound cricket for the paralysed one. 
The Spliex, on finding this new and lively victim, proceeds at 
once to sting it, and pounces on the cricket, which, after a brief 
struggle, is overcome by the wasp ; this holds it supine, and then 
administers three stings, one in the neck, one in the joint between 
the pro- and meso-thorax, and a third at the base of the abdomen, 
these three spots corresponding with the situation of the three 
chief nervous centres governing the movements of the body. 
The cricket is thus completely paralysed, without, however, being- 
killed. Fabre proved that an Insect so treated would survive for 
several weeks, though deprived of all power of movement. 
Three or four crickets are placed by the wasp in each cell, 100 
individuals or upwards being thus destroyed by a single wasp. 
Although the sting has such an immediate and powerful effect 
on the cricket, it occasions but a slight and evanescent pain to a 
human being ; the sting is not barbed, as it is in many bees and 
true wasps, and appears to be rarely used by the Insect for any 
other purpose than that of paralysing its victims. The egg is 
laid by the Spliex on the ventral surface of the victim between 
the second and third pairs of legs. In three or four days the 
young larva makes its appearance in the form of a feeble little 
worm, as transparent as crystal ; this larva does not change its 
place, but there, where it was hatched, pierces the skin of the 
cricket with its tiny head, and thus begins the process of feed- 
ing ; it does not leave the spot where it first commenced to feed, 
but gradually enters by the orifice it has made, into the interior 
of the cricket. This is completely emptied in the course of 
six or seven days, nothing but its integument remaining ; the 
wasp-larva has by this time attained a length of about 1 2 milli- 
metres, and makes its exit through the orifice it entered by, chang- 
ing its skin as it does so. Another cricket is then attacked and 
rapidly consumed, the whole stock being devoured in ten or twelve 
days from the commencement of the feeding operations ; the con- 
sumption of the later-eaten crickets is not performed in so delicate 
a manner as is the eating of the first victim. When full-grown, 
the process of forming a cocoon commences : this is a very ela- 
borate operation, for the encasement consists of three layers, in 



I I O HVMENOPTERA CHAP. 

addition to the rough silk that serves as a sort of scaffolding on 
the exterior : the internal coat is polished and is of a dark colour, 
owing to its being coloured with a matter from the alimentary, 
canal : the other layers of the cocoon are white or pale yellow. 
Fabre considers that the outer layers of the cocoon are formed 
by matter from the silk-glands, while the interior dark coat is 
furnished by the alimentary canal and applied by the mouth of 
the larva : the object of this varnish is believed to be the exclu- 
sion of moisture from the interior of the cocoon, the subterranean 
tunnels being insufficient for keeping their contents dry through- 
out the long months of winter. During the whole of the pro- 
cess of devouring the four crickets, nothing is ejected from the 
alimentary canal of the larva, but after the cocoon is formed 
the larva ejects in it, once for all, the surplus contents of the 
intestine. Nine months are passed by the Insect in the cocoon, 
the pupal state being assumed only towards the close of this 
period. The pupa is at first cpuite colourless, but gradually 
assumes the black and red colour characteristic of the perfect 
wasp. Fabre exposed some specimens of the pupa to the light 
in glass tubes, and found that they went through the pupal meta- 
morphosis in just the same manner as the pupae that remained in 
the darkness natural to them during this stage of their existence. 
Sphex coendetis is frequently stated to have the habit of pro- 
visioning its nests with both Orthoptera and Spiders; but Kohl 
considers with reason that this record is, as regards spiders, a 
mistake, arising probably from a confusion with some other 
Insect of similar appearance, such as Pelopacus (Sceliphron) 
coendcus. S. coeridcus is no doubt the same as S. (Chlorion) 
Inlmtt/s, which Eothney observed in East India, provisioning its 
nests with Orthoptera. He discovered a nest in process of con- 
struction, and during the absence of the mother-wasp abstracted 
from the burrow a large field-cricket that she had placed in it ; 
he then deposited the Orthopteron near the cell ; the parent 
X/i/K'.i- on returning to work entered the tunnel and found the 
provision placed therein had disappeared ; she came out in a state 
of excitement, looked for the missing cricket, soon discovered it, 
submitted it to the process of malaxation or kneading, and again 
placed it in the nest, after having cleared it from some ants 
that had commenced to infest it. She then disappeared, and 
Itothney repeated the experiment ; in due course the same series 



in SPHEGIDAE SPHEGIDES I I I 

uf operations was performed, and were repeated many times, the 
Sphex evidently acting in each case as if either the cricket had 
disappeared owing to its being incompletely stunned, or to its 
having been stolen by ants. Finally, the observer placed the 
cricket at a greater distance from the nest, when it recovered 
from the ill-treatment it had received sufficiently to make its 
escape. The points of interest in this account are the fact that 
the cricket was only temporarily paralysed, and that the wasp 
was quite able to cope with the two special difficulties that must 
frequently occur to the species in its usual round of occupations. 
The genus Ammophila is of wide distribution, and its species 
make vertical tunnels in the ground. The habits of some of the 
species found in France have been described by Fabre. The 
Insect does not inhabit the burrow while it is in process of 
formation, but quits it ; and some of the species temporarily 
close the entry to the incomplete nest with a stone. The 
tunnel is a simple shaft with a single cell at its termination ; 
this is stored with caterpillars, the different species of Ammo- 
phila selecting different grubs for the purpose. A. liirsuta hiber- 
nates in the perfect state, and carries on its work in the spring ; 
it chooses a single larva of considerable size belonging to one 
of the nocturnal Lepidoptera, and this it paralyses by a series of 
about nine stings, of which one is implanted in each segment 
from the first thoracic ring backwards ; it forms the burrow only 
after the food to be placed therein lias been obtained. The 
caterpillar used is subterranean in habit, and the Ammophila 
detects the larva by some sense, the nature of which appears at 
present quite uncertain. A. holosericea chooses smaller larvae of 
the family Geometridae, and uses only one or two stingings to 
paralyse each larva ; several caterpillars are used to provision a 
single cell, and they are often selected of different colours. 

Marchal has also published an important account of the 
proceedings of A. ajfinis ; he confirms Fabre's observations, and 
even adds to their interest by suggesting that the Ammophila 
administers special stings for the purpose of paralysing the 
mandibles of the caterpillar and depriving it of any power of 
afterwards injuring the larva that will feed on it. He thinks 
the motliQT-Ammojyhila herself profits by appropriating an exuda- 
tion from the victim. 

Some species of Sphegides have the curious habit of choosing 



112 HYMENOPTERA CHAP. 

the interiors of human habitations as the spots most suitable for 
the formation of their own domestic establishments. Fabre has 
given a charming account of the habits of Pelopaeus (Sceliphron} 
spirifex, a species that inhabits the South of Europe, and that forms 
its nests in the cottages of the peasants. The spot usually selected 
is a nook in the broad, open fireplace, out of reach of the names, 
though not of the smoke ; here the Pelopaeus forms a nest of 
earth, consisting of ten to fifty cells, the material being mud or 
clay brought in little balls by the aid of the Insect's mandibles ; 
about twenty visits are required in order to complete one cell, so 
that for the construction of a large nest of fifty cells, about one 
thousand visits must be made by the Insect. It flies in and 
out of the house apparently not at all incommoded by the 
human habitants, or by the fact that the peasant's potage may 
be simmering on the fire quite close to where the fearless little 
creature is carrying on its architectural operations. The cells 
are stored with spiders, of which the wasp has to bring a plentiful 
supply, so that its operations extend over a considerable period. 
The prey is captured by the Pelopaeus whilst on the wing, and 
carried off at once, being probably stung by the wasp during 
the process of transit ; apparently it is killed by the operation, 
not merely paralysed. Only small spiders are taken by this 
species, and the larva of the Pelopaeus consumes them in a short- 
time, one by one, before the process of decomposition sets in ; 
the egg, too, is laid on the first spider introduced, and this is of 
course at the bottom of the cell, so that the spiders are eaten by 
the wasp's larva in the order in which they were brought to the 
cell. The cell is sealed up when full, the number of spiders 
placed in it being on the average about eight. The larva 
completes its task of consuming the store in about ten days, and 
then forms a cocoon for its metamorphosis. TW T O or thivr 
generations are produced in a single year, the autumnal one 
passing eight or nine months in the clay cells, which are lodged 
in a nook of the peasant's hearth, and exposed to the smoke of 
his fire during all the months of winter. Pelopae.us (Scdipliroti) 
is a genus including many species ; * several of them are known 

1 Pelopaeus disappears from the new catalogue of Hymenoptera as the name of a 
valid genus; its species being assigned to ,sv/7/y>A/-n// \\\\<\ various other genera. 
We have endeavoured, as regards this name, to reconcile the nomenclature of 
previous authors with that used in the new catalogue by placing the generic name 
adopted in the latter in brackets. 



in SPHEGIDAE SPHEGIDES PELOPAEUS 113 

to be specially .attached to the habitations of human beings. 
Both has given an account of the habits of P. (Scelipliron} 
laetus in Australia ; he says that in some parts it is very 
difficult to keep these wasps out of the houses ; the nest is 
formed of mud, and constructed on the furniture or in any part 
of a room that suits the fancy of the Insect. This it must be 
admitted is, according to human ideas, liable to the charge of 
being very capricious. Both timed a wasp building its nest, 
and found that it brought a fresh load of mud every two or 
three minutes. If the wasp be allowed to complete the nest 
undisturbed, she does so by adding to the exterior diagonal 
streaks of mud, so giving to the nest the look of a small piece 
of the bark of a common acacia. The construction consists of 
from ten to twenty cells, and when completed is provisioned 
with spiders for the use of the young. This wasp is much 
pestered by parasites, some of which prevent the development of 
the larvae by consuming the spiders intended by the mother- 
wasp for its young. A fly, of the Order Diptera, is said to follow 
the wasp when carrying a spider, and to deposit also an egg on 
the food : as the Dipterous larvae have more rapid powers of 
assimilation, the Pelopaeus larvae are starved to death ; and their 
mildewed remains may be found in the cell, after their enemies 
have become fully developed and have flown away. Another 
parasite is said to eat the wasp-larva, and attains this end by 
introducing an egg through the mud wall and the cocoon of the 
wasp a habit that seems to indicate a Leucospid parasite. 
Tachytes australis, a wasp of the sub-family Larrides also dis- 
possesses this Pelopaeus in a manner we shall subsequently 
describe. This fragment of natural history from Australia has 
a special interest, for we find repeated there similar complex 
biological relations to those existing in the case of the European 



congeners. 



P. (SceUpliroti) madraspcdanus is common in the north-west 
provinces of Hindostan, and is called the " mud-dauber " by the 
European residents. According to Home it constructs its cells in 
the oddest places, but chiefly about the inhabited apartments in 
houses. It is perfectly fearless when engaged in building : the 
cells are four to six in number, and are usually provisioned with 
spiders to the number of about twenty. On one occasion 
it was observed that green caterpillars were stored instead of 

VOL. VI T 



HYMENOPTERA 



CHAP. 



spiders. The species is said to be protected by a peculiar odour 
as well as by its sting ; it is also stated that it disguises its 
edifice when completed by making it look like a dab of mud, 
and on one occasion " rays of mud were observed round the nest, 
even more exactly imitating a lump of mud thrown with some 
force." P. (Sceliphroii) bilineatus, formerly thought to be a variety 
of P. 'inadraspatanv-S, builds its nests in hedges and trees. 

Sub-Fam. 2. Ampulicides. Prothorax long and narrow. forming 
a neck in front ; chjpeus beak-like ; four submarginal cells, 
the outer one being complete ; metathorax elongate, the pos- 
terior part of the metasternum deeply divided to allow a 
perfect inflection of the abdomen. 

This is one of the smallest of the divisions of the Sphegidae, 
but has a very wide distribution, being represented in both 
the Eastern and Western Hemispheres. It is allied to the 
Sphegides, but differs by the prolongation of the neck and of the 
head, and by the articulation between the petiole and thorax 
being placed on the under surface of the body ; the wing- 
nervures are said to be of inferior importance owing to their 

frequently differing in indi- 
viduals of the same species. 
These Insects appear to be 
rare in individuals, as well as 
few in species, and but little 
has been recorded as to their 
habits ; but it is known that 
they live on cockroaches. Per- 
kins has given a brief sketch 
of the habits of Ampulex sibi- 
rica that is of great interest, 
but requires confirmation. He 
says that this Insect, in West 
Africa, enters apartments where 
cockroaches abound, and attack- 
ing one, that may probably be 
Male. four times its own size, suc- 
ceeds, after a struggle, in sting- 
ing it; the cockroach instantly becomes quiet and submissive, and 
suffers itself to lie lei] away and placed in confinement in some 




Fid. 4i. Am/in/' ' cnmpressa. 
East India. 



in SPHEGIDAE AMPULICIDES I I 5 

spot such as a keyhole, and in one case was apparently pre- 
vented from afterwards escaping, by the wasp carrying some 
heavy nails into the keyhole. The larva of the Ampulex may 
be presumed to live on the Blattid, as it is added that dead 
bodies of the cockroaches are frequently found with the empty 
cocoon protruding from them. This account, if correct, points to 
some features in the habits of this Insect that are unique. A 
remark made by Eothney in reference to the habits of A. (Elii- 
nopsis) rujicornis seems to indicate some similar instinct on the 
part of that species ; he says, " I also saw two or three of these 
wasps collar a peculiar cockroach by the antennae and lead it off 
into a crack in the bark, but as the cockroach reappeared smiling 
each time, I don't know what was up." The same observer records 
that this species associates with Sima rufonigra, an ant it greatly re- 
sembles in appearance, as well as with a spider that is also of similar 
appearance (Fig. 72). Schurr has given a brief account of the 
proceedings of Ampulex compressa, and his statements also tend 
to confirm the correctness of Perkins' report. The habits of a 
species of Ampulex \vere partially known to Reaumur, who 
described them on the authority of M. Cossigni. The species is 
believed to be A. compressa, which occurs not only in East India, 
but also in the island of Bourbon, the locality where M. Cossigni 
made his observation : his account is, like the others, a mere 
sketch of certain points observed, the most important of which 
is that when Ampidex cannot introduce the cockroach into a 
hole that it has selected as suitable, it bites off some portions 
of the body in order to reduce the poor Insect to the necessary 
extent. 

From these fragmentary observations it would appear that 
the sting of the Ampulex has not so powerful a paralysing effect 
as that of most other Fossores ; and that the Ampulex does 
not form any nest, but takes advantage of suitable holes and 
crevices to store the victim in ; also that it displays consider- 
able ingenuity in the selection of materials with which to block 
up the cavity in which it has placed the partially incapacitated 
creature. 

The genus Dolicliurus is by some entomologists considered 
the type of a sub-family allied to the Ampulicides ; it long 
consisted of a small and rare European Insect, but some exotic 
species have recently been added to it. It will probably prove not 



Il6 HYMENOPTERA 



CHAP. 



sufficiently distinct from Ampulicides, although the pronotum is 
much shorter, but Handlirsch has recently observed that the 
European species attacks Blattidae as do the normal Ampulicides ; 
and Ferton has recorded that D. haeinorrhous lives at the expense 
of Loloptera deei/iicn*, the wasp depositing its egg on the left 
intermediate femur of the prey. This is placed in a solitary 
cell, and is entirely consumed by the larva, life being preserved 
till within a few hours of the end of the repast, which occupies 
altogether eight days. 

Sub-Fam. 3. Larrides. Hind loch/ not pt'<licell<itf, or with 
only a short pedicel ; one spur on the mi<l<ll>' fi/n'n ; 
inconspicuous. Marginal cell of the front wings 
culate, 1 or mandibles excised externally, or loth. 

This group is by some writers called Tachy tides instead of 
Larrides, as owing to a change of nomenclature Txcliytcs may now 
be considered its principal genus. It is in connection with 
this and the neighbouring sub -families of Sphegidae that some 
of the greatest taxonomical difficulties exist. We include in 
Larrides the " Miscoplt us group " of Kohl. 

The species of the genus Tachy tes seem to have habits very 
similar to those of the genus Sphex ; they form shafts in the 
earth and provision them with Orthoptera ; like the Spltex and 
other Fossores, they have the habit, when they fly to their tunnel 
with a victim, of depositing it for a short time on the ground 
close to the mouth of the burrow while they turn round and 
enter backwards ; and, after doing this they again seize their 
prey and drag it into the burrow. Fabre availed himself of an 
opportunity to remove the prey while the Hymenopteron w;is 
entering the hole alone ; as a result it had to come out again to 
seek the object ; this it soon found, and carried to the hole, 
relinquishing it again as usual while it turned round ; Fabre 
repeated the operation several times, and always with the same 
result; the wasp, though it might have kept hold of the victim 
while it turned, and thus have saved itself from losing the 
precious object, never did so. 

1 "When a second cell is more or less perfectly marked out, the cell with which 
it is connected is said to lie appendiculate. Tin- nervnres I'reijiiently extend beyond 
the complete cells towards the outer margin, forming " incomplete " cells: only 
complete cells are counted, except when "incomplete" is mentioned. 



Ill 



SPHEGIDAE LARRIDES 



117 




FIG. 45. Tachytes pectinipes ?. 

Britain. 



One species of Tachytes in the south of France selects as its prey 
Orthoptera of the family Mantidae, Insects of a highly ferocious 
disposition, and provided with 
most powerful front legs, 
capable of cutting in two by 
a single act the body of an 
aggressor like the Tachytes; 
the latter is, however, by no 
means dismayed by the arms 
of its future victim, but hover- 
ing above the latter for some 
time, as if to confuse it, and 
causing it repeatedly to turn 
its very mobile head, the 
Tacliytcs at last pounces down 
and instantaneously stings the Mantis in the nerve centre 
between the formidable arms, which at once are reduced to in- 
capacity ; subsequently the Tachytes paralyses each of the other 
pairs of legs, and then carries off its victim. 

Larra anathema chooses mole-crickets as the viand for its 
young, and Tachysphex panzeri selects grasshoppers of the family 
Acridiidae. Larra pompiliformis ( = Tachytes nigcr, Fabre) some- 
times associates itself with Sphex flavipennis (? S. maxillosus, 
according to Kohl), forming its burrow amidst the works of a 
colony of that species, and making use, like the Sphex, of crickets 
for provender. This led Fabre to believe that the Larra stole 
its prey from the Spliex, but he has since withdrawn this indict- 
ment, and declares that the Larra obtains its crickets by the more 
honourable, if not more humane, process of catching and stinging 
them itself. Smith has informed us, on the faith of his own 
observation, that L. pompiliformis uses both Lepidopterous larvae 
and grasshoppers for its stores. 

T. (Larrada) australis, according to Whittell, plays the part of 
a burglar, breaking open the cells of Pelopaeus (Sceliphrori) laetus 
after they have been completed and stored with spiders ; it then 
takes possession of the cell, and curiously enough the Pelopaeus 
permits this, although the cell contains its egg and the store 
of food that is intended for the use of its own young. To us 
this seems very strange, but it is probable that the Pelopaeus 
has no idea of the consecpuences of the intruder's operations ; 



I I 8 HYMENOPTERA 



CHAP. 



it being one of the strange facts of nature that these highly 
endowed creatures never even see the offspring for whose welfare 
they labour with such extraordinary ingenuity and perseverance. 
Neither can we suppose that they have a conception of it 
derived from a knowledge of their own individual history ; for 
their very complete metamorphosis is scarcely reconcilable with 
any such recollection on their part. It may possibly therefore 
be the case that, having no idea whatever of the offspring, they 
are equally destitute of any conception that it will lie destroyed by 
the operations of the Larrada. However this may be, "Whittell 
informs us that both wasps skirmish about for a little as if each 
were mistrustful and somewhat afraid of the other ; this ends by 
the Pelopaeus withdrawing its opposition and by the Larrada 
taking . possession of the cell, which it then proceeds to divide 
into two, using for the purpose of the partition portions of the 
material of the nest itself; possibly it is only a contraction of 
the size of the cell, not a true division, that is effected ; however 
this may be, after it is accomplished the Larradn deposits its 
own egg in the cell, having, it is believed by "Whittell, previously 
destroyed that of the Pelopaeus. Judging from what occurs in 
other species it is, however, more probable that the destruction 
of the egg or young of the Pelopaeus is carried out by the larva 
of the Larrada and not by the parent-wasp. From a remark 
made by Maindron as to the proceedings of Larrada nivde^in, 
in Ternate, it seems probable that its habits may prove to be 
similar to those of L. australis, for it frequents the nests of 
Pelopaeus after they have been completed. 

Sub-Fam. 4. Trypoxylonides. Differ from Lan-ide* by the 
inner margin of the eyes being concave, and the marginal 
cell not appendiculate. (In Trypoxylon there is only one 
distinct submarginal and' one distinct discoidal cell, a second 
of each being indicated faintly.} 

The nervuration of Trypoxylon is very peculiar, and differs 
from that of the widely-distributed genus Pison, though according 
to Kohl's views the two may be correctly associated to form 
this sub-family. The species of Trypoxylon ;m- apparently rather 
fond of human propinquity, and build clay- or mud-nests in or 
inMi' houses. T. albitarse has this habit, and is well known in 
Southern Brazil under the name of " Marimbouda da casa " 



Ill 



SPHEGIDAE ASTATIDES BEMBECIDE 



119 



this Insect, like Pelopaeus, stores its nest with spiders, and 
Peckholt has remarked that however great may be the number of 
spiders placed by the mother-wasp in a cell, they are all consumed 
by the larva, none ever being found in the cell after the perfect 
Insect escapes therefrom. The European T. figulus forms a nest 
either in bramble-stems or in sandy soil or walls ; it makes use 
of spiders as provisions. 

Sub-Fam. 5. Astatides. Eyes very large in the male, meeting 
'broadly on the vertex ; two spurs on the middle tibia. 

We have two species of the genus Astata in Britain : one of 
them A. loops is known to form burrows in the ground, each 
of which contains only a single cell ; this, it appears, is usually 
provisioned with bugs of the 
genus Pentaloma, Insects re- 
markable for their strong and 
offensive odour. St. Fargeau 
records that this species also 
makes use of a small cockroach 
for forming the food - store : 
thus exhibiting an unique 
catholicity in the toleration of 
the disagreeable ; almost the 
only point of connection be- 
tween bus and cockroaches 




FlG ' 46 Astata 



male - Britaln - 



being their disagreeable char- 
acter. According to Smith, Oxybelus, another genus of Fossores, 
is also used. Authorities are far from agreement as to the 
validity and relations of the sub-family Astatides. It consists 
only of the widely-distributed genus Astata, with which the 
North American Diploplectron (with one species) is doubtfully 
associated. 

Sub - Fam. 6. Bembecides. - - Labrum frequently elongate ; 
wing - nervures extending very near to the outer margin; 
marginal cell of front wing not appendiculate ; mandibles 
not emarginale externally ; kind body stout, not pedicellate. 

The elongation of the labruni, though one of the most trust- 
worthy of the characters of the Bembecides, cannot be altogether 



I2O 



HYMENOPTERA 



CHAP. 



relied on owing to the variation it presents both in this and the 
allied sub-families. The Bembecides carry their prey to their 
young tucked underneath their own bodies and hugged to the 
breast ; they affect loose, sandy soils for nidification ; make use, 
in the great majority of the cases where the habits are known, 
of Diptera for provisions, and give these dead to the young ; 
making repeated visits to supply fresh food to the progeny, which 
notwithstanding this fact, are distributed in isolated burrows. 

One of the most interesting of Fabre's studies of the instincts 
of Hymenoptera is devoted to Bembex rostrata. The Bembecides 
have the habit of forming their nests in the ground in wide 

expanses of sand, and of cover- 
ing them up, they leave them 
so that there appears to be 
absolutely nothing by which 
the exact position of the nest 
can be traced ; nevertheless the 
!:< in hex flies direct to it with- 
out any hesitation. How neces- 
sary it is to these Insects to 
possess this faculty of finding 
their nests will be understood 
when we recall that the Bembex 




FIG. 47. Bembex rostrata 



Europe. 



does not provision its nest once and for all, but supplies the young 
at first with only insufficient food, and has therefore to return at 
daily, or other intervals, with a fresh store of provisions. The burrow 
is made in the sand by means of the fore-legs ; these work with 
such rapidity and skill that a constant stream of sand flows out 
behind the Insect while it is engaged in the act of excavation. 
The nest or cell in which the larva is to live, is formed by this 
process of digging ; but no fastening together of the material 
occurs, nor does any expedient seem to be resorted to, other 
than that of making a way through the sand by clearing out all 
the pieces of stick or stone that might diminish facility of access. 
The cell being formed, the Bembex leaves the spot in search of 
prey, and when it has secured a victim in the- shape of a two- 
winged fly, it returns therewith to the burrow, and the booty is 
placed therein, an egg being deposited on it. The wasp then 
leaves the burrow, disguising, however, the spot where it is 
situate, and Hies away: to proceed possibly with the formation 



in SPHEGIDAE BEMBECIDES 121 

of other burrows. 1 In the course of twenty -four hours the 
egg hatches, and the larva in two or three days completely 
devours the stock provided for it. The mother - wasp then 
returns with another fly this time probably a larger one- 
penetrates rapidly to the bottom of the burrow, and again re- 
treats, leaving the second stock of provisions for the benefit of 
the greedy larva. These visits of supply are repeated with increased 
frequency, as the appetite of the larva for the benefit of which 
they are made increases with its growth. During the fourteen 
or fifteen days that form this portion of the life-cycle, the single 
larva is supplied with no less than fifty to eighty flies for food. 
To furnish this quantum, numerous visits are made to each 
burrow, and as the mother Benibex has several burrows though 
how many does not appear to be known her industry at this 
time must be very great. All the while, too, a great danger has 
to lie avoided, for there is an enemy that sees in the booty 
brought by the Bembex to its young, a rich store for its own 
progeny. This enemy is a feeble, two-winged fly of the family 
Tachinidae and the genus Miltogramma ; it hangs about the 
neighbourhood of the nests, and sooner or later finds its 
opportunity of descending on the prey the Bembex is carrying, 
choosing for its purpose a moment when the Bcmlex makes a 
brief delay just at the mouth of the burrow ; then down comes 
the Miltogramma, and lays one, two, or three eggs on some portion 
of the booty that may be projecting from beneath the body of 
the wasp. This latter carries in the food for its own young, but 
thus introduces to the latter the source of its destruction, for the 
Miltogramma larvae eat up the supply of food intended for the 
Bembex larvae, and if there be not enough of this provender they 
satisfy their voracity by eating the Bembex larva itself. It is a 
remarkable fact that notwithstanding the presence of these 
strange larvae in the nest the mother Bembex continues to bring 
food at proper intervals, and, what is stranger still, makes no 
effort to rid the nest of the intruders : returning to the burrow 
with a supply of food she finds therein not only her legitimate 
offspring, a single tenant, but several others, strangers, it may 
be to the number of twelve ; although she would have no 
difficulty in freeing the nest from this band of little brigands, 
she makes no attempt to do so, but continues to bring the 

1 See on this point the note on p. 130. 



122 HVMENOPTERA CHAP. 

supplies. In doing so she is fulfilling her duty ; what matters 
it that she is nourishing the enemies of her race ? Both race 
and enemies have existed for long, perhaps for untold periods of 
time, why then should she disturb herself, or deviate from her 
accustomed range of duties ? Some of us will see in such pro- 
ceedings only gross stupidity, while others may look on them as 
sublime toleration. 

The peculiar habits of Bmibex rostrata are evidently closely 
connected with the fact that it actually kills, instead of merely 
paralysing, its prey ; hence the frequent visits of supply are neces- 
sary that the larvae may have fresh, not putrefying, food ; it may 
also be because of this that the burrow is made in a place of loose 
sand, so that rapid ingress may be possible to the Bembex itself, 
while the contents of the burrow are at the same time protected 
from the inroads of other creatures by the burrow being filled 
up with the light sand. Fabre informs us that the Bemlex 
larva constructs a very remarkable cocoon in connection with 
the peculiar nature of the soil. The unprotected creature has 
to pass a long period in its cocoon, and the sandy, shifting soil 
renders it necessary that the protecting case shall be solid and 
capable of keeping its contents dry and sound. The larva, how- 
ever, appears to have but a scanty supply of silk available for 
the purpose of constructing the cocoon, and therefore adopts the 
device of selecting grains of sand, and using the silk as a sort of 
cement to connect them together. For a full account of the 
ingenious way in which this difficult task is accomplished the 
reader should refer to the pages of Fabre himself. Bembe- 
cides appear to be specially fond of members of the Tabanidae 
(or Gad-fly family) as provender for their young. These 1 flies 
infest mammals for the purpose of feasting on the blood they 
can draw by their bites, and the Bembecides do not hesitate to 
capture them while engaged in gratifying their blood-thirsty pro- 
pensities. In North America a large species of Bembecid some- 
times accompanies horsemen, and catches the flies that come to 
attack the horses ; and Bates relates that on the Amazons a 
Bembecid as large as a hornet swooped down and captured one 
of the large blood-sucking Motuca flies that had settled on his 
neck. This naturalist has given an account of some of the 
Bembecides of the Amazons Valley, showing that the habits there 
are similar to those of their European congeners. 



in SPHEGIDAE NYSSONIDES 123 

Sphecius speciosus is a member of the Stizinae, a group recog- 
nised by some as a distinct sub-family. It makes use, in North 
America, of Insects of the genus Cicada as food for its young. 
Burrows in the ground are made by the parent Insect ; the egg- 
is deposited on the Cicada, and the duration of the feeding-time 
of the larva is believed to be not more than a week ; the pupa 
is contained in a silken cocoon, with which much earth is incor- 
porated. Eiley states that dry earth is essential to the well- 
being of this Insect, as the Cicada become mouldy if the earth 
is at all damp. As the Cicada is about twice as heavy as the 
Sphecius itself, this latter, when about to take the captured 
burden to the nest, adopts the plan of climbing with it to the 
top of a tree, or some similar point of vantage, so that during its 
flight it has to descend with its heavy burden instead of having 
to rise with it, as would be necessary if the start were made from 
the ground. 

Sub-Fam. 7. Nyssonides. Lcibrum short; mandibles entire on 
the outer edge; hind liody usually not pedicellate; wing 
with the marginal cell not appendicidate. 

This group has been but little studied, and there is not much 
knowledge as to the habits of the species. It is admitted to be 
impossible to define it accurately. It is by some entomologists 
considered to include MeUinus, in which the abdomen is pedi- 
cellate (Fig. 48), while others treat that genus as forming a 
distinct sub-family, Mellinides. Kohl leaves MeUinus unclassified. 
Gerstaecker has called attention to the fact that many of the 
Insects in this group have the trochanters of the hind and middle 
legs divided : the division is, as a rule, not so complete as it 
usually is in Hymenoptera Parasitica ; but it is even more marked 
in some of these Nyssonides than it is in certain of the parasitic 
groups. 

MeUinus arvensis is one of our commonest British Fossores, 
and we are indebted to the late F. Smith for the following 
account of its habits : " It preys upon flies, and may be commonly 
observed resorting to the droppings of cows in search of its prey; 
it is one of the most wary and talented of all its fraternity ; were 
it at once to attempt, by a sudden leap, to dart upon its victim, 
ten to one it would fail to secure it ; no, it does no such thing, 
it wanders about in a sort of innocent, unconcerned way, amongst 




124 HYMENOPTERA CHAP. 

the deluded flies, until a safe opportunity presents itself, when 

its prey is taken without any 
chance of failure ; such is its 
ordinary mode of proceeding. At 
Bournemouth the flies are more 
active, more difficult to capture, 
or have they unmasked the 
treacherous Mellinus ? and is it 
found necessary to adopt some 
fresh contrivance in order to 
* accomplish its ends ( if so, it is 

FIG. 48. Mellinus arcensis ?. Britain. i .c i T 

not deficient in devices. 

noticed once or twice, what I took to he a dead specimen of 
Mellinus, lying on patches of cow-dung ; but on attempting 
to pick them up off they flew ; I at once suspected the crea- 
ture, and had not long to wait before my suspicions were 
confirmed. Another, apparently dead fellow, was observed ; and 
there, neither moving head or foot, the treacherous creature lay, 
until a fine specimen of a Bluebottle ventured within its grasp, 
when, active as any puss, the Mellinus started into life, and 
pounced upon its victim." 

Lucas states that in the north of Prance Mellinus salnilosus 
provisions its nest with Diptera, which it searches for on the 
flowers of Umbelliferae, and then carries to its nest. This is a 
burrow in the earth, and when it is reached the Hymeuopteron 
deposits its Insect burden for a moment on the ground while it 
turns round in order to enter the burrow backwards. The same 
writer states that two varieties of this Insect live together or 
rather in the same colonies and make use of different species 
of Diptera, even of different genera, as food for their young. 
These Diptera are stung before being placed in the nest. The 
stinging does not kill the Insect, however, for Lucas was able to 
keep one specimen alive for six weeks after it had passed this 
trying ordeal. 

Sub-Fam. 8. Philanthides. Laltnun small ; anterior icings ivith 
three complete, submarginal o /As ; him I l>t/// constricted at 
the base Imt not so as to form a slender pedicel. 

This sub-family contains Insects resembling wasps or Tra- 
limnides in appearance, and is, as regards the pronotal structure, 



Ill 



SPHEGIDAE- -PHILANTHIDES 



125 



intermediate between the two great divisions of the Fossores, for 
the pronotal lobe extends nearly or quite as far back as the 
tegulae, and in Philanthus the two come into almost actual 
contiguity. 

The species of the genus Cerceris are numerous in Europe, 
and several of them are known to make burrows in the ground, 
and store them with beetles for the benefit of the future larvae. 
The beetles chosen differ in family according to the species of 
Cerceris ; but it appears from the observations of Fabre and Dufour 
that one kind of Cerceris 
never in its selection goes 
out of the limits of a 
particular family of 
beetles, but, curiously 
enough, will take Insects 
most dissimilar in form 
and colour provided they 
belong to the proper 
family. This choice, so 
wide in one direction and 
so limited in another, 
seems to point to the 
existence of some sense, 




FIG. 49. Philanthus triaiiyulttm 6- Britain. 



of the nature of which we are unaware, that determines the 
selection made by the Insect. In the case of our British species 
of Cerceris, Smith observed C. arenaria carrying to its nest Cur- 
culionidae of very diverse forms ; while C. labiata used a beetle 
Haltica tabida of the family Chrysomelidae. 

The beetles, after being caught, are stung in the chief 
articulation of the body, that, namely, between the pro- and 
mesothorax. Cerceris lupresticida confines itself exclusively to 
beetles of the family Buprestidae. It was by observations on 
this Insect that Dufour first discovered the fact that the Insects 
stored up do not decay : he thought, however, that this was due 
to the liquid injected by the wasp exercising some antiseptic 
power; but the observations of Fabre have shown that the pre- 
servation in a fresh state is due to life not being extinguished ; 
the stillness, almost as if of death, being due to the destruction 
of the functional activity of the nerve centres that govern the 
movements of the limbs. 



126 IIYMENOPTERA 



CHAP. 



It has long been known that some species of Cerceris prey on 
bees of the genus Hal ictus, and Marchal has recently described 
in detail the proceedings of C. ornata. This Insect catches 'a 
Halictus on the wing, and, holding its neck with the mandibles, 
bends her body beneath it, and paralyses it by a sting admin- 
istered at the front articulation of the neck. The Halictus is 
subsequently more completely stunned or bruised by a process of 
kneading by means of the mandibles of the Cerceris. Marchal 
attaches great importance to this " malaxation " ; indeed, he is of 
opinion that it takes as great a part in producing or prolonging 
the paralysis as the stinging does. Whether the malaxation 
would be sufficient of itself to produce the paralysis he could not 
decide, for it appears to be impossible to induce the Cerceris to 
undertake the kneading until after it has reduced the Halictus 
to quietude by stinging. 

Fabre made some very interesting observations on Cerceris 
tnlerculiita, their object being to obtain some definite facts as to 
the power of these Insects to find their way home when removed 
to a distance. He captured twelve examples of the female, 
marked each individual on the thorax with a spot of white 
paint, placed it in a paper roll, and then put all the rolls, with 
their prisoners, in a box ; in this they were removed to a distance 
of two kilometres from the home and then released. He visited 
the home five hours afterwards, and was speedily able to assure 
himself that at any rate four out of the twelve had returned to 
the spot from whence they had been transported, and he enter- 
tained no doubt that others he did not wait to capture had been 
equally successful in home- finding. He then commenced a second 
experiment by capturing nine examples, marking each with two 
spots on the thorax, and confining them in a dark box. They 
were then transported to the town of Carpentras, a distance of 
three kilometres, and released in the public street, " in the centre 
of a populous quarter," from their dark prison. Each Cerceris 
on being released rose vertically between the houses to a sufficient 
height, and then at once passed over the roofs in a southerly 
direction the direction of home. After some hours he went 
back to the homes of the little wasps, but could not find that 
any of them hail then returned; the next day he went again, 
and found that at any rate five of the Cerceris liberated the 
previous day were then at home. This record is of considerable 



in SPHEGIDAE MIMESIDES I2/ 

interest owing to two facts, viz. that it is not considered that the 
Cerceris as a rule extends its range far from home, and that the 
specimens were liberated in a public street, and took the direc- 
tion of home at once. 

Philanthus apivorus is one of the best known of the members 
of this sub -family owing to its habit of using the domestic 
honey-bee as 'the food for its offspring. In many respects its 
habits resemble those of Cerceris ornata, except that the Pkil- 
anthus apparently kills the bee at once, while in the case of the 
Cerceris, the' Hal-ictus it entombs does not perish for several days. 
The honey-bee, when attacked by the Philanthus, seems to be 
almost incapable of defending itself, for it appears to have no 
power of finding with its sting the weak places in the armour, 
of its assailant. According to Fabre, it has no idea of the Phil- 
anthus "being the enemy of its race, and associates with its 
destroyer on amicable terms previous to the attack being made on 
it. The Philanthus stings the bee on the under- surface of the 
mentum ; afterwards the poor bee is subjected to a violent process 
of kneading, by which the honey is forced from it, and this the 
destroyer greedily imbibes. The bee is then carried to the nest 
of the Philanthus. This is a burrow in the ground ; it is of 
unusual depth about a yard according to Fabre and at its ter- 
mination are placed the cells for the reception of the young ; in 
one of these cells the bee is placed, and an egg laid on it : as 
the food iu this case is really dead, not merely in a state of 
anaesthesia, the Pkilanthus does not complete the store of food 
for its larvae all at once, but waits until the latter has consumed 
its first stock, and then the mother-wasp supplies a fresh store 
of food. In this case, therefore, as in Bembex, the mother really 
tends the offspring. 

Sub-Fam. 9. Mimesides. - - Small Insects with pedicellate hind 
body, tlie pedicel not cylindric ; mandibles not excised ex- 
ternally ; inner margin of eyes not concave ; middle tibia 
with one spur ; wings with two, or three, submarginal cells. 

Mimesides is here considered to include the Pemphredonides 
of some authors. Mimesides proper comprises but few forms, and 
those known are small Insects. Psen concolor and P. a.tratus 
form their nests in hollow stems, and the former provisions its 
nest with Homopterous Insects of the family Psyllidae. Little 



128 



HYMENOPTERA 



CHAP. 




FIG. 50. Mimesa bicolor f, 
Britain. 



information exists as to their liabits ; but Verhoeff states that 

the species of Pscn like mem- 
bers of the Pemphredoninae 
do not form cocoons. 

The Pemphredonine subdivi- 
sion includes numerous small and 
obscure Insects found chiefly in 
Europe and North America (Fig. 
51, P. lugubris)', they resemble 
the smaller black species of Cra- 
bronides, and are distinguished 
from them chiefly by the exist- 
ence of at least two complete, 
submarginal cells on the an- 
terior wing instead of one. 
The species of Passaloecus live in the burrows that they form 

in the stems of plants ; Pemphredon lugubris frequents the decayed 

wood of the beech. The larva and pupa of the latter have been 

described by Verhoeff; no 

cocoon is formed for the 

metamorphosis. Both these 

genera provision their nests 

with Aphidae. This 

also the case with Stt 

pendulus, but the burrows 

of this species form a com- 

plex system of diverticula 

proceeding from an irregu- 

lar main channel formed in 

the pithy stems of bushes. 

Cemonus unicolor, according to Giraud, forms its burrows in 

bramble - stems, but it also takes advantage, for the purposes 

of nidification, of the abandoned galls of Ct/nij>n, and also ol 

a peculiar swelling formed by a fly Llpm-n lucens on the 

common reed, Arundo p/i /</>/ mites. This species also makes 

use of Aphidae, and Verhoeff states that it has only an imperfect 

instinct as to the amount of iood it stores. 

Sub-Fam. 10. Crabronides --Pi'onoitun short, front u-rmj u-iih 
one co////>/i fr submarginal and two discoidal cell* : hind body 



s 




FIG. 51. Pemphredon luyitbris 9. Britain. 



Ill 



SPHEGIDAE CRABRONIDES 



129 




/(triable in form, pedicellate in some abnormal forms, but more 
usually not stalked. 

The Crabronides ( Vespa crabro, the hornet, is not of this sub- 
family) are wasp-like little Insects, with unusually robust and 
quadrangular head. They frequently have the hind tibiae more 
or less thickened, and the clypeus covered with metallic hair. 
It appears at present 
that they are specially 
attached to the tem- 
perate regions of the 
northern hemisphere, 
but this may possibly be 
in part due to their 
having escaped attention 
elsewhere. In Britain 
they form the most im- 
portant part of the 
fossorial Hymenoptera, 
the genus Crabro 
(with numerous sub- 
genera) itself comprising thirty species. The males of some of 
the forms have the front tibiae and tarsi of most extraordinary 
shapes. They form burrows in dead wood, or in pithy stems, 
(occasionally in the earth of cliffs), and usually store them with 
Diptera as food for the larvae : the wings and dried portions of 
the bodies of the flies consumed by Crabronides are often exposed 
to view when portions of old wood are broken from trees. 

The genus Oxybelus is included by some systematists, but 
with doubt, in this sub-family ; if not placed here, it must form a 
distinct sub-family. It has the metathorax spinose, and the sub- 
marginal and first discoidal cells are not, or are scarcely, separated. 

Crabro leucostomus has been observed by Fletcher to form 
cells for its larvae in the soft wood of broken willows : the food 
stored therein consists of two-winged flies of the family Dolicho- 
podidae. This Crabro is parasitised by an Ichneumonid of the 
genus Tryphon, and by a two-winged fly of uncertain genus, but 
belonging to the family Tachinidae. The metamorphoses of 
Crabro clirysostomus have been briefly described by Verhoeff: 
the food stored consists of Diptera, usually of the family Syr- 

VOL. VI K 



FIG. 52. Crabro cephalotes ? . Britain. 



130 HYMENOPTERA CHAP, in 

phidae ; the larva spins an orange-red cocoon, passes the winter 
therein, and assumes the pupal form in the spring ; there is, he 
says, a segment more in the female pupa than there is in the male. 

The species of the sub-genus Crossocerus provision their nests 
with Aphididae, but C. wesmaeli makes use, for the purpose, 
according to Ferton, of an elegant little fly of the family 
Tipulidae ; according to Pissot this same wasp also makes use 
of a species of Typldocylta, a genus of the Homopterous division of 
Rhynchota. Supposing there to be no mistake as to this latter 
observation, the choice of Diptera and of Homoptera by the same 
species indicates a very peculiar habit. 

Fertonius (Crossocerus} luteicollis in Algeria forms cells at a 
slight depth in sandy soil, and provisions them with ants. The 
ant selected is Tapinoma erraticum, and the individuals captured 
are the wingless workers. The mode of hunting has been de- 
scribed by Ferton ; the \vasp hovers over one of the ant-paths at a 
distance of a few millimetres only above the surface, and when 
an ant that is considered suitable passes, the Fertonius pounces 
on it, stings it, and carries it off to the burrow ; forty or fifty 
ants are accumulated in a cell, the egg is laid in the heap of 
victims about one-third of the depth from the bottom; the 
resulting larva sucks the ants one by one, by attaching itself to 
the thorax behind the first pair of legs. There is a very 
interesting point in connection with the habits of this species, 
viz. that the ants are not only alive, but lively ; they have, 
however, lost the power of co-ordinating the movements of the 
limbs, and are thus unable to direct any attack against the feeble 
larva. Ferton thinks there are three generations of this species 
in a single year. 

NOTE. In a note on p. 99 we have mentioned the new publication of 
Mr. and Mrs. Peckham on the habits of Fossores. We may here add that it 
contains much fresh information on these Insects, together with criticisms of 
the views of Fabre and others. One of the points most noteworthy is that 
they have observed Crabro stirpicola working night and day for a period of 
forty-two consecutive hours. They made experiments on Sembex spinolae 
with a view of ascertaining whether the female provisions two nests simul- 
taneously ; as tlie result they think this improbable. If the female Beinbeeid 
make nests only consecutively, it is clear it must have but a small fecundity. 
The larval life extends over about fifteen days ; and if we allow three months 
as the duration of life of a female, it is evident that only about six young 
can be produced in a season. 



CHAPTER IV 



HYMENOPTERA ACULEATA COXTIXUED- 

OR ANTS 



-DIVISION IV. FORMICIDAE 




B 



Division IV. Heterogyna or Formicidae Ants. 

The segment, or the two segments, behind the propodeum, either 
small or of irregular form, so that if not throughout of 
small diameter, the articulation until the segment behind is 
slender, and there is great mobility. 
The trochanters undivided. The 
individuals of each species are 
usually of three kinds, males, females 
and workers ; the latter h are no 
wings, but the males and females 
(ire usually winged, though the 
females soon lose the flying organs. 
They live in communities of various 
numbers, the majority being workers. 
The larvae are helpless maggots fed 
<f nd tended by the ivorkers or by 

" FIG. 53 Abdomens of ants. 

A, Of Camponotus ruli-i/^.-i 

IN ants the distinction between the (Formicides) ; B, of Ecta- 

,-, . ^,111- tnmma auralum (Ponerides); 

three great regions of the body is very c> of Aphmnog^i., bar- 
marked. The abdomen is connected bara (Myrmicides). , Pro- 

. . podeum ; b, first abdominal 

with the propodeum in a peculiar manner, segment forming a scale or 
one or two segments being detached node : c, second ; rf, third 

abdominal segment. 

from the main mass to form a very 

mobile articulation. This is the most distinctive of the char- 
acters of ants. The structure and form of these parts varies 






132 HYMENOPTERA CHAP. 

greatly in the family : and the Amblyoponides do not differ in 
a marked manner from the Scoliidae in ibssorial Hymenoptera. 

The arrangement of the parts of the mouth is remarkable, 
and results in leaving the mandibles quite .free and unconnected 
with the other trophi ; the mouth itself is, except during feeding, 

closed completely 
by the lower lip 
and maxilla assum- 
ing an ascending 
vertical direction, 
while the upper 

FIG. 54.-Front of head of Dinopon y h d 

A, Mouth closed ; B, opi-n. 

and overlaps the 

lower lip, being closely applied to it ; so that 1 in Ponerides 
the palpi, except the apices of the maxillary pair, are enclosed 
between the upper and lower lips (Fig. 54, A). In Cryptocerini 
the palpi are not covered by the closed lips, but are protected by 
being placed in chinks at the outsides of the parts closing the 
mouth. The mandibles of ants can thus be used in the freest 
manner without the other parts of the mouth being opened or even 
moved. The mandibles close transversely over the rest of the 
mouth, and when shut are very firmly locked. There are, 
however, some ants in which the lips remain in the position 
usual in mandibulate Insects. 

The antennae, except in the males of some species, have a 
long basal joint and are abruptly elbowed at its extremity. The 
eyes and ocelli vary excessively, and may be totally absent or 
very highly developed in the same species. The winged forms 
are, however, never blind. The size of the head varies extremely 
in the same species ; it is frequently very small in the males, 
and largest in the workers. In some ants the worker-caste 
consists of large-headed and small- headed individuals ; the former 
are called soldiers, and it has been supposed that some of them 
may act the part of superior officers to the others. It should be 
clearly understood that there is no definite distinction between 
soldiers and workers; so that in this respect they are widely 
different from Termites. 

The complex mass forming the thorax is subject to great 
change of structure in the same species, according as the indi- 
viduals are xvinged or wingless. The sutures between the dorsal 



FORMICIDAE ANTS 



133 



(notal) pieces are frequently obliterated in the workers, while 
they are distinct in the males and females, and the pieces them- 
selves are also much larger in size in these sexed individuals. 
The pro-mesothoracic stigma is 
apparently always distinct ; the 
meso-metathoracic one is distinct 
in the male Dorylus, but can scarcely 
be detected in the winged forms of 
other ants, owing to its being en- 
closed within, and covered by, the 
suture between the two segments : 
in the workers, however, it is usually 
quite conspicuous. The posterior 
part of the thoracic mass, the pro- 
podeum or median segment, is of 
considerable size ; no transverse 
suture between the component pieces 
of this part can be seen, but its 
stigma is always very distinct. The Fia ^_ 0ecodoma cephaMes . South 

peduncle, Or pedicel, formed by the America. A, Worker major ; B, 

, ., female after casting the wings. 

extremely mobile segment or seg- 
ments at the base of the abdomen (already noticed as form- 
ing the most conspicuous character of the family), exhibits much 





FIG. 56. Stridulating organ of an ant, Myrmica rubra, var. laevinodis. Sagittal section 
of part of the 6th and 7th post-cephalic segments. (After Janet.) a, a 1 , muscles ; 
6, connecting membrane (corrugated) between 6th and 7th segments : c, 6th seg- 
ment ; d, its edge or scraper ; e, striate area, or file on 7th segment ; /, posterior 
part of 7th segment ; g, cells, inside body ; ,h, trachea. 

variety. Sometimes the first segment bears a plate or shield 
called a scale (Fig. 53, A, V) ; at other times there are two 



134 



HYMENOPTERA 



CHAP. 




small segments (Fig. 53, B, C, l>, c~) forming nodes or knots, of 

almost any shape. The articulations between these segments are 

of the most perfect description. In many ants these parts bear 

4 highly developed stridulating organs, and the delicacy and perfection 

of the articulations allow the parts to be 
moved either with or without producing 
stridulation. In the male sex the peduncle 
and its nodes are much less perfect, and 
possess comparatively little capacity for 
movement ; in the male of Dorylus (Figs. 
79, A, and 80, /) the single node is 
only imperfectly formed. The eyes and 
ocelli of the males are usually more 
largely developed than they are in the 
female, though the head is much smaller. 
The legs of ants are elongate, except 
in a few forms ; the Cryptocerini and 
the males of Dorylides being the most 
conspicuous exceptions. The tarsi are 
five -jointed, the basal joint being dis- 
proportionately elongate, so that in use 
Fro. 57. Combs and brushes : . . / , 

on front leg of an ant, Dino- ^ acts in many species as it it were a 
ponera grandis (tip of tibia, portion of the tibia, the other four joints 

1 .earing the comb-like spur, -in in r rl " f 

and the base of the first joint forming the functional loot. ilie Il'Ollt 

of the tarsus ; cf. fig. 75). tibiae are f urn i s hed with a beautiful 

A, Inner, B, outer aspect. 

combing apparatus (Fig. 57) 

Features of Ant-life. In order that the reader may realise 
the nature of ant-life we may briefly recount its more usual and 
general features. Numerous eggs are produced in a nest by one 
or more queens, and are taken care of by workers. These eggs 
hatch and produce helpless maggots, of which great care is 
taken by the workers. These nurses feed their charges from 
their own mouths, and keep the helpless creatures in a fitting- 
state by transporting them to various chambers in conformity 
with changes of temperature, humidity, and so on. When full 
grown the maggots change to pupae. In some species the 
maggots form cocoons for themselves, but in others this is not 
the case, and the pupae are naked. 1 After a brief period of 

1 The pupae and cocoons of ants are usually called by the uninstructed, "ants' 
eggs." In this country they are used as food for pheasants. 



iv FORMICIDAE ANTS 135 

pupal life a metamorphosis into the perfect Insect occurs. The 
creatures then disclosed may be either winged or wingless ; the 
wingless are the workers and soldiers imperfect females the 
winged are males or females fully developed. The workers re- 
main in or near the nest they were produced in, but the winged 
individuals rise into the air for a nuptial flight, often in great 
numbers, and couple. When this is accomplished the male 
speedily dies, but the females cast their wings and are ready to 
enter on a long life devoted to the production of eggs. From 
this account it will be gathered that males are only found in 
the nests for a very short time ; the great communities consist- 
ing at other periods entirely of the two kinds of females and of 
young. The imperfect females are themselves in some species 
of various kinds ; each kind being restricted, more or less com- 
pletely, to a distinct kind of duty. 

No Insects are more familiar to us than ants ; in warm 
countries some of them even invade the habitations of man, or 
establish their communities in immediate proximity to his 
dwellings. Their industry and pertinacity have, even in remote 
ages, attracted the attention and admiration of serious men ; some 
of whom we need scarcely mention Solomon as amongst them 
have not hesitated to point out these little creatures as worthy 
of imitation by that most self-complacent of all the species of 
animals, Homo sapiens. 

Observation has revealed most remarkable phenomena 
in the lives of these Insects. Indeed, we can scarcely avoid 
the conclusion that they have acquired in many respects 
the art of living together in societies more perfectly than 
our own species has, and that they have anticipated us in the 
acquisition of some of the industries and arts that greatly 
facilitate social life. The lives of individual ants extend over a 
considerable number of years in the case of certain species at 
any rate so that the competence of the individual may be 
developed to a considerable extent by exercise ; and one genera- 
tion may communicate to a younger one by example the arts 
of living by which it has itself profited. The prolonged life of 
ants, their existence in the perfect state at all seasons, and the 
highly social life they lead are facts of the greatest biological 
importance, and are those that we should expect to be accom- 
panied by greater and wider competence than is usually exhibited 



136 HYMENOPTERA 



CHAP. 



by Insects. There can indeed be little doubt that ants are really 

t a 

not only the " highest " structurally or mechanically of all Insects, 
but also the most efficient. There is an American saying to 
the effect that the ant is the ruler of Brazil. We must add a 
word of qualification ; the competence of the ant is not like that 
of man. It is devoted to the welfare of the species rather than to 
that of the individual, which is, as it were, sacrificed or specialised 
for the benefit of the community. The distinctions between the 
sexes in their powers or capacities .are astonishing, and those 
between the various forms of one sex are also great. The differ- 
ence between different species is extreme ; we have, in fact, the 
most imperfect forms of social evolution coexisting, even locally, 
with the most evolute. 

These facts render it extremely difficult for us to appreciate 
the ant ; the limitations of efficiency displayed by the individual 
being in some cases extreme, while observation seems to elicit 
contradictory facts. About two thousand species are already 
known, and it is pretty certain that the number will reach at 
least five thousand. Before passing to the consideration of a 
selection from what has been ascertained as to the varieties of 
form, and of habits of ants we will deal briefly with their habita- 
tions and polymorphism, reserving some remarks as to their 
associations with other Insects to the conclusion of this chapter. 

Nests. Ants differ greatly from the other Social Hymenoptera 
in the nature of their habitations. The social bees construct cells 
of wax crowded together in large numbers, and the wasps do the 
like with paper ; the eggs and young being placed, each one in a 
separate cell, the combinations of which form a comb. Ants 
have, however, a totally different system ; no comb is constructed, 
and the larvae are not placed in cells, but are kept in masses and 
are moved about from place to place as the necessities of tempera- 
ture, air, humidity and other requirements prompt. The habita- 
tions of ants are in all cases irregular chambers, of which there is 
often a multiplicity connected by galleries, and they sometimes 
form a large system extending over a considerable area. Thus 
the habitations of ants are more like those of the Termites than 
those of their own allies among the Hymenoptera. They are 
chiefly remarkable for their great variety, and for the skilful 
manner in which they are adapted by their little artificers to 
particular conditions. The most usual form in Europe, is a 



IV 



ANTS' HABITATIONS 



'37 




number of subterranean chambers, often under the shelter of a 
stone, and connected by galleries. It is of course very difficult to 
trace exactly the details of such a work, because when excavations 
are made for the purposes of examination, the construction becomes 
destroyed : it is known, however, that some of these systems 
extend to a considerable depth in the earth, it is said to as much 
as nine feet, and it is thought the object of this is to have access 
to sufficiently moist earth, for ants are most sensitive to variations 
in the amount of moisture ; 
a quite dry atmosphere is in 
the case of many species very 
speedily fatal. This system 
of underground labyrinths 
is sometimes accompanied by 
above-ground buildings con- 
sisting of earth more or less 
firmly cemented together by 
the ants ; this sort of dwell- 
ing is inOSt frequently adopted FlG . 58. Portion of combined nest of For- 
whell the soil ill which the ' aml Solenopsis fugax (Alter 

nests are placed is sandy ; it 
is probable that the earth is 
in such cases fastened together 
by means of a cement pro- 
duced by the salivary glands 
of the ants, but this has not 
been determined with certainty ; vaulted galleries or tunnels of 
this kind are constructed by many species of ants in order to 
* -liable them to approach desired objects. 

In South America Camponotus ruftpes and other species that 
habitually dwell in stumps, in certain districts where they are 
liable to inundations, build also nests of a different nature on 
trees for refuge during the floods. In Europe, a little robber-ant, 
Solenopsis fugax, constructs its dwelling in combination with that 
of Fn' i/i ir/f f<i*ca (Fig. 58), in such a manner that its chambn> 
cannot, on account of the small size of the orifices, be entered by 
the much larger Fur mint. Hence the robber obtains an easy 
living at the expense of the larger species. The Sauba or Sauva 
ants of South America (the genus Atta of some, Oecodoma of 
other authors) appear to be most proficient in the art of sub- 



Forel.) x |. f, f, Chambers of Formica, 
recognisable by the coarser shading ; 
s, s', chambers of the Solenoftsis (with 
finer shading) ; s", opening in one of the 
chambers, the entrance to one of the 
galleries that connects the chambers of the 
Solenopsis; ic, walls forming the founda- 
tions of the nest and the limits of the 
chambers. 



'38 



HYMENOPTERA 



CHAP. 



terranean mining. Their systems of tunnels and nests are known 
to extend through many square yards of earth, and it is said on 
the authority of Hamlet Clark that one species tunnelled under' 
the bed of the river Parahyba at a spot where it was as broad 
as the Thames at London Bridge. 

A considerable number of ants, instead of mining in the 
ground, form chambers in wood; these are usually very close to 
one another, because, the space being limited, galleries cannot be 
indulged in. Ctimponotus ligniperdus in Europe, and 0. pennsyl- 
vanicus in North America, work in this way. 

Our British Lasius fuliginosus lives in decayed wood. Its 
chambers are said by Forel to consist of a paper-like substance 
made from small fragments of wood. Cryptocerus burrows in 

branches. Colobopsis lives in a 
similar manner, and Forel in- 
forms us that a worker with 
a large head is kept stationed 
within the entrance, its great head 
acting as a stopper ; when it sees 







a nest-fellow desirous of entering 



the nest, this animated and intel- 
ligent front-door then retreats a 
little so as to make room for 
ingress of the friend. Forel has 
observed that in the tropics of 
America a large number of species 
of ants live in the stems of grass. 
There is also quite a fauna of 
ants dwelling in hollow thorns, 
in spines, on trees or bushes, or 
in dried parts of pithy plants ; 
and the tropics also furnish a 
number of species that make nests 
of delicate paper, or that spin 
together by means of silk the 
leaves of trees. One eastern 
species Polyrhachis spiniycra fabricates a gauze-like web of 
silk, with which it lines a subterranean chamber after the 
manner of a trap-door spider. 

Some species of ants appear to lind both food and shelter 



Kii;. . r i9. Ant-plant, Hydnophytwn mon- 
taniim. Java. (After Forel.) 



iv ANTS 139 

entirely on the tree they inhabit, the food being usually sweet 
stuff secreted by glands of the plant. It is thought that the 
ants in return are of considerable benefit to the plant by defend- 
ing it from various small enemies, and this kind of symbiosis has 
received much attention from naturalists. A very curious con- 
dition exists in the epiphytic plants of the genera Myrmecodia 
and Hydnophytum ; these plants form large bulb-like (Fig. 59) 
excrescences which, when cut into, are found to be divided into 
chambers quite similar to those frequently made by ants. Though 
these structures are usually actually inhabited by ants, it appears 
that they are really produced by the plant independent of the 
Insects. 

Variability and Polymorphism of Ants. Throughout the 
Hymenoptera there are scattered cases in which one of the sexes 
appears in dimorphic form. In the social kinds of bees and 
wasps the female sex exists in two conditions, a reproductive 
one called queen, and an infertile one called worker, the limits 
between the two forms seeming in some cases (honey-bee) to be 
absolute as regards certain structures. This sharp distinction 
in structure is rare ; while as regards fertility intermediate con- 
ditions are numerous, and may indeed be induced by changing 
the social state of a community. 1 In ants the phenomena of the 
kind we are alluding to are very much more complex. There are 
no solitary ants ; associations are the rule (we shall see there are 
one or two cases in which the association is with individuals of 
other species). In correlation with great proclivity to socialism 
we find an extraordinary increase in the variety of the forms of 
which species are made up. In addition to the male and female 
individuals of which the species of Insects usually consist, there 
are in ants workers of various kinds, and soldiers, all of which 
are modified infertile females. But in addition to the existence 
of these castes of infertile females, we find also numerous cases 
of variability or of dimorphism of the sexual individuals ; and 
this in both sexes, though more usually in the female. Thus 
there exists in ants an extraordinary variety in the polymorphism 
of forms, as shown by the table on p. 141, where several very 
peculiar conditions are recorded. 

The complex nature of these phenomena has only recently 

1 The parthenogenetic young produced by worker females are invariably of the 
male sex. 



140 HYMENOPTERA CHAP. 

become known, and as yet has been but little inquired into. The 
difference between the thoracic structure in the case of the winged 
rind wingless females of certain species (Fig. 55, and in vol. v. . 
tiir. .S-'39) is enormous, but in other species this difference appears 
to be much less. The ordinary distinctions between the queen- 
female and worker-females appear to be of two kinds; firstly, 
that the former is winged, the latter wingless ; 1 and secondly, that 
the former possesses a receptaculum seminis, the latter does not. 
In a few cases it would seem that the dimorphism of winged 
;ind wingless forms is not complete, but that variability exists. 
Intermediate conditions between the winged and wingless forms 
are necessarily rare; nevertheless a certain number have already 
been detected, and specimens of Lasius aliemts have been found 
with short wings. In rather numerous species some or all of 
the fertile females depart from the usual state and have no wings ; 
(a similar condition is seen, it will be recollected, in Mutillides 
and Thynnides of the neighbouring family Scoliidae). A di- 
morphism as regards wings also exists in the male sex, though it is 
only extremely rarely in ants that the males are wingless. Never- 
less a few species exist of which only wingless males have been 
found, and a few others in which both winged and wingless 
individuals of this sex are known to occur. The wingless males of 
course approach the ordinary workers ( == infertile wingless females) 
in appearance, but there is not at present any reason for 
supposing that they show any diminution in their male sexual 
characters. The distinction between workers and females as 
based on the existence or non-existence of a receptaculum seminis 
has only recently become known, and its importance cannot yet 
be estimated. The adult, sexually capable, though wingless forms, 
are called ergatoid, because they are similar to workers ('E^ar?;?, 
a worker). 2 

1 The student must recollect that the winged female ants cast their wings 
previously to assuming the social life. The winglessness of these females is a 
totally different phenomenon from that we here allude to. 

- See Fun 1, Verh. Ges. deutsch. Xalurf. Ixvi. 1894, 2, pp. 142-147 ; and Emery 
/>'/<>/. ( '< nt i'ii II, I. xiv. 1S<)1, p. 53. The term ergatoid applies to both sexes ; a species 
with worker-like female is ergatogynous ; with a worker-like male ergatandrous. 



IV 



ANTS 



141 



Table of the Chief Forms of Polymorphism in Ants. 



1-9 




v 







. 




M 
g 




| 


S 


""S ^ 




,. 


M 








* 


flj * 


^ S S 




o 


c . 




Cj <v* 




\*~ ^ 


^j ^ 


.5 3; T~ 


"S 


-- ^- ' 


Xame of Ants. 


^"5 


^ 


T s 


sl 


"S^^ 




" & c 

r- : ^ "~ 




a 


O a; 


i'l 




1 i^ 


t,' , t. 






'5 


>s 


^ 


% 


C +3 rt 


is 


o 


C J 




O 


a 


o 


H 


M ^ 


CO 


^ 


~ 


Miinaica, Polyrhachis, ~\ 


















etc. 


"*" 
















Camponotus, Atta,\ 


















Phcidologeton, etc. . J 


"*" 
















Pheidole, subg. Colo-} 


















bopsis . . . / 


















EC i t o n h a m atu m, "j 


















E. quadriglume, - 


+ 






+ 




+ 


+ 


+ 


E. foreli, etc. . 


















Cryptocerus disco- \ 


















ccplmlus, etc. . J 


+ 




+ 






+ 






Strongylognathus 


+ 




+ 






+ 






Carebara and Solen- ~j 


















opsis (except S. r 


+ 




+ 










+ 


geminata) . . | 


















Solenopsis geminata . 


+ 




+ 








+ 


+ 












f + 








Formica rufa . 


+ 




+ 




- excep- 


















( tionally 








Ponera punctatissima 


+ 


+ 


+ 








+ 




Ponera ergatandria . 


? 


+ 


+ 








+ 




Cardiocondyla cmeryi 


+ 




+ 


+ 






+ 




C. wroughtonii and ) 














, 




C. stambuloffi . . j 


















Formicoxentis niti- ~\ 














- 




dulus . . . J 




+ 


+ 












Tomognathus 


+ 






+ 






+ 




Odontomachus Jtae-} 








( 


. 




+ 




matodcs . 


+ 




+ 


' excep- 
[ tionally 










Polyergus . 


+ 




+ 


+ 










Dorylus, Anomma,} 














+ 


+ 


Eciton part. . 


















Aenictus . 


-f- 














+ 


Leptogenys, Dia-\ 














, 




camma . . .j 


















Myrmecocystus mel-} 














1 and 




ligcr, M. mexicanusj 














"j honey- 
\_ pots 


















f + 


f + 


Ponera eduardi . 


+ 




+ 








1 eyes 
[ large 


I'Vi-s 

[obsolete 


Ancr gates . 




+ 


+ 













In addition to the above there are apparently cases of females with post-meta- 
niorphic growth in Dorylides, but these have not yet been the subject of investiga- 
tion. 



14- HYMENOPTERA CHAI'. 

Much has been written about the mode in which the variety 
of forms of a single species of ant is produced. As to this there 
exists but little actual observation or experiment, and the 
subject has been much complicated by the anxiety of the writers 
to display the facts in a manner that will support some general 
theory. Dewitz was of opinion that workers and queens of ants 
were produced from different kinds of eggs. This view finds 
but little support among recent writers. Hart in recording the 
results of his observations on the parasol ant (of the genus Attct) 
one of the species in which polymorphism is greatest says 1 
that these observations prove that "ants can manufacture at 
will, male, female, soldier, worker or nurse," but he has not 
determined the method of production, and lie doubts it being 
" the character of the food." There is, however, a considerable 
1 Hjdy of evidence suggesting that the quality or quantity of the 
food, or both combined, are important factors in the treatment 
by which the differences are produced. The fact that the social 
Insects in which the phenomena of caste or polymorphism occur, 
though belonging to very diverse groups, all feed their young, is 
of itself very suggestive. When we add to this the fact that in 
ants, where the phenomena of polymorphism reach their highest 
complexity, the food is elaborated in their own organs by the 
feeders that administer it, it appears probable that the means 
of producing the diversity may be found herein. Wasmann has 
pointed out that the ants'-nest beetle, Lomechusa, takes much 
food from the ants, and itself destroys their young, and that in 
nests where Lomeclmsa is abundant a large percentage of erga- 
togynous forms of the ants are produced. He attributes this to 
the fact that the destruction of the larvae of the ant by the 
beetle brings into play the instinct of the ants, which seek to 
atone for the destruction by endeavouring to produce an 
increased number of fertile forms ; many ergatogynous individuals 
being the result. This may or may not be the case, but it is 
clear that the ants' instinct cannot operate without some 
material means, and his observation adds to the probability that 
this means is the food supply, modified either qualitatively or 
quantitatively. 

The existence of these polymorphic forms led Herbert 
Spencer to argue that the form of an animal is not absolutely 

1 Nature li. 1894, p. 12:.. 



iv ANTS 143 

determined by those " Aulau'en " or rudiments that Weismann 
and his school consider to be all important in determining the 
nature or form of the individual, for if this were the case, how 
can it be, he asked, that one egg may produce either a worker, 
nurse, soldier or female ant ? To this "Wasmann (who continued 
the discussion) replied by postulating the existence of double, triple 
or numerous rudiments in each egg, the treatment the egg receives 
merely determining which of these rudiments shall undergo de- 
velopment. 1 Forel seems to have adopted this explanation as being 
the most simple. The probability of Weismann's hypothesis being 
correct is much diminished by the fact that the limit between the 
castes is by no means absolute. In many species intermediate forms 
are common, and even in those in which the castes are believed to 
be quite distinct, intermediate forms occur as very rare excep- 
tions. 2 Emery accounts 3 for the polymorphism, without the 
assistance of the Weismannian hypothetical compound rudiments, 
by another set of assumptions ; viz. that the phenomenon has 
been gradually acquired by numerous species, and that we see it 
in various stages of development ; also that variation in nutrition 
does not affect all the parts of the body equally, but may be 
such as to carry on the development of certain portions of the 
organisation while that of other parts is arrested. Speaking 
broadly we may accept this view as consistent with what we 
know to be the case in other Insects, and with the phenomena of 
post-embryonic development in the class. But it must be ad- 
mitted that our knowledge is at present quite inadequate to 
justify the formulation of any final conclusions. 

The geological record of Formicidae is not quite what we 
should have expected. They are amongst the earliest Hymen- 
optera ; remains referred to the family have been found in the 
Lias of Switzerland and in the English Purbecks. In Tertiary 
times Formicidae appear to have been about the most abundant 
of all Insects. At Florissant they occur in thousands and form 
in individuals about one-fourth of all the Insects found there. 
They have also been met with numerously in. the European 
Tertiaries, and Mayr studied no less than 1500 specimens found 

1 B'wl. Ccntralbl. xv. 1895, p. 640. 

2 Prof. Forel has favoured the writer by informing him of several cases of these 
rare intermediate forms he has himself detected. 

3 Biol. CcntralU. xiv. 1894, p. 53. 



144 HYMENOPTERA CHAP. 

in amber. Formicides and Myrmicides are more abundant than 
Ponerides, but this latter group has the larger proportion of 
extinct genera ; conditions but little dissimilar to those existing 
at present. 

Classification of Ants. Ants are considered by many ento- 
mologists to form a series called Heterogyna. They can, 
however, be scarcely considered as more than a single family, 
Formicidae, so that the serial name is superfluous. Their 
nearest approach to other Aculeates is apparently made, by 
AmUyopone, to certain Mutillides (e.g. Apterogynct) and to the 
Thyiinides, two divisions of Scoliidae. Emery considers Dory- 
lidt'.s rather than Amblyoponides to be the most primitive form 
of ants, but we are disposed to consider Forel's view to the effect 
we have above mentioned as more probably correct. The point 
is, however, very doubtful. The condition of the peduncle is in 
both the sub-families we have mentioned very imperfect compared 
with that of other ants. Both these sub-families are of very 
small extent and very imperfectly known. We shall also 
follow Forel in adopting six sub-families, Camponotides, Dolicho- 
derides, Myrmicides, Ponerides, Dorylides, and Amblyoponides. 
Emery rejects the Amblyoponides as being merely a division of 
the Ponerides. This latter group displays the widest relations 
of all the sub-families, and may be looked on as a sort of central 
form. The Camponotides and Dolichoderides are closely allied, 
and represent the highest differentiation of the families in one 
direction. The Myrmicides are also highly differentiated, but 
are not allied to the Camponotides and Dolichoderides. 1 

Sub-Fam. 1. Camponotides. Hind body furnished with lni.t one 
constriction, so that only a single scale or node exists on the 
pedicel. Poison-sac forming a cushion of convolutions, on 
which is situate the modified sting, which forms merely an 
ejaculatory orifice for the poison. 

The members of this very extensive division of ants can be 
readily distinguished from all others, except the Dolichoderides, 
by the absence of a true sting, and by the peculiar form of the hind 
body ; this possesses only a single scale at the base, and has no 

1 Forel's latest views on this subject will be found in the Ann. Sac. cnt. Bdgique 
xxxvii. 1893, p. 161 ; the very valuable paper by Emery, in ZooL Jahrb. Syst. 
viii. 1896, p. 760. 



iv ANTS CAMPONOTIDES 145 

constriction at all on the oval, convex and compact mass of the 
abdomen behind this. The cloacal orifice is circular, not, as in 
other ants, transverse. These characters are accompanied by a 
difference in habits. The Camponotides, though they do not 
sting, produce poison in large quantity, and eject it to some dis- 
tance. Hence, if two specimens are confined in a tube they are 
apt to kill one another by the random discharges they make. 
Janet suggests that in order to neutralise the effect of this very 
acid poison, they may have some means of using, when they are 
in their natural abodes, the alkaline contents of a second gland 
with which they are provided. We shall mention the characters 
by which the Camponotides are distinguished from the small 
sub-family Dolichoderides when we deal with the latter. 

The sub -family includes 800 or more species. Camponotus 
itself is one of the most numerous in species of all the genera of 
Formicidae, and is distributed over most parts of the earth. We 
have no species of it in Britain, but in the south of Europe the 
Camponotus become very conspicuous, and may be seen almost 
everywhere stalking about, after the fashion of our British wood- 
ant, Formica n/fa, which in general appearance Camponotus much 
resembles. 

Until recently, the manner in which fresh nests of ants were 
founded was unknown. In established nests the queen-ant is 
fed and tended by the workers, and the care of the helpless larvae 
and pupae also devolves entirely on the workers, so that the 
queens are relieved of all functions except that of producing eggs. 
It seemed therefore impossible that a fresh nest could be estab- 
lished by a single female ant unless she were assisted by workers. 
The mode in which nests are founded has, however, been recently 
demonstrated by the observations of Lubbock, M'Cook, Adlerz, 
and more particularly by those of Blochmann, who was successful 
in observing the formation of new nests by Camponotus ligni- 
perdus at Heidelberg. He found under stones in the spring 
many examples of females, either solitary or accompanied only 
by a few eggs, larvae or pupae. Further, he was successful in 
getting isolated females to commence nesting in confinement, and 
observed that the ant that afterwards becomes the queen, at first 
carries out by herself all the duties of the nest : beginning by 
making a small burrow, she lays some eggs, and when these hatch, 
feeds and tends the larvae and pupae ; the first specimens of these 

VOL. VI L 



146 HYMENOPTERA 



CHAP. 



latter that become perfect Insects are workers of all sizes, and 
at once undertake the duties of tending the young and 
feeding the mother, who, being thus freed from the duties of 
nursing and of providing food while she is herself tended and 
fed, becomes a true queen-ant. Thus it seems established that 
in the case of this species the, division of labour found in the 
complex community, does not at first exist, but is correlative with 
increasing numbers of the society. Further observations as to the 
growth of one of these nascent communities, and the times and 
conditions under which the various forms of individuals composing 
a complete society first appear, would be of considerable interest. 

An American species of the same genus, C. pennsylvanicus, 
the carpenter-ant, establishes its nests in the stumps of trees. 
Leidy observed that solitary females constructed for themselves 
cells in the wood and closed the entrances, and that each one in 
its solitary confinement reared a small brood of larvae. The 
first young produced in this case are said to be of the dwarf 
caste, and it was thought by the observer that the ant remained 
not only without assistance but also without food during a period 
of some weeks, and this although she was herself giving food to 
the larvae she was rearing. 

Adlerz states that the females or young queens take no food 
while engaged in doing their early work, and that the large 
quantity of fat-body they possess enables them to undergo several 
months of hunger. In order to feed the young larvae they use 
their own eggs or even the younger larvae. It is to the small 
quantity of food rather than to its nature that he attributes the 
small size of the first brood of perfect workers. M. Janet : has 
recently designed an ingenious and simple apparatus for keeping 
ants in captivity. In one of these he placed a solitary female of 
Lasiii* (t/ic/ius, unaccompanied by any workers or other assistants. 
;tii(l he found at the end of 98 days that she was taking care of 
a progeny consisting of 50 eggs, 2 larvae, 5 pupae in cocoons, 5 
without cocoons. On the 102nd day workers began to emerge 
from the cocoons. 2 From these observations it is evident that 
the queen-ant, when she begins her nest, lives under conditions 
extremely different from those of the royal state she afterwards 
reaches. 

1 Ann. Sod. cut. France, 1893, p. 467. 
z Ann. Soc. cat. France, 1893, JJii/L p. cclxiv. 



JV 



ANTS CAMPONOTIDES 



147 



In many kinds of ants the full-grown individuals are known 
to feed not only the larvae by disgorging food from their own 
mouths into those of the little grubs, but also to feed one another. 
This has been repeatedly observed, and Forel made the fact the 
subject of experiment in the case of Camponotus ligniperdus. 
He took some specimens and shut them up without food for 
several days, and thereafter supplied some of them with honey, 
stained with Prussian blue ; being very hungry,- they fed so 
greedily on this that in a few hours their hind bodies were dis- 
tended to three times their previous size. He then took one of 
these gorged individuals and placed it amongst those that had 
not been fed. The replete ant w r as at once explored by the 
touches of the other ants and surrounded, and food was begged 
from it. It responded to the demands by feeding copiously a 
small specimen from its mouth : when this little one had received 
a good supply, it in turn communicated some thereof to other 
specimens, while the original well-fed one also supplied others, 
and thus the food was speedily distributed. This habit of receiv- 
ing and giving food is of the greatest importance in the life- 
history of ants, and appears to be the basis of some of the 
associations that, as w r e shall subsequently see, are formed with 
ants by numerous other Insects. 

OecopliyllcL smaragdina, a common ant in Eastern Asia, forms 
shelters on the leaves of trees by curling the edo;es of leaves 

t/ O O 

and joining them together. In doing this it makes use of an 
expedient that would not be believed had it not been testified by 
several competent and independent witnesses. The perfect ant 
has no material with which 
to fasten together the edges 
it curls ; its larva, how- 
ever, possesses glands that 
secrete a supply of material 
fur it to form a cocoon 
with, and the ants utilise 
the larvae to effect their 
purpose. Several of them 
combine to hold the foliage in the desired position, and while 
they do so, other ants come up, each one of which carries a 
larva in its jaws, applies the mouth of the larva to the parts 
where the cement is required, and makes it disgorge the sticky 




FIG. 60. Oecoplrylla smarciyiUna. Worker 
using a larva for spinning. 



148 HYMENOPTERA CHAP. 

material. Our figure is taken from a specimen (for which we 
are indebted to Mr. E. E. Green) that was captured in the act 
of bearing a larva. 

Formic tt ri'ftt, the Red-ant, Wood-ant, or Hill-ant, is in this 
country one of the best-known members of the Formicidae. It 
frequents woods, especially such as are composed, in whole or 
part, of conifers, where it forms large mounds of small sticks, 
straws, portions of leaves, and similar material. Although at 
first sight such a nest may appear to be a chaotic agglomeration, 
yet examination reveals that it is arranged so as to leave many 
spaces, and is penetrated by galleries ramifying throughout its 
structure. These mound-nests attain a considerable size when the 
operations of the industrious creatures are not interfered with, or 
their work destroyed, as it too often is, by ignorant or mischievous 
persons. They may reach a height of three feet or near it, and 
a diameter of twice that extent. The galleries by which the 
heaps are penetrated lead down to the earth below. From the 
mounds extend in various directions paths constantly traversed 
by the indefatigable ants. M'Cook observed such paths in the 
Trossachs ; they proceed towards the objects aimed at in lines 
so straight that he considers they must be the result of some 
sense of direction possessed by the ants ; as it is impossible to 
suppose they could perceive by the sense of sight the distant 
objects towards which the paths were directed : these objects in 
the case M'Cook describes were oak-trees up which the ants 
ascended in search of Aphides. 

M'Cook further observed that one of the oak-trees was reached 
by individuals from another nest, and that each of the two parties 
was limited to its own side of the tree, sentinels being placed on 
the limits to prevent the trespassing of an intruder ; he also 
noticed that the ants saw an object when the distance became 
reduced to about an inch and a half from them. This species is 
considered to be wanting in individual courage ; but when acting 
in combination of vast numbers it does so with intelligence and 
success. It does not make slaves, but it has been observed by 
Bignell and others that it sometimes recruits its numbers by 
kidnapping individuals from other colonies of its own species. 
Its nests are inhabited by forty or fifty species of guests of various 
kinds, but chiefly Insects. Another ant, Myrmica laevinodis, 
sometimes lives with it in perfect harmony, and Formicdxenus 



iv CAMPONOTIDES SLAVE-MAKING ANTS 149 

nilididus lives only with F. rufa. Amongst the most peculiar 
of its dependants we may mention large beetles of the genera 
Cetonia and Clytlira, which in their larval state live in the 
hills of the wood-ant. It is probable that they subsist on some 
of the vegetable matter of which the mounds are formed. Adlerz 
has given some attention to the division of labour amongst the 
different forms of the workers of ants, and says that in F. rufa it 
is only the bigger workers that carry building and other materials, 
the smaller individuals being specially occupied in the discovery 
of honey -dew and other Aphid products. In Camponotus it 
would appear, on the other hand, that the big individuals leave 
the heavy work to be performed by their smaller fellows. 

The wood-ant and its near allies have been, and indeed still 
are, a source of great difficulty to systematists on account of the 
variation that occurs in the same species, and because this differs 
according to locality. Our European F. rufa has been supposed 
to inhabit North America, and the interesting accounts pub- 
lished by M'Cook of the mound-making ant of the Alleghanies 
were considered to refer to it. This Insect, however, is not F. 
rufa, as WAS supposed by M'Cook, but F. exsectoides, Forel. It 
forms colonies of enormous extent, and including an almost in- 
credible number of individuals. In one district of about fifty 
acres there was an Ant City containing no less than 1*700 of 
these large ant-hills, each one teeming with life. It was found 
by transferring ants from one hill to another that no hostility 
whatever existed between the denizens of different hills ; the 
specimens placed on a strange hill entered it without the least 
hesitation. Its habits differ in some particulars from those of 
its European congener ; the North American Insect does not close 
the formicary at night, and the inquilines found in its nest are 
very different from those that live with F. rufa in Europe. 
Whether the typical wood-ant occurs in North America is 
doubtful, but there are races there that doubtless belong to 
the species. 

F. sanguinea is very similar in appearance to its commoner 
congener F. rufa, and is the only slave-making ant we possess in 
lUitain. This species constructs its galleries in banks, and is 
of very courageous character, carrying out its military operations 
with much tactical ability. It is perfectly able to live without 
the assistance of slaves, and very frequently does so : indeed it 



HYMENOPTERA CHAP. 



has been asserted that it is in our own islands (where, however, 
it is comparatively rare) less of a slave-owner than it is in 
Southern Europe, but this conclusion is very doubtful. It ap- 
pears when fighting to be rather desirous of conquering its 
opponents by inspiring terror and making them aware of its 
superiority than by killing them ; having gained a victory it 
will carry off the pupae from the nest it has conquered to its 
own abode, and the ants of the stranger-species that develop from 
these pupae serve the conquerors faithfully, and relieve them of 
much of their domestic duties. The species that F. sanguinea. 
utilises in this way in England are F. fusca, F. cunicularia, and 
possibly Lasius flatus. Huber and Forel have given graphic 
accounts of the expeditions of this soldier-ant. In the mixed 
colonies of F. sanguinea and F. fusca the slaves do most of the 
house-work, and are more skilful at it than their masters. Adlerz 
says that one of the slaves will accomplish twice as much work 
of excavation in the same time as the slave-owner ; these latter 
being lazy and fond of enjoyment, while the slaves are very 
industrious. 

Polyergus rv/escens, an European ant allied to Formica, is 
renowned since the time of Huber (1810) as the slave-making 
or Amazon ant. This creature is absolutely dependent on its 
auxiliaries for its existence, and will starve, it is said, in the 
midst of food unless its servitors are there to feed it. "Wasmann, 
however, states that PO////-/V///.S dues possess the power of 

feeding itself to a certain extent. Be 
this as it may, the qualities of this ant 
as warrior are superb. When an indi- 
vidual is fighting alone its audacity is 
splendid, and it will yield to no superi- 
ority of numbers ; when the creatures are 
acting as part of an army the individual 
boldness gives place to courage of a more 
suitable sort, the ants then exhibiting the 

FIG. 61. Head of Polyergws ae t of retreating or making flank move- 
nifcscens. (After Andre. ) J~, 

ments when necessary. It a Jrolyergus 

that is acting as a member of a troop finds itself isolated, and 
in danger of being overpowered, it has then no hesitation in 
seeking safety even by flight. This species is provided with 
mandibles of a peculiar nature : they are not armed with teeth, 




iv CAMPONOTIDES SLAVE-MAKING ANTS I 5 I 

but are pointed and curved ; they are therefore used after the 
manner of poignards, and when the ant attacks a foe it seizes 
the head between the points of these curved mandibles, and driv- 
ing them with great force into the brain instantly paralyses the 
victim. 

Mandibles of this shape are evidently unfitted for the purposes 
of general work, they can neither cut, crush, nor saw, and it is 
not impossible that in their peculiar shape is to be found the 
origin of the peculiar life of Polyergus : we find similar mandi- 
bles reappearing amongst the aberrant Dorylides, and attaining a 
maximum of development in the ferocious Eciton ; they also 
occur, or something like them, in a few aberrant Myrmicides ; 
and in the male sex of many other ants, which sex exercises no 
industrial arts, this sort of mandible is present. 

The ants that Polyergus usually attacks in order to procure slaves 
are Formica fusca and F.fusca, race auricularia ; after it has routed 
a colony of one of these species, P. rufescens pillages the nest and 
carries off pupae and some of the larger larvae to its own abode. 
When the captives thus deported assume the imago state, they 
are said to commence working just as if they were in their own 
houses among their brothers and sisters, and they tend their 
captors as faithfully as if these were their own relatives : possibly 
they do not recognise that they are in unnatural conditions, and 
may be quite as happy as if they had never been enslaved. The 
servitors are by no means deficient in courage, and if the place 
of their enforced abode should be attacked by other ant-enemies 
they defend it bravely. The fact that P. ru/escens does not feed 
its larvae has been considered evidence of moral degeneration, 
but it is quite possible that the Insect may be unable to do so 
on account of some deficiency in the mouth -parts, or other 
similar cause. The larvae of ants are fed by nutriment regurgi- 
tated from the crop of a \vorker (or female), and applied to the 
excessively minute mouth of the helpless grub : for so delicate 
a process to be successfully accomplished, it is evident that a 
highly elaborated and specialised arrangement of the mouth- 
parts must exist, and it is by no means improbable that the 
capacity of feeding its young in true ant-fashion is absent in 
Polyergus for purely mechanical reasons. 

M'Cook states that the North American ant, 7Wy/v///x />/< idus, 
which some entomologists consider to be merely a variety of 



152 



IIYMENOPTERA 



CHAP. 



the European species, makes slaves of Formica scJiaufussi, itself 
does no work, and partakes of food only when fed by its servi- 
tors. He did not, however, actually witness the process of feed- 
ing. AVhen a migration takes place the servitors deport both 
the males and females of P. hi.cidus. M'Cook adds that the 
servitors appear to be really mistresses of the situation, though 
they avail themselves of their power only by working for the 
advantage of the other species. 

The honey-ant of the United States and Mexico has been in- 
vestigated by M'Cook and others; the chief peculiarity of the 

species is that certain individuals are 
charged with a, sort of honey till they 
become enormously distended, and in fact 
serve as leather bottles for the storage 
of the fluid. The species Myrmecocyslus 
liortideorum and M. mclliger, are mode- 
rate-sized Insects of subterranean habits, 
the entrance to the nest of M. 1 1 or tide or u in 
being placed in a small raised mound. 
The honey is the product of a small gall 
found on oak leaves, and is obtained by 
i- the worker-ants during nocturnal expedi- 
fooiis, f rom which they return much dis- 

v 

tended ; they feed such workers left at 
home as may be hungry, and then apparently communicate the 
remainder of the sweet stuff they have brought back to already 
partly charged ; ' honey-bearers " left in the nest. The details 
of the process have not been observed, but the result is that the 
abdomens of the bearers become dis- 
tended to an enormous extent (Figs. 
62, 63), and the creatures move but 
little, and remain suspended to the 
roof of a special chamber. It is 
considered by M'Cook that these 
living honey-tubs preserve the fond 
till a time when it is required for 
the purposes of feeding the com- 
munity. The distension is pro- 
duced entirely by the overcharging 
of the honey-crop, the other contents of the abdomen being 




FIG. 62. J/y /-//. * 

m ii nn. Honey - pot ant, 
dorsal view. 




Flu. 63. Mi/rmecocystus 
Lateral view. 



iv ANTS CAMPONOTIDES 153 

forced by the distention to the posterior part of the body. 
Lubbock has since described an Australian ant, Melophorus 
i njiatus, having a similar peculiar habit, but belonging to the 
allied tribe Plagiolepisii. Quite recently a South African honey- 
tub ant belonging to the distinct genus Plagiolepis (Ptrimeni 
For.) has been discovered, affording a proof that an extremely 
specialised habit may arise independently of relation between the 
Insects, and in very different parts of the world. 

Species of the genus Lasius are amongst the most abundant 
of the ant-tribe in Britain. They are remarkable for their con- 
structive powers. L. niger, the common little black garden-ant, 
forms extensive subterranean galleries, and is extremely successful 
in the cultivation of various forms of Aphidae, from the products 
of which the species derives a large part of its subsistence. The 
ants even transport the Aphidae to suitable situations, and thus 
increase their stock of this sugary kind of cattle, and are said to 
take the eggs into their own dwellings in the autumn so that 
these minute and fragile objects may be kept safe from the 
storms and rigours of winter. These little creatures are brave, 
but when attacked by other ants they defend themselves chiefly 
by staying in their extensive subterranean galleries, and blocking 
up and securing these against their assailants. 

Z. fuliginosus, another of our British species, has very different 
habits, preferring old trees and stumps for its habitation ; in the 
hollows of these it forms dwellings of a sort of card ; this it 
makes from the mixture of the secretions of its salivary glands 
with comminuted fragments of wood, after the fashion of wasps. 
It is a moderate-sized ant, much larger than the little L. niger, 
and is of a black colour and remarkably shining ; it gives off 
a very strong but by no means disagreeable odour, and may be 
seen on the hollow trees it frequents, stalking about in large 
numbers in a slow and aimless manner that contrasts strikingly 
with the active, bustling movements of so many of its congeners. 
When this species finds suitable trees near one another, a colony 
is established in each ; the number of individuals thus associated 
becomes very large, and as the different colonies keep up inter- 
communication, this habit is very useful for purposes of defence. 
Forel relates that he once brought a very large number of 
Formica pratensis and liberated them at the base of a tree in 
which was a nest of L. fuliginosus ; these latter, finding them- 



154 HYMENOPTERA CHAP. 

selves thus assaulted and besieged, communicated in some way, 
information of the fact to the neighbouring colonies, and Forel 
soon saw large columns of the black creatures issuing from the 
trees near by and coming with their measured paces to the 
assistance of their confreres, so that the invaders were soon dis- 
comfited and destroyed. Although the European and North 
American representatives of the sub-family Camponotides live 
together in assemblies comprising as a rule a great number of 
individuals, and although the separate nests or formicaries which 
have their origin from the natural increase of a single original 
nest keep up by some means a connection between the members, 
and so form a colony of nests whose inhabitants live together on 
amicable terms, yet there is no definite information as to how 
long such association lasts, as to what is the nature of the ties 
that connect the members of the different nests, nor as to the 
means by which the colonies become dissociated. It is known that 
individual nests last a long time. Charles Darwin has mentioned 
in a letter to Forel that an old man of eighty told him he had 
noticed one very large nest of Formica n/fx in the same place 
ever since he was a boy. But what period they usually endure 
is not known, and all these points probably vary greatly according 
to the species concerned. It has been well ascertained that when 
some ants find their nests, for some unknown reason, to be unsuit- 
able the inhabitants leave their abodes, carrying with them their 
young and immature forms, and being accompanied or followed by 
the various parasites or commensals that are living with them. 
Wasmann and other entomologists have observed that the ants 
carry bodily some of their favourite beetle-companions, as well 
as members of their own species. Forel observed that after 
a nest of Formica pratensis had been separated into two nests 
placed at a considerable distance from one another so as to have 
no intercommunication, the members yet recognised one another 
as parts of the same family after the lapse of more than a month ; 
but another observation showed that after some years of separation 
they were no longer so recognised. 

Although it is now well ascertained that ants are able to 
distinguish the individuals belonging to their own nests and 
colonies from those that, though of their own species, are not so 
related to them, yet it is not known by what means the recogni- 
tion is effected ; there is, however, some reason to suppose that it 



IV 



ANTS CAMPONOTIDES 



155 



is by something of the nature of odour. One observer has noticed 
that if an ant fall into water it is on emerging at first treated as 
if it were a stranger by its own friends ; but other naturalists have 
found this not to be the case in other species. Contact with 
corrosive sublimate deprives ants for a time of this power of 
recognising friends, and under its infhience they attack one 
another in the most ferocious manner. 

The nests and colonies of the species of Camponotides we 
have considered are all constructed by societies comprising a 
great number of indi- 
viduals ; there are, 
however, in the tropics 
numerous species that 
form their nests on 
foliage, and some of 
these contain only a 
few individuals. The 
leaf -nests (Fig. 64) of 
certain species of Poly- 
/'//acltis are said to be 
formed of a paper-like 
material, and to con- 
tain each a female and 
about 8 or 10 worker 
ants. Forel : has ex- 
amined nests of several 
Indian species, and finds they differ from those of other ants in 
consisting of a single cavity, lined with silk like that of a spider. 
These nests are further said to be constructed so as to render 
them either inconspicuous or like other objects on the" leaves ; 
P. argentea covers its small dwelling with little bits of vegetable 
matter, and a nest of P. rastdla was placed between two leaves 
in such a manner as to be entirely hidden. All the species of 
the genus do not, however, share these habits, P. mayri making 
a card-nest, like Dolichoderus and some other ants. The species 
of the genus Polyrhachis are numerous in the tropics of the Old 
World. 

Forbes noticed that a species of this genus, that makes its 
paper-like nest on the underside of bamboo-leaves produces a noise 
1 Forel, J. Bombay Soe. viii. 1893, p. 36. 




FIG. 64. Nest of Polyrhachis sp. (After Smith.) 



HYMENOPTERA 



CHAP. 



by striking the leaf with its head in a series of spasmodic taps. 
The same observer has recorded a still more interesting fact in 
the case of another species of this genus a large brown ant- 
found in Sumatra. The individuals were " spread over a space, 
perhaps a couple of yards in diameter, on the stem, leaves, and 
branches of a great tree which had fallen, and not within sight 

of each other ; yet the tapping. was 
set up at the same moment, con- 
tinued exactly the same space of 
time, and stopped at the same in- 
stant ; after the lapse of a few 
seconds all recommenced at the 
same instant. The interval was 
always of about the same duration, 
though I did not time it ; 
each ant did not, however, beat 
synchronously with every other 
in the congeries nearest to me ; 
there were independent tappings, 
so that a sort of tune was played, 
each congeries dotting out its 
own music, yet the beginnings 
and endings of the musical parties 
were strictly synchronous." 
Mr. Peal has also recorded that an ant the name is not 
mentioned, but it may be presumed to be an Assamese species- 
makes a concerted noise loud enough to be heard by a human 
being at twenty or thirty feet distance, the sound being produced 
by each ant scraping the horny apex of the abdomen three times 
in rapid succession on the dry, crisp leaves of which the nest is 
usually composed. These records suggest that these foliage-ants 
keep up a connection between the members of different nests 
somewhat after the same fashion as do so many of the terrestrial 
Camponotides. Although the species of Cainponotides have no 
special organ for the production of sound in the position in which 
one is found in Mvrmicides and Ponerides, yet it is probable that 

v > t/ x 

they are able to produce a sound by rubbing together other parts . 
of the abdomen. 




FIG. 65. P<->h/,-/u:i-//i.i JHI, nl ii 
worker. Singapore. 



IV 



ANTS 



157 



Sub-Fam. 2. Dolichoderides. Hind //^/y furnished u-ith lut 
one constriction so that only a single scale or node is formed ; 
tiling rudiincntari/ ; the poison-sac without cushion. 

The Dolichoderides are similar to Camponotides in appearance, 
and are distinguished chiefly by the structure of the sting and the 
poison apparatus. To this we may add that Forel also considers 
the gizzard to be different in the two sub-families, there being no 
visible calyx in the Dolichoderides, while this part is largely 
developed in the Camponotides. This is one of the least extensive 
of the sub-families of ants, not more than 150 species being- 
yet discovered. Comparatively little is 
known of the natural history of its 
members, only a very small number of 
species of Dolichoderides being found in 
Europe. The best known of these (and 
the only British Dolichoderid) is Tapi- 
noma erraticum, a little ant of about the 
size of Lasius niger, and somewhat similar 
in appearance, but very different in its 
habits. T. erraticum does not cultivate 
or appreciate Aphides, ' but is chiefly 
carnivorous in its tastes. Our knowledge 
of it is due to Forel, who has noticed that 
it is very fond of attending the fights 
between other ants. Here it plays the 
part of an interested spectator, and 
watching its opportunity drags off the 
dead body of one of the combatants in 
order to use it as food. Although desti- 
tute of all power of stinging, this Insect 
has a very useful means of defence FlG - 66 - fapinoma en-ati- 

. . . cum, worker. Britain. 

m the anal glands with which it is Upper side and profile, 
provided ; these secrete a fluid having 

a strong characteristic odour, and possessing apparently very 
noxious qualities when applied to other ants. The Tn/ii- 
noma has no power of ejecting the fluid to a distance, but is 
very skilful in placing this odorous matter on the body of an 
opponent by touching the latter with the tip of the abdomen ; 
on this being done its adversary is usually discomfited. This 




i;8 HYMENOPTERA 



CHAP. 



Insect is subterranean in its habits, and is said to change its 
abode very frequently. T. erraticum occurs somewhat rarely in 
Britain. Forel has also noted the habits of Liometopum micro- 
cephalum, another small European species of Dolichoderides. It 
is a tree-ant, and by preference adopts, and adapts for its use, the 
1 mrrows made by wood-boring beetles. It forms extremely populous 
colonies which may extend over several large trees, the inhabitants 
keeping up intercommunication by means of numerous workers. 
Xo less than twelve mighty oaks were found to be thus united 
into a colony of this ant in one of the Bulgarian forests. The 
species is very warlike, and compensates for the extreme minute- 
ness of its individuals by the skilful and rapid rushes made by 
combined numbers on their ant-foes of larger size. 

Fritz M tiller has given a brief account, under the name of 
the Imbauba ant, of a Brazilian- arboreal ant, that forms small 
nests in the interior of plants. The species referred to is no 
doubt an Azteca, and either A. instabilis, or A. mulleri. The nests 
are founded by fertilised females which may frequently be found 
in the cells on young Cecropia plants. Each internode, he says, 
has on the outside, near its upper part, a small pit where the wall 
is much thinner, and in this the female makes a hole by which 
she enters. Soon afterwards the hole is completely closed by a 
luxuriant excrescence from its margins, and it remains thus closed 
until about a dozen workers have developed from the eggs of 
the female, when the hole is opened anew from within by the 
workers. It is said that many of the larvae of these ants are 
devoured by the grubs of a parasite of the family Chalcididae. 
Tbis Insect is thought to protect the plant from the attacks of 
leaf-cutting ants of the genus Atfn. 

We may here briefly remark that much has been written 
about the benefits conferred on plants by the protection given to 
them in various ways by ants : but there is reason to suppose 
that a critical view of the subject will not support the idea of 
the association being of supreme importance to the trees. 1 

Sub-Fam. 2. Myrmicides. Pedicel <>f ahdomcn formed of two 
in 11- marked nodes (knot-like segments*). Sting present (absent 
i a Hie Cryptocerini and Attini'). (It should be noted that the 

1 See von Iliering, Berlin, cut. Zcitschr. xxxix. 1894, p. 364 ; and Forel, Ann. 
Soc. cut. /:./,/;,{//,, \\. 1896, p. 170. 



IV 



ANTS MYRMTCIDES 



159 



workers of the genera Eciton and Aenictus of the sub-family 
Dorylides have, like the Myrmicides, tivo nodes in the 




This sub-family consists of about 1000 species, and includes 
a great variety of forms, but, as they are 
most of them of small size, they are less 
known than the Camponotides, and much 
less attention has been paid to their 
habits and intelligence. Forel, until re- 
cently, adopted four groups : Myrmiciui, 
Attini, Pseudomyrmini and Cryptocerini ; 
but he is now disposed to increase this 
number to eight. 1 They are distinguished 
by differences in the clypeus, and in the 
form of the head ; but it must be noted 
that the characters by which the groups , 

J FIG. 6/. Pheulologeton labo- 

are defined are not in all cases fully riosus, large and small 
applicable to the males. The Crypto- workers> East Iudia - 
cerini are in external structure the most highly modified of 
Hymenoptera, if not of all the tribes of Insecta. 

i. The MYRMICINI proper are defined by Forel as having the 
antennae inserted near the middle, a little behind the front, of 

the head, which has carinae on the inner 
sides, but none on the outer sides, of the 
insertions of the antennae ; the clypeus ex- 
tends between the antennae. 

Certain genera of small European ants of 
the group Myrmicini display some most 
anomalous phenomena. This is especially 
the case in Formicoxenus, Anergates and 
Tomognathus. The facts known have, how- 
ever, been most of them only recently dis- 
covered, and some obscurity still exists as to 
many of even the more important points 
i u these extraordinary life-histories. 

it has long lieen known that the little 
fonmcoxenus mtiduius lives as a guest 
in the nests of Formica rufa, the wood -ant; and another 
similar ant, Xtenamma westiuoodi, which shares the same life, 




FIG. 68. Formicoxenus 
mtiduius, male. (After 

Adlerz. ) 



Ann. Soc. cut. Belgique, xxxvii. 1893, p. 163. 



i6o 



HVMENOPTERA 



CHAP. 



was declared by Nylander and Smith to be its male ; it was 
however shown some years ago by Andre that this is a mistake, 
and that S. westwoodi is really the male of another ant that 
had till then been called Asemorhoptrum lippulinn. This correc- 
tion left the workers and females of Formicoxenus nitidulus 
destitute of a male, but Acllerz has recently discovered that the 
male of this species is wingless and similar to the worker, the 
female being a winged Insect as usual. It is very curious that 
the characters by which the male is distinguished from the 
worker should vary in this species ; but according to Adlerz this 
is the case, individuals intermediate in several points between 
the males and workers having been discovered. This pheno- 
menon of quite wingless males in species where the female is 
winged is most exceptional, and is extremely rare in Insects ; but 
it occurs, as we shall see, in one or two other Myrmicides. Charles 

Darwin made the very 
reasonable suggestion 
that winged males may 
be developed occasion- 
ally as an exceptional 
phenomenon, and it is 
very probable that this 
may be the case, though 
it has not yet been 
demonstrated. Formi- 
coxenus nitidulus occurs 
in England in the nests 
of Formica rufa and of 
F. congerens, but we 
are not aware that the 
male has ever been 

Fia.GQ.Anergatesatratulus. Europe. A. male, with found ill this country, 
part of hind leg broken off ; B, female, with wings.: C, T , .., Awernnt^ is 
female, after casting the wings and becoming a queen. 

allied to Formicoxenus, 

and occurs in Central Europe, but has not been found in Britain ; 
the female, as in Formicoxenus, is winged and the male wingless, 
but there is no worker-caste ; the male is a rather helpless creature, 
and incapable of leaving the nest. The species lives in company 
with Tetru'inoritnn cawjtitintt, a little ant very like M//n/t!c</, and not 
uncommon in South-East England. The female Anergates is at 




I 



IV 



M VRMICIDES ANERGA TES I 6 I 



first an active little creature with wings, but after these are lost 
the body of the Insect becomes extremely distended as shown in 
Fig. 69, C; the creature is in this state entirely helpless, and as 
there are no workers, the Anergates, is completely dependent, for 
the existence of itself and its larvae, on the friendly offices of the 
Tetramorium that lives with it. The mode of the association 
of these two Insects is at present both unparalleled and inexplic- 
able, for only workers of the Tetramorium are found in company 
with the and 5 Anergates ; the community, in fact, consisting 
of males and females of one species and workers of another. The 
nests of Anergates are so rare that only a few naturalists have 
been able to observe them (Schenk, von Hagens, and Forel may 
be specially mentioned), but in the spots where they occur, 
nests of the Tetramorium, containing all the forms of that species, 
are numerous, and it therefore seems probable that a young fertile 
female of the Anergates may leave a nest in which it was born, 
enter a nest of the Tetramorium, and, destroying the queen thereof, 
substitute herself in the place of the victim ; but if this be really 
the case, the larvae and pupae of the Tetramorium must also be 
destroyed, for no young of the Tetramorium are ever found in 
these strange associations. It is very difficult to believe that 
the Tetramorium workers should be willing to accept as their 
queen a creature that commenced her acquaintance with them by 
destroying their own queen or queens and a number of their 
young sisters ; especially as the Tetramorium is a more powerful 
ant than the Anergates, and could readily dispose of the murderous 
intruder if it were disposed to do so. It is known, however, 
that colonies of Tetramorium completely destitute of queens some- 
times occur, and Wasmann has suggested that the female Aner- 
gates may seek out one of these, and installing herself therein as 
a substitute, may be accepted by the orphaned colony. This 
plausible hypothesis has still to be verified. 

The genus Cardiocondyla also exhibits the phenomenon of 
apterous, worker-like males, while in one species, C. emeryi, a 
winged male is also known to exist. 

Tomognathus sublaevis is a little Myrmicid ant, found rarely 
in Denmark and Sweden, where its habits have recently been 
studied by Adlerz. A band of the Tomognathus attack the nest 
of another little Myrmicid, Leptothorax acervorv. m , and succeed 
by their own pertinacity and the fears of the Leptothorax in 

VOL. VI M 



1 62 HYMENOPTERA 



CHAP. 



obtaining possession of it ; the legitimate owners disappear, leav- 
ing the Tomognathus in possession of their larvae and pupae ; 
these complete their development only to find themselves the 
slaves of Tomognathus. The subsequent relations of the two 
ants are friendly, the slaves even preventing their masters from 
wandering from the nest when they wish to do so. If an estab- 
lished mixed community of this nature is in want of additional 
servitors, the Tomognatlius secure a supply by raids after the 
fashion of the Amazon-ant, bringing back to their abode larvae 
and pupae of Leptothoraoc to be developed as slaves. It was for- 
merly supposed that the Tomognathus continued its species by 
perpetual parthenogenesis of the workers, for neither males nor 
females could be found. Adlerz l has, however, now discovered the 
sexual individuals. The male is an ordinary winged ant, and is 
so like that sex of the Lcptoilwrax, that Adlerz had failed to 
distinguish the two before he reared them. The females are 
apterous, and in fact like the workers. It would perhaps be 
more correct to say that the workers of this species vary greatly 
but never become winged ; some of them have ocelli and a struc- 
ture of the thorax more or less similar to that of winged females, 
though none have been found with wings. Certain of these 
females possess a receptaculum seminis, and Adlerz treats this as 
the true distinction between female and worker. In accordance 
with this view the female of Tomognatli us may be described as a 
worker -like individual possessing a receptaculum seminis, and 
having more or less of the external structures of winged females, 
though never being actually winged. It is probable that other 
workers reproduce parthenogenetically. The males of this species 
will not unite with females from the same nest, thus differing 
from many other ants, in which union between individuals of the 
same nest is the rule. Finally, to complete this curious history, 
we should remark that the larvae of the Tomot/nathus are so 
similar to those of the Leptothorax that Adlerz is quite unable to 
distinguish the two. 

Stronyylognatlius testaceusand S. kuhcri live in association with 
Tetramorium caespit-um, and are cared for by these latter ants; it 
is notable that as in the case of the slave-making Polyergus 
rufescens the mandibles of the Strongylognathus are cylindrical 
and pointed, and I her* 'tore unsuitable for industrial occupations. 

1 Jiili. Hvenska Ale. x.\i. 1896, Afd. iv. Xo. -1. 



iv ANTS MYRMICIDES 163 

S. testaceus is a weak little ant, and lives in small numbers in the 
nests with T. caespitum, which it is said to greatly resemble in 
appearance. The proportions of the forms of the two species 
usually associated is peculiar, there being a great many workers 
of T. caespitum both in the perfect and pupal states, and also all 
the sexes of the Strongylognathus, of which, however, only a few 
are workers. This would seem to suggest that S. testaceus attacks 
and pillages the nests of T. caespitum in order to carry off worker 
pupae, just as Polyergus ru/escens does. But the facts that S. 
testaceus is a weaker Insect than the Tetramorium, and that only 
a few of its worker-caste are present in a community where there 
are many workers of the Tetramorium, seem to negative the view 
that the latter were captured by the former ; and the mode in 
which the associated communities of these two species are started 
and kept up is still therefore in need of explanation. 

Strongylognathus huberi is a much stronger Insect than its 
congener, S. testaceus, and Forel has witnessed its attack on 
T/'framorium caespitum. Here the raid is made in a similar 
manner to that of Polyergus ru/escens on Formica ; the Tctramo- 
r in in is attacked, and its pupae carried off to the abode of the 
Strongylognathus to serve in due time as its slaves. The man- 
dibles of S. huberi, being similar in form to those of Polyergus 
ru/escens, are used in a similar manner. 

Although T. caespitum is common enough in South-East Eng- 
land, it is to be regretted that none of the guests or associates we 
have mentioned in connection with it occur in this country. It 
is a most variable species, and is distributed over a large part of 
the globe. 

Our British species of Myrmicides, about ten in number, 
all belong to the group Myrmicini ; none of them are generally 
common except Murmica rubra, which is a most abundant Insect, 
and forms numerous races that have been considered by some 
entomologists to be distinct species ; the two most abundant of 
these races are M. ruginodis and M. scabrinodis, which sometimes, 
at the time of the appearance of the winged individuals, form vast 
swarms. 

The tiny Mb-nomorium pharaonis is a species that has 
been introduced into Britain, but now occurs in houses in 
certain towns ; it sometimes accumulates on provisions in such 
numbers as to be a serious nuisance. Seventeen thousand 



164 



HYMENOPTERA 



CHAP. 



individuals weigh 1 gramme, and it is probable that a nest 

may include millions of specimens. 

The genus Aphaenogaster^ and its immediate- allies include the 

harvesting ants of Europe and North America : they form subter- 

ranean nests consisting of iso- 
lated chambers connected by 
galleries ; some of the chambers 
are used as store -houses or 
granaries, considerable quantities 
of corn, grass, and other seeds 
being placed in them. A. 
structor and A. lar/i/i/'//* have 
been observed to do this in 
Southern Europe by Lespes, 
Moggridge, and others. 

In the deserts about Algeria 
and Tunis a harvesting ant, 
Aphaenogaster (Messor) aren- 
arius, is an important creature : 
its subterranean dwellings are 
very extensive, and are placed 
at a depth of several feet from 
the surface. Entrance to these 
dwellings is obtained by small 

holes, which are the orifices of 
Flu. 70. Aphaenogaster (Messor) barbarus. , , , 

Algeria. A, male ; B, winged female : galleries many feet in length : 




C, large worker or soldier; D, small 
worker, x -I}. 



the holes are SUlTOimded by 

. . 

pellets ot sand projecting some- 
what above the general surface, and consequently making the 
places conspicuous. The subterranean works occupy an area of 
iifty or a hundred square yards excavated at a depth of three to 
six feet. In these immense nests there exists a form of worker, 
of very small size, that never conies to the surface. 2 

Pogonomyrmex l><irl><itns and other species have been observed 
to do harvesting in North America. After the workers of P. 
!>/ 1, -I in Ins have taken the seeds into the nest they separate the 
husks and carry them out, depositing them on a heap or kitchen- 

1 Until I'crcully this genus was generally known as Atta, but this name is now 
applied to the leai'-futting ants, that were formerly called Oeeodoma. 

2 Foivl, /;,///. Soc. Vaudoise, xxx. pp. 29-30, 1S91. 



IV 



ANTS MYRMICIDES 165 



midden, formed near by. M'Cook has witnessed and described 
the process of stripping the seeds. 

Certain genera e.g. Aphaenogaster, Plieidole exhibit great 
disparity in the forms of the workers, some of which are of 
size much superior to the others, and possess disproportionately 
large heads ; these large individuals are found in the same nest 
as the smaller forms. All the intermediate forms may frequently 
be found, and at the same time, in the genus Apliaenogaster ; but 
in Pheidole intermediates are of the utmost rarity. 

The genus Cremastogaster is remarkable on account of the 
shape of the hind body and its articulation, which give the 
abdomen the appearance of being put on upside down. This 
mode of articulation may allow the Insect to threaten its enemies 




X 

A B 

Fio. 71.' Oremastogaster tricolor, worker. A, with abdomen extended ; B, uplifted. 

when they are in front of it ; but it is doubtful whether the 
Cremastogaster possesses an effective sting. 

ii. The group ATTINI is distinguished by the presence of a 
carina near the eye, by the antennae being inserted at a moderate 
distance from one another, by the clypeus being prolonged back- 
wards between them ; and by the absence of a sting. The 
group is not represented in Europe, but in Tropical America the 
ants belonging to it are amongst the most important of natural 
objects. The species of the genus Atta (usually styled Oecodomct) 
are the formidable leaf-cutting ants of America. They occur in 
enormous colonies in certain places, and will in a short time 
completely strip a tree of its leaves. As they appear to prefer 
trees of a useful kind, especially those planted by man, their 
ravages are often of the most serious nature. The natives, 
feeling it hopeless to contend with these Insect hordes, only too 
frequently abandon all attempts to cultivate the trees and 
vegetables the Insects are fond of. Both Bates and Belt have 



1 66 HYMENOPTERA 



CHAP. 



given accounts of some points in the economy of these ants. 
They are amongst the largest of the Formicidae, the females in 
some cases measuring about two and a half inches across their 
expanded wings ; the males are much smaller, but are less dis- 
similar to their partners than is usual among ants. The workers, 
on the other hand, are so extremely different, that no one would 
suppose them to he at all related to the males and females (see 
vol. v. Fig. 339). 

The mode of operation of these ants is to form paths from 
their formicary extending for a considerable distance in various 
directions, so that they have a ready access to any spot in a 
district of considerable extent ; when a tree or bush is found 
bearing leaves suitable for their purposes, the worker ants ascend 
it in large numbers and cut up the leaves by biting out of them 
pieces similar in size and shape to a small coin ; these pieces are 
then carried back in the jaws of the ants to their nests ; the ant- 
paths are therefore constantly traversed by bands of little creatures 
carrying burdens homewards, or hurrying outwards in search of 
suitable trees. 

The formicaries are of considerable size, and are described 
as consisting of low mounds of bare earth of considerable extent. 
Bates speaks of as great a circumference as forty yards ; these 
accumulations of earth have frequently an appearance different 
from the adjoining soil, owing to their being formed of subsoil 
brought up from below ; they are kept bare by the ants con- 
stantly bringing to and depositing on the surface fresh material 
resulting from their subterranean excavations. The true abodes, 
beneath the earth, are of greater extent than the mounds them- 
selves, and extend to a considerable depth ; they consist of 
chambers connected by galleries. 

The leaf -cutting ants extend their range to North 
America, and M'Cook has recently called attention to a case 
there in which A. fen-ens made an underground route at an 
average depth of 18 inches, and at an occasional depth oT 
feet, extending 448 feet entirely beneath the earth, after which 
it was continued for 185 feet to reach a tree which the ants were 
engaged in defoliating. This route, extending altogether to a length 
of more than GOO feet, presented only a very slight deviation 
from a straight line drawn between the point of departure and 
the object to be attained. By what sense this ant was enabled 



IV 



MYRMICIDES LEAF-CUTTING ANTS I 6/ 



to make a subterranean tunnel in a straight line to a desired 
object situated at so great a distance, we know not. 

The use the leaf-cutting ants make of the enormous amount 
of material they gather was for long a subject of debate, and has 
only recently been ascertained by the observations of Moller. 
After being carried to the nest the pieces of leaves are cut into 
small fragments by another set of workers and formed into balls, 
which are packed in various parts of the nest, and amongst 
which the mycelium of a fungus Rozites gongylophora -ramifies. 
This fungus the ants cultivate in the most skilful manner : 
they manage to keep it clear from mouldiness and bacterial 
agents, and to make it produce a modified form of growth in 
the shape of little white masses, each one formed by an agglomera- 
tion of swellings of the mycelium. These form the chief food of 
the colony. Moller ascertained by experiment that the results 
were due to a true cultivation on the part of the ants : when 
they were taken away from the nests, the mycelium produced 
two kinds of conidia instead of the ant-food. 

Many details of the economy of these leaf-cutting ants are 
still very imperfectly known. The large-headed forms, called 
soldiers, have been the subject of contradictory statements ; 
Bates having concluded from his own observations that they 
are harmless, while Mr. J. H. Hart assures us that they are very 
fierce and vindictive, and inflict very serious wounds by biting 
(the Attini do not sting). We anticipate that the observations 
of both these naturalists will prove to be substantially correct, 
and that the differences in habits will be found to be owing to 
distinctions in the conditions of the community. In connection 
with this point we may remark that the function of the ex- 
cessively large heads of certain kinds of soldier-ants is still 
obscure. In the East Indian Pheidologeton diversus the big- 
soldiers are quite one hundred times as large as the smaller 
workers. As these latter bite viciously it would naturally be 
supposed that their gigantic confreres with enormous heads would 
be warriors of a most formidable nature ; but, as a matter of fact, 
the giants are unable to bite even when they try to do so. 
Aitken has somewhere suggested that these enormous individuals 
play the part of state elephants ; and we have been informed by 
Colonel Bingham that the small ants may frequently be seen 
riding in numbers on their unwieldy fellows. We anticipate 



1 68 HYMENOPTERA CHAP. 

however, that some other function will be found to exist for 
these forms with enormous heads. An examination of their 
organs of sense and of voice is very desirable. 

Details of the modes in which the great communities of the 
leaf- cutting Attidae are maintained, are still wanting. The 
females do not, we have been informed by Mr. Hart, possess 
any considerable powers of aftergrowth, so that there is no reason 
to suppose them to be unusually prolific. At certain seasons 
great swarms of winged individuals are produced, and after 
leaving the nests pair in the manner of our European Myrmica. 
Possibly the females may, after losing their wings, again enter 
the large communities. Yon Ihering states that the workers of 

o o 

Atta lundi are fertile. 

iii. The group PSEUDOMYRMIXI includes the genera Pscudo- 
ini/riiia and Himn, which are by some entomologists treated as 
but a single genus. The antennae are inserted near together on the 
front of the head ; there is no carina on the head external to 
their insertion, and the clypeus does not extend forwards between 
them. The Insects are usually of elongate form, possess a sting, 
and have a naked pupa. The group occurs in both hemispheres, 
but is exclusively exotic, and but little is known of the habits of 
its members. Forel has recently observed that numerous species 
live inside dried stems of grass or in hollow twigs, and are 
beautifully adapted for this mode of life by their elongate form, 
some of them being as slender as needles. Some interesting 
observations have been made in Nicaragua by Belt on Pscudo- 
mynna bicolor and its relations with an acacia-tree, in the thorns 
of which it lives. The acacia in question is called the bull's-horn 
thorn, because the branches and trunk are armed with strong 
curved spines, set in pairs, and much resembling the horns of 
the quadruped whose name they bear. The ant takes possession 
of a thorn by boring a small hole near the distal extremity, and 
forms its nest inside. The leaves of this plant are provided 
with glands that secrete a honey-like fluid, which it appears 
forms the chief, if not the sole, subsistence of the ant. I'.elt 
considers that the presence of the ant is beneficial to the acacia ; 
he supposes that the ants assume the rights of proprietors, and will 
not allow caterpillars or leaf-cutting ants to meddle with their 
property ; the leaves are, he thinks, so preserved to the benefit 
of the tree. 



IV 



ANTS MYRMICIDES 



169 



Kothney has given some particulars of the habits of Sima 
rufo-nigra, an ant of this group that appears to be not uncommon 




FIG. 72. Sima rufo-nigra and 
its associates. A, winged 
female ; B, worker, of the 
ant ; C, Kliinoijsis ruji- 
cornis, a fossorial wasp of 
the sub-family AmjinH- 
cicles ; D, a spider, Salticus 
sp. The coloration is ex- 
tremely similar in all these 
creatures. 



near Calcutta, where it lives on the trunks of trees in company 
with a spider and a wasp that greatly resemble it in form and 
in colour. The three creatures seem to associate together 
on amicable terms ; indeed the wasp and the ant occasionally 
indulge in wrestling matches without doing 
one another any serious harm. In connection 
with this fact we may observe that other 
species of ants have been observed to indulge 
in sports and feats of agility. 

S. leviceps, an Australian species of this 
genus, is furnished with a stridulating file 
that has the appearance of being constructed 

so as to produce two very different kinds of Fit;. 73. stridulating file 
j of Sima lerlceps. 

iv. The CRYPTOCERINI are distinguished from other ants by their 
antennae being inserted at the sides of the head, where they are 
placed between ridges or in a groove into which they can be 
withdrawn ; when in some cases they are entirely concealed. 
These ants assume a great variety of shapes and forms, some of which 
look almost as if they were the results of an extravagant imagin- 
ation. The skeleton is usually much harder than it is in other 




170 HYMENOPTERA CHAP. 

ants ; the abdomen consists almost entirely of one very large seg- 
ment, there being, however, three others visible at its extremity ; 
these segments can be only slightly protruded, and the ants have 
110 power of stinging. They are probably most of them arboreal 
in their habits. Nearly all of the known forms are exotic. 
According to the observations of Bates the species of the genus 

Cryptocerus in the Amazons Valley 
may frequently be observed in dry 
open places on low trees and 1 tushes, 
or running on branches of newly felled 
trees ; they also visit flowers abund- 
antly. The species generally are 
wood-borers, usually perforating the 
dead branches of trees. C. atmttis has 
been observed to construct its nests 
in the dead, suspended branches of 
woody climbers : a number of neatly 
drilled holes are all that can he seen 
pressed first joint of the haul externally ; but, inside, the wood is 

foot is shown at a and b in P -, c ,-, 

different positions. freel j perforated with intercommuni- 

cating galleries. Each community 

appears to consist of a single female and two kinds of workers ; 
the latter in some species are quite unlike each other, differing 
in the form of the head, and in the armature of the thorax and 
nodes of the peduncle. The species of Cryj>t<-fi'nx appear to be 
omnivorous, and are frequently attracted by the excrement of birds. 
The pupae are not enclosed in a cocoon. In the South of Europe 
two very minute ants, of the genera /S'/v////////o///.s and k'jri.tritus, 
belonging to this family, are met with under very large stones 
partly embedded in the earth. They are of the greatest rarity. 

Sub-fam. 4. Ponerides. --Hind body elongate,furnished i'-itJi one 
node at the base, n<l liri n</ <ih<> i/i-ntt fnjmriti/ <//' inurement 
between the first nd second segments, bct/'-t^'/i <>// ir/i there is 
a .s7 ////// constriction, Rtin<j wrl! 




This suit-family includes numerous genera and about 400 
species. The Ponerides have an elongate hind-body; the second 
segment behind the node is eapable of great movement in and out 
of the preceding segment, and for this purpose is furnished with 
a basal portion slightly more slender than the apical part; this 



iv ANTS PONERIDES 17 I 



basal part is usually concealed within the more anterior segment, 
the hind margin of which embraces it very closely. On the 
middle of the dorsal aspect of this articulation there is usually 
placed a stridulating organ, consisting of an elongate band or 
patch of very fine lines ; this gives out a sound when the second 
segment is moved in and out of the first at a time when the 
posterior edge of the latter is slightly depressed. 

We follow Forel in including the Australian bull-dog ants 
Mynnecia in Ponerides, as well as the Odontomachi. The 
former have, however, a definite pedicel, consisting of two nodes 
(Fig. 76). In the Odontomachi the mandibles are approximate 
at their bases, being inserted on the middle of the front of the 
head (Fig. 77). 

This sub-family includes a considerable number of species, 
and is found in all parts of the world. Extremely little is 
known as to the habits, but the true Ponerides do not, so 
far as is known, occur in large communities, and it seems 
probable that they are destitute of the powers of combined 
action that are so remarkable in the Camponotides, and in some 
of the Myrmicides and Dorylides. Most of the species that 
have been described are known by only one sex, so that very 
little knowledge exists as to the sexual distinctions ; but from 
the little that is kno\vn it would appear that the three sexual 
forms are not so differentiated as they are in most of the Cam- 
pouotides and Myrmicides. 

The species of the genus Leptoyenys are believed by Emery 
and Forel to possess an 
apterous female. Mr. 
Perkins has observed that 
the Hawaiian L.falciyera 
has workers with differ- 
ent kinds of sting, but 
no true female. Males of 
this species are, however, 
abundant. "\Vroughton 
has recently discovered 
that one member of this 

m ., FIG. 75. 1)11111/11,11, 'I'll </rtiu<lix, worker. Amazons. 

genus is of Termito- 

phagous habits, but this is not the case with L.fulcigera. Dino- 

ponera grandis (Fig. 75) is the largest of the Ponerides, its 




1/2 HYMENOPTERA CHAP. 

workers attaining an inch and a quarter in length. This Insect, 
according to Bates, marches in single file in the thickets at Para ; 
its colonies consist of a small number of individuals, and are 
established at the roots of slender trees. The effects of its 
powerful sting are not so serious as is the case with some of the 
smaller ants. 

In Britain we have only two representatives of the sub-family, 
viz. Ponera contracta, a small ant of dirty-yellow colour, found 
rarely in the Southern counties, living in moss or under stones. 
Its colonies consist of only a few individuals ; Forel giving fifty 
as the highest number he has observed. The second species, P. 
pwnctatissima, presents the almost unique peculiarity of possess- 
ing two forms of the male sex, one of them resembling the 
worker in most of its peculiarities, and in being destitute of 
wings, while the other is winged, as is usual in male ants. In 
the island of St. Vincent another species of Ponera has been dis- 
covered having an apterous and worker-like male, and was named 
by Forel P. t'l'f/nfifiidria. 1 The discovery of this form has led 
him to express some doubt as to whether Ponera pu nchiti^i nm 
has two forms of males ; but it seems probable that it really is 
so, the ergatoid males being produced under somewhat different 
circumstances from the normal males. We have already said 
that Ci'ti'diocnittfi/la and a few other Myrmicides exhibit an 
analogous peculiarity. 

The genus Myrmccia is confined to the Australian continent 
;nul Tasmania, and includes a considerable number of species of 
large and moderate-sized ants, the classification of which has 
been a subject of difference of opinion-. This has arisen from 
the fact that the nodes of the abdominal pedicel are more similar 
to those existing in the Myrmicides than to those of the typical 
Ponerides. There are, however, some American members of the 
latter sub-family (Para /m in / <-lr<it, <v/.) that differ but little 
in this point from J///v///.r/^, and, moreover, the pupae of Jfyr- 
mecia are enclosed in a cocoon, while in the Myrmicides they 
are usually naked. On the other hand the nests are, it appears, 
very large and populous, more like what exists in the Myrmi- 
cides ; there is no true stridulating organ on the first abdominal 
segment. The genus is therefore one of those interesting 
anomalies that form so large a proportion of the Australian 

1 Tr. ent. Soc. l.nn,],,ii. is;i:;, j.j.. :i6"i-467. 



iv PONERIDES BULL-DOG ANTS 173 

fauna, and will probably be ultimately treated as a distinct sub- 
family. There are about thirty species. 

The ants of this genus are well known to the residents in 
Australia, where they are called " 1 mil- 
dog ants." They form large mounds of 
earth for their nests. The workers, and 
females (Fig. 76) are much alike except 
during the period when the latter are 
still carrying their wings. The males, 
however, differ considerably, being of 
more slender form, and possessing only 
insignificant mandibles, and straight 
antennae with a quite short basal joint. 

Forel considers Myrmecia to be the 
most formidable of all the ants ; the 
hills are said to be sometimes five feet 




FIG. / 6. Myrmeaa jtyrifo. 

high, and the colonies are immense in Australia. Female after 
numbers, while the Insect is an inch or casting wings ' 
more in length, and armed with a very powerful sting, the use 
of which on the human body is said to give rise in some cases 
to serious symptoms. On the other hand, we have seen state- 
ments to the effect that the sting of Myrmecia, has only very 
evanescent sequelae ; it is also said that the ant-hills have only 
a slight elevation, so that probably both these points differ 
according to the species. It appears from a communication of 
Miss Shepherd's that the formidable Myrmecia, forficata has its 
larvae destroyed by a parasitic Hymenopteron (Eucharis myr- 
meciae*) of brilliant colour and considerable size, so that we have 
the curious fact of the hordes of this most formidably armed ant. 

e/ 

which possesses also large eyes, falling a victim to a brilliant and 
very conspicuous Insect. Particulars of this case of parasitic attack 
are still wanting. There are other cases known of the larvae of 
ants being destroyed by parasitic Diptera and Hymenoptera, but 
in none of them have any sufficient observations been made as to 
the mode in which the attack is made. Lowne says that M. t/n/uxn 
itself attacks large beetles of the genus Anoplognathus and buries 
them ; and he also adds the very curious statement that M. niyro- 
cincta, when running, is able to take leaps of a foot in length. 
The Odontomaehi were formerly considered a distinct sub- 

v 

family, distinguished by the peculiar mandibles (Fig. 77). 



1/4 HYMENOPTERA CHAP. 

Many of the Ponerides have elongate mandibles, but they are 
inserted at the sides of the front of the head, not in the middle 
of the front. These organs in some species of Odontomachi 
serve as levers:, by aid of which the Insect can execute considerable 
leaps. In only a few species are the males known ; Mayr and 
Forel state that they are destitute of the peculiar mandibles 
characteristic of the worker. 

The unique European representative of the Odontomachi, 
Anochetus ghiliani, occurs in Andalusia. Near Tangier Mr. 
George Lewis found it to be not uncommon ; but the sexes are 

not known, and it even appears doubt- 
ful whether there exists any well- 
, marked division between workers and 

female. Lewis observed, among the 
ordinary forms, individuals with longer 
bodies, usually one in a nest, and he 
supposed these to be females ; Saun- 
ders, on examining these examples, 
found them to possess distinct ocelli, 
and therefore agreed with Lewis as to 
their being the female sex. Dr. Emery 

subsequently examined these same 

FIG. 77. Anochetus ghiliani, j i i ^ 

worker. Tangier specimens, and took what is scarcely 

a different view, viz. that they are not 

females but an intermediate form; and he also expressed the 
opinion that " the true female may not exist." The male of 
Anochetus is not known. The female of A. mayri, a Neotropical 
.species, has rudimentary wings. 




Sub-fam. 5. Dorylides. Clypens extreme/// *///>///, the 

in *< rt <'<1 very near flic front margin of the head. Hind 
laxly nxniilly elongate anil subcylindricol, with an imperfect 
l><'ilircl formed by f/ie constriction of tlie Imrk of tlie first 
segment, but occasionally tin re nee tiro tiodrs in the workers. 
Distinctions bctn-een tlie hro sexes, mnl between the workers 
i/n</ xe,ee<] forms, enormous. tlie ijiieen* truly ivingless. The 
female* <i n<l worker* nxiiiilly hfiiif/, or <tt onyrnte destitute of 
fiieefteil rt/rs. (In Ecilonini /lie antennae are not inserted, 
i/nite ill I lie front of the head, and there arc ttco nudes in the 
pedicel.^) 



IV 



DORYLIDES WANDERING ANTS 



175 



We have reserved to the end of the ants the consideration of 
the two groups Dorylides and Amblyoponides, recent investigations 
having rendered it somewhat doubtful whether they can be 
maintained as distinct from Ponerides. The chief character of 
the Dorylides is that the males are much less ant-like in form 
than they are in the other groups, and that the distinction 
between the females and workers are enormous. The little that 
is known as to the males and females of this group suggests 
the view that these sexes may offer sufficient reason for keep- 
ing the Dorylides as a group distinct from the other ants ; but 
it must be admitted that it is very difficult to find satisfactory 
characters to distinguish the workers of the Dorylides in some 
cases from the Ponerides, in others (Eciton} from the Myrmicides. 1 

The Dorylides are of great interest, for they exhibit the remark- 
able phenomenon of a nomadic 
social life, accompanied by im- 
perfect sight in the wanderers. 
The sub-family includes two 
apparently distinct groups : 
(1) the Ecitonini, peculiar to 
the New World, and having 
a close relationship with the 
Myrmicides ; and (2) the 
Dorylini existing chiefly in 
the eastern hemisphere, and 
related closely by its workers 
to the Ponerides and Ambly- 
oponides. (i.) The ECITONINI 
consist of the species of the 
genus Eciton, the wandering 
ants of America, and of Labi- 
dus, which there is now good 
reason for believing to con- 
sist of the males of Eciton. 
The female is still uncertain. The Eciton are nomad ants 
having no fixed abode, but wandering from place to place in 
search of prey, and forming temporary resting-places. The 




FIG. 78. Various forms of worker of 
hamatum. Guatemala. 



1 For a valuable revision of Dorylus and its allies see Emery, Zool. Jahrb. Syst. 
viii. 1895, pp. 685, etc. We, however, doubt the wisdom of extending the sub- 
family so as to include Cerapachys, Parasyscia, etc. 



176 HVMENOPTERA 



CHAP. 



species are rather numerous, and the habits of several have been 
described by Bates, who, however, was not acquainted with some 
of the most peculiar features in their biology, these having been 
since revealed by Belt and AY. Miiller. 

These ants are predaceous in their habits, and some of the 
species travel in vast hordes ; they occasionally enter houses and 
clear them of much of the vermin with which they may be 
infested. They have no facetted eyes, some of the forms being 
quite blind, while others have a pair of peculiar lenses in the 
position normally occupied by the compound eyes. Usually 
there are two castes of the workers, and in some species these are 
very different from one another, the mandibles being in the 
larger form very elongate, cylindrical and unfit for industrial 
purposes, while the individuals of the smaller caste have the 
outer jaws shorter, with their edges apposed and coadapted : in 
other species individuals with mandibles differentiated from the 
normal form do not exist. The nomad habits of these ants were 
described by Bates, but the detection of their temporary resting- 
places was reserved for Belt, who found that, after their plundering 
raids, they retired to a place of concealment, and there clustered 
t< >gether in a compact mass like a swarm of bees. Belt says : 
" They make their temporary habitations in hollow trees and 
sometimes underneath large fallen trunks that offer suitable 
hollows. A nest that I came across in the latter situation was 
open at one side. The ants were clustered together in a dense 
mass, like a great swarm of bees, hanging from the roof, but 
reaching to the ground below. Their innumerable long legs 
looked like brown threads binding together the mass, which 
must have been at least a cubic yard in bulk, and contained 
hundreds of thousands of individuals, although many columns 
were outside, some bringing in the pupae of ants, others the legs 
and dissected bodies of various Insects. 1 was surprised to see 
in this living nest tubular passages leading down to the centre 
of the mass, kept open, just as if it had been formed of inorganic 
materials. Down these holes the ants who were bringing in 
booty passed with their prey. I thrust a long stick doWn to 
tlic centre of the cluster and brought out clinging to it many 
ants holding larvae and pupae." 

Turning now to the LnhnJuf, question: many American 
species of this genus have long been known, though all of them 



iv DORYLIDES WANDERING ANTS I // 

by the male sex only. The discoveries (to be subsequently 
alluded to) made in the Old World as to the relations between the 
driver ants and Dorylus raised a suspicion that Labidus might 
In.- the male of Eciton, the distinctions in the two cases being- 
very analogous : this conjecture has been almost proved to be 
correct by the recent observations of Hetschko and W. Miiller. 
The latter, w r ho observed the temporary nests of Eciton liamatum, 
confirms Belt's statements as to the ants hanging together in 
clumps, like swarms of bees ; he also states that the change from 
one temporary abode to another takes place at night, though, as 
is well known, the hunting forays of this ant are carried on in 
the daytime. The periods of migration appear to be determined 
by the time at which all the larvae have assumed the pupal 
state, this at any rate being the time chosen in the case observed 
by Miiller. This naturalist bagged a part of one of the nests 
by the aid of ether, and found the larger portion to consist of 
pupae ; there were also some larvae and eggs ; a specimen of 
Labidus (L. burchelli) was also found on friendly terms with the 
EC it on- workers ; and myrmecophilous Coleoptera were discovered. 
The pupae are enclosed in cocoons. Persistent search failed to 
reveal any female, but the examination was made under great 
difficulties. Miiller also states that the earliest pupated larvae 
yield soldiers, the latest the smallest forms of workers. From 
observations made by Forel on a pupa, it seems probable that 
a wingless form of male may be found to exist. If there- 
i'i >re, as appears practically certain, Labidus is the winged male of 
Eciton, it is probable also that males of more or less worker-like 
form exist, as is now known to be the case in some other 
Formicidae. 

We may here notice a peculiar apterous female ant recently 
described by Andre under the name of Pscudodictkadia incerf". 
He thought this might prove to be the female of Eciton- 
Lubidus ; but his description and figure are imperfect, and do not 
greatly support his idea of a connection between Eciton and 
Psc ti dodic tkadia. 

ii. The group DOKYLINI includes the genus Dorylus, which was 
founded many years ago for Insects very like Labidus. As in 
the case of the American Insect named, males only were known ; 
two or three allied genera, consisting exclusively of individuals 
of the sex mentioned, were subsequently described. In the 

VOL. VI N 



178 



HYMENOPTERA 



CHAP. 



I 



regions inhabited by these males numerous species of blind ants 

are known, but only in the 
worker form, and were, or still 
are, referred to genera called 
Typhlopone and Anomma. No- 
thing that could be considered 
to be a female pertaining to 
any of these Insects was dis- 
covered until Gerstaecker de- 
scribed under the generic name 
Dicthadia an extraordinary 
apterous female ant found in 
Java, and it was suspected 
that it might be the long- 
expected female of the male 
Don/lus and of the worker 




Typhlopone or Anomma. This 
remained for many years with- 
out confirmation, but in 1880 
Trinien announced the dis- 

TiG.'iV.Darylushelcolus. Africa. A, male; covery ill South Africa of ail 
B, female (Dicthadia) ; C, worker major 

(T>/j>?i/<ij><>,<c) ; D, worker minor. (After enormous apterous female ant, 
Emer >"-) allied to Dicthadia; it had 

been disinterred from a nest of small red ants believed (wrongly) 
to be Anomma. As Dorylus had been previously found in con- 
nection with allied worker ants it has since then been clear that 
notwithstanding the enormous differences existing between these 
three forms they may all pertain to one (or to closely allied) 
species. From this summary the student shotild understand that 
he will find in myrmecological literature many references to two 
or three genera that really belong to one species. 

The workers of the Dorylini at present known are without 
exception quite blind, and are believed to be all of predaceous 
habits; it is thought by some that they have no fixed abodes, 
but, like the Ecitonini, frequently change their residence, and it 
has been suggested that in doing so they make use of the nests 
of other ants as temporary abodes ; all these points are, however, 
still unsettled, and as there are several genera it is not unlikely 
that considerable variety will be found to prevail. The driver 
ants of Africa, belonging to the genus Anomma, are in some 



IV 



DORYLIDES DRIVER ANTS 



179 




FIG. 80. Body of male of Dorylus sp. Uelagoa Bay. 
a, pronotuni ; t>, r, divisions of mtsonotum ; d, 
nietanotum ; e, propodeum ; ./', first abdominal seg- 
ment ; (j. //, points of insertion of anterior and pos- 
terior wings. 



respects similar to Eciion in luibits, as they enter human habita- 
tions and cause nearly everything else to quit ; it is probable 
that they are also exclusively carnivorous. Savage detected the 
nests of A. arc ens, but the account he has given of them is too 
vague to permit one to decide whether the assemblages he saw 
were of a nomad kind. The workers of this species vary greatly 
in size, and Emery has 
recently stated that 
lie believes all the 
supposed species of 
the genus to be 
merely varieties of A. 
burmeisteri. The 
female of the driver 
ants is still quite 
unknown. A Dorylus 
has been ascertained 
to be the male of Typlilopone. The male Dorylus (Figs. *79, A, 
and 80) is of great interest, for the propodeum is in a more primi- 
tive form than it is in any other petiolate Hymenopteron known to 
us, while at the same time the pronotum and mesonotum are very 
highly developed. The genus Typldatta Sin. has been recently 
identified by Wroughton and Forel as the worker-condition of 
which Aenictus is the winged male. The genus Alaopone will 
probably be found to have some species of Dorylus as its 
male. 

The females of the Dorylides are amongst the rarest of Insects, 
and are also amongst the greatest of natural curiosities. Although 
worker ants and female ants are merely forms of one sex the 
female yet in this sub-family of ants they have become so 
totally different from one another in size, form, structure, and 
habits that it is difficult to persuade oneself they can possibly 
issue from similar eggs. In the Insect world there are but few 
cases in which males differ from females so greatly as the 
workers of Dorylides do from the females, the phenomena finding 
their only parallel in the soldiers and females of Termites ; the 
mode in which this difference is introduced into the life of the 
individuals of one sex is unknown. The largest of all the 
Dorylides are the African Insects of the genus fihognnts. Only 
the male is known. 



I 80 HYMENOPTERA 



CHAP. 



The specimens of female Dorylides that have been detected may, 
after fifty or sixty years of research, be still counted on the fingers. 
As the greatest confusion exists in entomological literature 
owing to the forms of a single species having been described as 
two or three genera, the following summary of the principal 
names of genera of Dorylides may be useful : 

Eciton = the workers, Labidus = male : ? unknown. 

J'fii-Hi/ixJirf/iitt/ii/ : female only known, possibly that of Eciton. 

Cheliomyrmex : workers and soldiers only known. 

Aenictus = the male, Typlilatt worker : unknown. 

Rliogmus : male ; female unknown. (According to Emery 
the worker is very small and like Alaopone.} 

Anomma : only worker known ; male probably a Dorylus. 

Dorylvs = male ; Dicthadia = 9 : Alaopone and TypJilopone - 
workers. 



Sub-Fam. 6. Amblyoponides. Abdomen destitute of distinct 
IH'ilirf] ; f/ie articulation between the first and second 
segments behind the true petiole being broad. 

"We follow Forel in separating Amblyorjone and a few allies 
from the Ponerides, because the abdominal pedicel is more 

imperfect than in any other ants. It is, 
indeed, very difficult to frame a definition 
that will include the Amblyoponides 
among ants, and at the same time sepa- 
rate Formicidae and Scoliidae. Forel con- 
siders the Amblyoponides to approach 
closely to certain divisions of the Scoliidae 
(Thyimides, e.g.}. Little is known of these 
\jf) I \. Insects, though they are widely distri- 
1 mtecl. A in bli/npone, is found in Australia 
and Xew Zealand ; the allied genus 

f, worker. Sfio ma fo mma h as a ^ide distribution, 
Tasmania. 

occurring even in Europe. The social 

life is believed to be imperfect, and the habits subterranean and 
M'llmtary. The males and females are winged; the latter much 
resemble the workers, which are nearly blind, and have a con- 
siderable general resemblance to Anomma, in Dorylides. 

Association of Ants with other kinds of Insects. -We 
have already alluded to tin- fact that a few species of ants are 




iv INSECTS INHABITING ANTS'-NESTS I 8 I 

used liy other species as attendants, and that the two kinds then 
live together quite amicably ; and we have also seen that a few 
ants live in association with other species on terms that are not 
yet understood. One little ant, Formicoxenus niti<lntux, lives 
only in the large nests of Formica rufa ; these ants tolerate 
the little Form-iro,irn./>s, which so far as is known does them 
neither sjood nor harm. There are also a considerable number of 

O 

species of small ants that are in the habit of choosing the neigh- 
bourhood of larger species for their dwelling-places; in some 
cases the nests are constructed actually within a portion of the 
edifice of the more powerful species, and the rule then appears 
to be that these neighbours do not molest one another. X< it- 
withstanding the militant lives that many of them lead, ants 
cannot be considered as of generally ferocious disposition. 

But the most remarkable point in connection with their 
toleration consists in the fact that the nests of many species are 
inhabited by quite a colony of foreign Insects of various Orders ; 
many of these, being found nowhere else, are spoken of as ants'- 
nest or Myrmecophilous Insects. 1 The relations of ants with 
other Insects are of the most varied and complex character ; 
some of their guests live with them on terms of the most intimate 
association, being indeed absolutely dependent for their existence 
on the good offices of their hosts ; others of the ants'-nest Insects 
are enemies, while others are neutral or indifferent to the ants. 
We have already mentioned that the guests migrate in company 
with their hosts. 

Many species of ants derive a considerable portion of their 
sustenance from the sweet substances excreted by Aphidae. 
Ants may constantly be seen occupied with clusters of Aphidae, 
and it is said that the ingenious little creatures defend from 
enemies the manufacturers of the sweet-stuff they are so fond of, 
even going so far as to form barricades and covered places for the 
isolation and protection of this peculiar kind of cattle ; a few 
ants keep some of the root -feeding Aphidae in their nests. 
Coccidae and other Homoptera, which also excrete much matter 
of a sugary nature, are likewise consorted with by ants; as are 
also the larvae of some butterflies of the family Lycaenidae ; 
these latter being believed to furnish to the ants some substance 

1 A Catalogue of Myrmecophilous and Termitophilous Arthropods was pub-' 
lished by Wasmanu, Berlin 1894. 



I 82 HYMENOPTERA CHAP. 




of ;i nutritious kind. The Insects we have spoken of are, how- 
ever, rather of the nature of ant-cattle, and the fondness of the 
ants for them is not very remarkable. The relations of the ants 
to the peculiar species of Insects that live only in or around their 
nests are much more extraordinary. The greater number of these 
guests belong to the Order Coleoptera, and of these there are many 
hundreds probably many thousands of species that depend on 
ants for their existence. The family Pselaphidae furnishes a 
large number of ants'-nest beetles, and it appears probable that 
most of them excrete some 
sugary substance of which the 
ants are fond. Many of these 
Pselaphidae are of the most 
fantastic shapes, more especi- 
ally the members of the sub- 
family Clavigerides. But the 

,, , , FIG. 82. The beetle, A temeles, soliciting 

most curious of all the ants- food from an ant. (After Wasmann.) 

nest beetles are the Paussidae, 

a family exclusively dependent on ants, and having the curious 
faculty, when disturbed, of bombarding that is, of discharging 
a small quantity of vapour or liquid in a state of minute 
subdivision accompanied by a detonation. Many species of 
Staphylinidae are peculiar to ant's-nests, and most of them are 
indifferent or inimical to their hosts, but some of them, such as 
Atemeles (Fig. 82) and Lomechusa, are doubtless producers of sweet 
stuff that is liked by the ants. The ants feed some of their special 
favourites amongst these guests in the same manner as they feed 
one another, viz. by opening the mouth, causing a drop of liquid 
to appear on the lip, and remaining passive while the guest 
partakes of the proffered bonne boucJie. This way of giving food 
to other individuals is a most remarkable feature in the character 
of ants; it is not the same system that they adopt in feeding the 
larvae, for they then make a series of actual movements, and 
force the nutriment into the mouths of the grubs. Besides the 
Insects we have mentioned there are also Orthoptera, Hemiptera, 
Poduridae and Thysanura,, Acari, and small Isopod crustaceans 
that live exclusively in company with ants. We have mentioned 
that a few Hymenopterous and Dipterous parasites have been 
detected living at the expense of ants ; it is probable that 
closer observation of the ant larvae and pupae in their nests 



iv INSECTS INHABITING ANTS'-NESTS 183 

will disclose a greater number of the parasites of this latter 
class. 

Much attention has been given to the relations between ants 
and their guests by Wasmann. 1 He arranges them in four 
categories ; 1, " Symphily " for the true guests, which are fed and 
tended by the ants, the guests often affording some substance 
the ants delight in ; 2, " Metochy," the class of tolerated guests, 
being so far as is known not disagreeable to the hosts ; 3, 
" Synecthry," including those Insects, etc., to which the ants 
are hostile, but which nevertheless maintain themselves in the 
midst of their foes ; 4, Parasites, dwelling in the bodies of the 
adult, or of the young ants. Many of these ants'-nest Insects 
present a more or less perfect resemblance to the ants in one or 
more points, such as sculpture, colour, size, or form. To these 
resemblances Wasmann attaches great importance. We should, 
too, notice that some of the inquilines ' 2 have become acquainted 
with the movements and habits of the ants, and stroke them (as 
the ants do one another) to induce them to disgorge food in the 
manner we have alluded to. According to Janet, ants of the 
genus Lasius are infested by Acari of the genus Antennophorus. 
The ants carry the mites, which assume positions so as not to 
cause greater inconvenience than is inevitable. Moreover, the 
ants give food to the mites when requested, and behave in a most 
obliging way to them, though there is not any reason for supposing 
that in this case the ants derive any benefit from the Symphily. 

The relations between ants and plants have been of late years 
much discussed. We have already briefly alluded to the subject 
when speaking of the Pseudomyrmini. We will here only remark 
that ants frequent plants not only for the purpose of securing the 
sweet stuff excreted by the Aphidae that live on them, but also 
for the sake of getting the sweet products the plants themselves 
afford. Mr. Aitken, speaking of ants in India, says : " I have 
come to the conclusion that one of the most important sources of 
food-supply which ants have is the sacchariferous glands to be 
found at the bases of so many leaves." It is supposed that the 
ants are on the whole beneficial to the plants that thus afford them 
supply ; and this fact is considered by many to afford an adequate 
explanation of the existence of these interesting relations. 

1 For a summary of this subject see Wasmann, Congr. internal. Zool. Hi. 1896, 
pp. 411-440. 2 For explanation of tliis term see vol. v. p. 524. 



CHAPTER V 

COLEOPTEKA OR BEETLES 

Order V. Coleoptera. 

Apparently wingless Insects when at rest, but really with four 
wings; the elytra, or anterior pair, shell-like, reposing 
an. the back of the body and fitted together accurately 
a font/ the middle I// a straight suture; the posterior pair 
membranous, folded together under the elytra. Mouth with 
mandibles; lower lip not divided along the middle. Meta- 
//i<i/'/>/iosis great and very abrupt ; the larra being a grub or 
maggot, which changes to a jmpa (usually soft} in which the 
external structure of the perfect Insect is conspicuous. 

COLEOPTERA or Beetles are chiefly distinguished from other 
Insects by the solidity of their external integument, and by the 
peculiar nature of the first pair of their alar organs, which do 
not serve as instruments of flight, but as shells for protecting the 
upper face of the after-body, which, unlike the other parts, remains 
as a rule soft and membranous. These modifications of structure, 
though apparently slight, must be really extremely advantageous, 
for beetles are the predominant Order of Insects in the existing 
epoch. They depart from most other Insects in being less aerial in 
their habits ; therefore, notwithstanding their enormous numbers, 
they do not meet the eye so frequently as flies, bees, or butter- 
flies. The parts of the hard outer skeleton are beautifully 
fitted together, and as their modifications are easily appreciated 
they offer as fascinating a subject for study as do the skeletons 
of Yertebrata. The habits of beetles are so extremely varied 
that it is but little exaggeration to say that Coleoptera are to 
be found everywhere, when looked for. The number of species 
at present known is probably about 150,000. Of these some- 
where about 3300 have been found in Britain. The structure 



CHAP. V 



COLEOPTERA BEETLES 



I8 5 



of the hard parts of the skeleton is of importance, as the classi- 
fication of this enormous number of species is entirely based 
thereon ; it will be readily understood from the accompanying 
diagram (Fig. 83). The general proportions of the chief parts 
of the body call for a few remarks. The prothorax is remarkably 
free, and is therefore capable of a much greater amount of move- 
ment independent of the after-body than it is in other Insects. 
The mesothorax is, on the other hand, much reduced ; its chief 
function in the higher forms is to support the elytra, and to 



Fru. 83. Under- surface of a beetle, 
Harpalus caliyinosus ; legs and 
antenna of one side, and some parts 
of the mouth removed. A, an- 
tenna ; B, mandible ; C, labruni ; 
D. ligula ; E, paraglossa ; F. labial 
palp ; G, inner lobe of maxilla ; 
H, outer lobe (palpiform) of 
maxilla ; I, maxillary palp ; K, 
mentum ; L, gena ; M, gula : N, 
huccal fissure ; V, plates of ven- 
tral segments. 1, Prostermim ; 
2. prosternal episternum ; 3, pro- 
sternal epimerou : 4, anterior and 
middle coxal cavities ; 5, iuflexed 
side of pronotum; 6, nifsosterimm ; 
7, mesosternal episternum ; 8, 
mesosternal epimeron ; 9, nieta- 
sternum ; 10, posterior division of 
metasternum or ante-coxal piece ; 
11, metasternal episternum ; 12, 
metasternal epimeron ; 13, epi- 
pleuron or inflexed margin of ely- 
tron ; 14, ventral or ambulatory 
setae ; 15, trochanter ; 16, pos- 
terior coxa ; 17, femur ; 18, tibia ; 
19, tarsus. (Modified from Le- 
coiite and Horu.) 



help to keep them together by means of its scutellum. The 
metathorax, on the contrary, is largely developed, except in the 
rather numerous forms that are entirely deprived of powers of 
night. The composition of the abdomen has been a subject of 
great difference of opinion. Its upper surface is usually entirely 
covered by the elytra; the parts visible on the lower surface are 
called ventral segments, and are usually five in number. Although 
these five plates may constitute all that is superficially visible of 
the abdomen, yet if the elytra are taken off it is found that a larger 
number of segments usually seven or eight are visible on 
the dorsum. This seeming discrepancy of number between the 




'-14 



I 86 COLEOPTERA 



CHAP. 



dorsal and ventral plates is due to two facts ; 1, that the 
hind coxae have a great and complex development, so that they 
conceal the true base of the venter, which, moreover, remains 
membranous to a greater or less extent, and thus allows much 
mobility, and at the same time a very accurate coadaptation 
between the hard parts of the venter and the metasteruum l ; 2, 
that the terminal segments are withdrawn into the interior of 
the body, and are correspondingly much modified, the modifica- 
tion being greater in the case of the ventral than in that of the 
dorsal plates. The anatomy of the parts of the abdomen that 
are not externally visible has not been adequately studied by 
coleopterists, but Yerhoeff has inaugurated a careful study of 
the comparative anatomy of the terminal segments 2 ; unfor- 
tunately, however, he has not so thoroughly studied the modifi- 
cations at the base, and as it is not clear that these are so 
uniform as he has taken for granted, it is possible that his num- 
bering of the segments may have to be in some cases modified. 
The retracted plates or segments are so intimately connected 
with the internal copulatory organs that it is no easy matter to 
interpret them. For the nomenclature of these parts we must 
refer the student to Verhoeff's later works. He considers the 
abdomen as composed of ten segments, the dorsal plates being 
demonstrable, while the tenth ventral plate is usually absent. 
The anal orifice is placed immediately beneath the tenth dorsal 
plate, and above the genital orifice, which lies behind and above 
the ninth ventral plate. Peytoureau admits a diversity in both 
the number of segments and the position of the orifice. These 
studies in comparative anatomy are surrounded with difficulties, 
and no morphological conclusions based on them can be con- 
sidered as final until they have been confirmed by observation 
of the development of the parts. 

The elytra or wing-cases frequently have a remarkable 
sculpture, the use of which is unknown. According to Hofbauer 
there are between the outer and inner layers, glands secreting a 

1 An interesting exception occurs in the Malacodermidae, where this coadaptation 
is wanting, or is imperfect ; they are frequently considered to be the most primitive 
of existing beetles. 

2 In a series of memoirs in various German periodicals during the last five or six 
years (see especially Deutsche ent. Zcit. 1893 and 1894, also subsequent years of 
Arch. Xaturgcs.}. It should be noticed that in the course of his studies Verhoetf 
has modified some of his earlier views. 



BEETLES 187 



fluid that reaches the surface through small pores. Hicks sup- 
posed that he detected nerve cells. Meinert is of opinion that 
the elytra correspond to the teguke of Hymenoptera rather than 
to the wings of other Insects, but the little evidence that exists is 
not favourable to this view. The two elytra are usually, in repose, 
very perfectly fitted together by a complete coadaptation along the 
middle of the body, so that it is difficult to separate them ; this 
line of junction is called the suture. There are forms in which 
the coadaptation is quite imperfect (Malacodermidae) and some 
in which it does not exist at all (Meloty. The wings proper of 
beetles correspond to the posterior pair in other Insects, and are 
much more irregular in nervuration than those of most other In- 
sects, correlative, it is supposed, with the folding they are subjected 
to in order to get them beneath the wing-cases. There are large 
numbers of species, genera, and groups of genera, all the members 
of which have the wings so much reduced in size as to be quite 
useless for purposes of flight. These forms are called apterous, 
though they are not really so, for the elytra (which are really 
the anterior wings) are present, and even the posterior wings 
are not truly absent in these cases, though they are sometimes 
so extremely rudimentary as to elude all but the most careful 
observation. The number of forms in which the elytra are 
absent is extremely small ; this condition occurs only in the 
female sex ; it is usually confined to cases in which the female 
is larva-like in form ; but in the extraordinary Mediterranean 
Lamellicorn genus, Packypus, the females are destitute of wings 
and elytra, though the anterior parts of the body are normally 
formed : these individuals live underground and rarely or never 
emerge. When the wings are absent the elytra are frequently 
soldered ; that is to say, united together along the suture by some 
sort of secondary exudation ; this union occurs in every degree of 
firmness, and appears to be variable in the individuals of one 
species ; probably in accordance with the age of the individual. 
In most beetles the elytra are not only themselves closely con- 
nected, but are also very accurately coadapted with the sides of 
the body, except at the tip. Sometimes a coadaptation occurs 
between the tips of the elytra and the body, but not at the tip of 
the latter. In such cases one or more dorsal plates are left ex- 
posed : the last of such exposed dorsal plates is termed pygidium : 
a similar plate anterior to the pygidium is called propygidium. 



I 88 COLEOPTERA 



CHAP. 




Larvae. Owing to the difficulty of rearing Coleoptera, less 
is perhaps known of their life-histories than of those of other 
Insects. They exhibit, however, extreme diversity correlative 
with the great specialisation of so many beetles to particular 
kinds of life. Most beetles must have exactly the right condi- 
tions to live in. The larvae of many forms are known. They 
are composed of a head, three thoracic segments (usually very 
distinct), and a number of abdominal segments varying from eight 
to ten. Coleopterous larvae are usually described as having nine 
abdominal segments; and it is but rarely that ten can be readily 
detected ; they are, however, visible in various forms, as is the 
case in the form figured (Fig. 84). A great many of them 
possess a peculiar pseudopod at the underside of the body near 
or at the extremity ; it can in many cases be entirely retracted 
into the body, and is generally described as being the pro- 
truded termination of the ali- 
mentary canal. Inspection of a 
series of larvae shows that it 
represents a body segment : it is 
FIG. 84. Larva of a i.eetie, Family sometimes armed with hooks. 

Cerambycidae (I.Aroniiti ii<i~in<-luitt(). mi p n J.T i 

The first spiracle is placed just at the Three pairs of small thoracic legs 
hind margin of the large prothoracic are often present, but are very 
segment. (From La Massaiie. ) Pj m , 

often completely absent. These 

thoracic legs may be present in the young larva, but not in the 
older (Bruchus). The usual number of spiracles is nine pairs, 
me prothoracic, eight abdominal; but this is subject to many 
exceptions, and mesothoracic and metathoracic stigmata are occa- 
sionally found. The figures we give in the following pages will 
enable the student to form some idea of the variety of form 
exhibited by beetle larvae. 

Pupation usually takes place in a cavity in the earth, or 
near the feeding-place, but a great many species form a cocoon, 
composed either of fragments of earth or of wood, and slightly 
cemented together. A few suspend themselves by the tail after 
the manner of butterfly caterpillars (Cassididae, Coccinellidae). 
The pupae are usually extremely soft, their appendages not 
being fastened to the body. But some pupae (Staphylinides) 
are truly obtected, having a hard shell and the rudimentary 
appendages fastened by exudation to the body, like Lepi- 
dopterous pupae, and others (Coccinellidae) are intermediate 



BEETLES 189 



between this state and the normal soft pupa. The pupal state 
lasts but a short time, from one to three weeks being the usual 
period. The perfect Insect is at first soft and almost colourless, 
and it is often some days before it attains its complete coloration 
and hardness. 

Classification. Owing to the hardness of the skeleton, beetles 
shrivel but little after death, so that the form and relations of 
the various sclerites can usually be detected with ease. These 
sclerites seem to be remarkably constant (except in the case of 
sexual distinctions) within the limits of each species, and are 
very useful for the formation of genera and groups of genera : 
but they vary so much outside the limits mentioned that it is 
very difficult to make use of them for defining the larger groups. 
Hence it is not easy to frame accurate definitions of the 
families, and still less so to arrange these families in more com- 
prehensive series. The natural difficulty has been much increased 
by the habit coleopterists have of framing their definitions 
on what is visible without the aid of dissection. Nevertheless 
considerable progress has been made. "We are obliged at present 
to adopt upwards of eighty families ; and we are able to dis- 
tinguish on positive characters five series; this leaves a large 
number of forms still unclassified, and these we have here 
associated as a sixth series, which we have called Coleoptera Poly- 
morpha. This series corresponds with the two series called in 
books Clavicornia and Serricornia. As it is admitted to be 
impossible to define these two series, we think it much better to 
act accordingly, and to establish for the present a great group 
that can only be characterised by the fact that its members do 
not belong to any of the other five series. No doubt a larger 
knowledge of development, coupled with the advance of com- 
parative anatomy, will ultimately bring about a better state of 
affairs. The Strepsiptera, with one family Stylopidae, are only 
provisionally included among the Coleoptera. These six series 
are fairly equal as regards extent. Though the Polymorpha 
includes the larger number of forms, yet a large part of them 
belong to four great families (Staphylinidae, Buprestidae, Elat- 
eridae, Malacodermidae), which are easily recognisable, so that 
the number of unmanageable forms is not really great. Indeed, 
an acquaintance with the external anatomy of two or three 
dozen species, selected as typical, would enable a student to classify 



1 90 COLEOPTERA CHAP. 

with tolerable certainty the vast majority of species that he 
would subsequently meet with. 

Series 1. Lamellicornia. Antennae with the terminal joints leaf-like (or 
broader than the others, if not actually leaf-like), and capable of 
separation and of accurate apposition. Tarsi five-jointed. 

Series 2. Adephaga (Caraboidea of some authors). Antennae never lamelli- 
f 01-111, thin at the end ; all the tarsi five-jointed, with the fourth 
joint quite distinct. Maxillae highly developed, with the outer 
lobe slender and divided into two segments so as to be palpiiorm. 
Abdomen with six (or more) ventral segments visible. 

Series 3. Polymorpha. Antennae frequently with either a club, i.e. the 
distal joints broader [Clavicorn series of authors], or the joints from 
the third onwards more or less saw-like, the serrations being on the 
inner face [Serricorn series of authors]; but these and all the other 
characters, including the number of joints in the feet, very variable. 

Series 4. Hetcromcra. Front and middle tarsi five-jointed, hind tarsi four- 
jointed. Other characters very variable. 

Series 5. Phytophaga. Tarsi four-jointed [apparently], but with a small addi- 
tional joint at the base of the fourth joint : sole usually densely 
pubescent [sometimes the feet are bare beneath or bristly, and 
occasionally the small joint at the base of the fourth joint is more 
distinct]. 

Series 6. Rhynchophora. Head prolonged in front to form a beak ; gula 
indistinguishable. [Palpi usually not evident.] Tarsi four-jointed 
[apparently], but with a very minute additional joint at the 
extreme base of the fourth joint. 
Streps'iptera (see p. '298). 

The first and second series, with much of the third, form the 
Pentainera, the fifth and sixth the Tetramera [or Pseudotetra- 
inera *]. The term Isomera was applied by Leconte and Horn 
to a combination of series 1, 2, 13, and 5. 

Series I. Lamellicornia. 

Tarsi five-jointed ; antennae with the terminal joints (usually three, 
sometimes more), broader on one side, so as to form a peculiar 
cluli, the leaves of which are movable, and in repose are 
more or less perfectly eoadapted so as to hare the 'appearance 
of being lut one piece. 

This series includes three families, Passalidae, Lucanidae, and 
Scarabaeidae : the latter includes an enormous majority of the 
>]><-<-ies, and in them the structure of the antennae characteristic 
<jf the scries is well developed; but in the other two families 

1 We consider this term inferior to Tetramera for nomenclatorial purposes. 



LAMELLICORNS 



the form of the antennae is not so widely different from that of 

other Coleoptera. The larvae live on decaying vegetable matter, 

roots or dung. They 

have three pairs of 

legs, and are thick 

clumsy 

curved 




grubs with 

bodies, the 
last two segments 
being of larger size 
than usual. Many of 
them possess organs 
of stridulation, and 
the structure of their 
spiracles is very 
peculiar, each one 
being more or less 
completely sur- 
rounded by a chitin- 
ous plate. The 
spiracles usually form 

a System 

C'losed except at the 

moment when the 
skin is shed and the tracheal exuviae are detached. Meinert T 
considers these spiracles to be organs of hearing. The life of the 
larvae is passed underground or in the decaying wood on which 
the Insect feeds. 

Most of the members of this series are remarkable on account 
of the great concentration of the nerve-centres. This is extreme 
in Rhizotrogus, where there are only two great ganglia, viz. the 
supra-oesophageal and a great ganglion situate in the thorax, 
and consisting of the conjoined infra-oesophageal, thoracic, and 
abdominal ganglia, According to Brandt 2 there are several 
distinct forms of concentration in the series ; the Lucanidae only 
participate in it to the extent that the perfect Insects exhibit 
fewer ganglia than the larvae ; the latter possess two cephalic, 
three thoracic, and eight abdominal ganglia, while the perfect 
Insect has the abdominal ganglia reduced in number to six, and 



FIG. 85. Antennae of Lamellicoms. 1. AWts inter- 

entirely niptus ; - 2, Lucan-us cerrus <3 ; 3, J'/K/I/KCUS splen- 

didulus 9; 4, Phileurus didymus ?; 5, 
fitllo (J. 



1 J >(mske SeM: Mr. (G), viii. No. 1, 1895. 

2 Horae Soc. ent. lloss. xiv. 1879, ]>. 15. 



I 9 2 



COLEOPTERA 



CHAP. 



they are placed partially in the thorax. The diminution in 
number takes place in this case by the amalgamation of the first 
two abdominal with the last thoracic ganglia. 

Fam. 1. Passalidae. Lcibrum large, mobile ; mentum deeply 
cut out in the middle for the accommodation of the liyida ; the 
lamellae of the anten/nt In-onnlit together by the curling up of the 
antenna. The elytra entirely cover the dorsal suifuce <>f the abdo- 
men,. There are four or five hundred species of this family known ; 
they are usually shining-black, unattractive beetles, of large size, 



-I- -I 



FIG. 86. View of one 
7 side of nieso- ami 
metathorax of a 
Passalid larva from 
Borneo showing the 
stridulating organs. 
a, b, Portions of the 
metathorax ; c, coxa 
of 2nd leg; d, striate 
or stridulating area 
thereon ; e, basal 
part of femur of 
middle leg ; /', hairs 
with chitiuous pro- 
cess at base of each ; 
g, the diminutive 
3rd leg modified 
for scratching the 
striated area, x 20. 




-I 



and are abundant in the decaying wood of tropical forests. They 
are quite absent from Europe, and there is only one species in 
the United States of North America. The larvae are very 
interesting, from the fact that they appear to have only four legs. 
This arises from the posterior pair being present only as very 
short processes, the function of which is to scrape striated areas on 
the preceding pair of legs and so produce sound. In the specie^ 
figured (Fig. 86) this short leg is a paw-like structure, bearing 
several hard digits ; but in other species it is more simple, and 
without the digits. The perfect Insect has no sound-producing 
organs, and it is very remarkable therefore to find the larvae 



v LAMELLICORNIA STAG-BEETLES 193 

provided with highly -developed stridulatoiy structures. No 
auditory organ is known, unless the peculiar spiracles be 
such. 

Fam. 2. Lucanidae (Stag-beetles). Labrum indistinct, fixed ; 
mentum not excised ; antennae not curled in repose, with but little 
coadaptation of the terminal joints ; the elytra, entirely cover the 
dorsal surface of the abdomen. The Stag-beetles are well known 
on account of the extraordinary development of the mandibles in 
the male sex, these organs being in some cases nearly as long as 
the whole of the rest of the Insect, and armed with projections 
or teeth that give the Insects a most formidable appearance. So 
far as we have been able to discover, these structures are put to 
very little use, and in many cases are not capable of being of 
service even as weapons of offence. The males are usually 
very much larger than the females, and are remarkable on 
account of the great variation in the stature of different indi- 
viduals of the same species ; correlative with these distinctions 
of individual size we find extreme differences in the development 
of the head and mandibles. Moreover, the small male specimens 
exhibit not merely reductions in the size of the mandibles, but also 
show considerable differences in the form of these parts, due, 
in some cases, apparently to the fact that only when a certain 

length of the mandible 
is attained is there any 
development of certain 
of the minor projec- 
tions : in other cases it 
is not possible to adopt 
this view, as the small 
B mandibles bear as many 

FIG. 87.-Head M anaprothoraxofformsof the male of projections ES the large 

a stag-beetle ; Homoeoderns mellyl (Africa). A, forms do, Or even more. 

Large. B., intermediate, C, small forms. (From a T 

photograph by ROberthtir.) In each species these 

variations fall, in the 

majority of cases, into distinct states, so that entomologists describe 
them as " forms," the largest developments being called teleodont, 
the smallest priodont ; the terms mesodont and amphiodont being 
applied to intermediate states. Leuthner, who has examined many 
specimens, states that in Odontolabis sinensis, no intermediates 
between the teleodont and mesodont forms occur, and as the 

VOL. VI 




194 



COLEOPTERA 



CHAP. 





two forms are very different they are liable to be mistaken for 
distinct species. 

There are at present between 500 and 600 species of stag- 
beetles known ; the Indo-Malayan and Austro-Malayan regions 
being richest in them. Australia possesses many remarkable 
and aberrant forms. In the Ceratognathini a group well re- 
presented in New Zealand as well as 
in Australia the structure of the 
antennae is like that of the Scara- 
baeidae, rather than of the Luca- 
nidae. The most aberrant form 
known is, however, our common 
Sinodendron cylindricum ; this de- 
parts in numerous features from 
other Lucauidae, and instead of 
the mandibles of the male being 
FIG. w. Sinodendron cyiindricum. more largely developed, there is a 

A, Larva; B, pupa. New Forest. 

horn on the head ; it is very 

doubtful whether this Insect should be allowed to remain in 
the family. Little is known of the habits and development of 
Lucanidee, except in the case of three or four species that are 
common in Europe. 

The common stag-beetle, Lucanus cervus, is our largest British 
beetle. The larva much resembles that of Melolontlia mdgaris, 
but attains a larger size, and the anal aperture is placed longitu- 
dinally instead of transversely ; it lives in decaying wood, or eats 
the roots of trees without being injurious ; its life in this state 
lasts about four years ; the pupal, period is passed through rapidly, 
and the perfect Insect may remain for months underground 
before it becomes active ; this occurs in June and July. This 
larva stridulates by scraping certain hard tubercular ridges on 
the third pair of legs, over a specially adapted rough area at the 
base of the second pair. 

The Passalidae and Lucanidae are united by some authorities 
as a group called Pectinicornia ; the term Lamellicornia being 
then confined to the Scarabaeidae. The Passalidae appear, how- 
ever, to be really more nearly allied to the Scarabaeidae than to 
the Lucanidae. 

Fam. 3. Scarabaeidae (Chafers'). The leaflets of the antennae 
are well coadapted, and are susceptible of separation. The elytra 



v LAMELLICORNIA SCARABAEIDAE 195 

usually leave the pygidimii uncovered. The number of visible 
i'1'ntral segments is usually six, or at the sides seven, not Jive, as in 
Lucanidae and Passalidae. This is one of the most important 
families of Insects. About 13,000 species are already known; 
as some of them are highly remarkable creatures on account of 
the males being armed with horns, they are figured in many 
works on natural history. There is great variety of form, and 
the following five sub-families may be adopted, though authorities 
are by no means agreed as -to the classification of this extensive 
family, which, moreover, lie it remarked, is increasing by the dis- 
covery of about 300 new species every year. 

Abdominal spiracles plaivd in a line on the connecting membranes, and en- 
tirely covered by the wing-cases (Laparosticti). Sub-fain. 1. CopRiDES. 1 

Abdominal spiracles placed almost in a line, but only the basal three on the 
connecting membranes ; the terminal one usually not covered by the 
wing-cases. Sub-fain. 2. MELOLONTHIDES. 

Abdominal spiracles placed in two lines, the basal three on the connecting 
membranes, the others on the ventral segments (Pleurosticti). 

The claws of the tarsi unequal. Sub-fam. 3. RUTELIDES. 

The claws of the tarsi equal ; the front coxae transverse, but little pro- 
minent in the descending axis. Sub-fam. 4. DYNASTIDES. 

The claws of the tarsi equal ; the front coxae more prominent, shorter 
transversely. Sub-fam. 5. CETONIIDES. 

i. The COPRIDES form an immense group of about 5000 species ; 
they differ somewhat in habits from other Lamellicorns, inasmucb 
as most of them live on dung, or decaying animal matter ; the 
sub-family connects with the Lucanidae, so far as superficial char- 
acters go, by means of two of its groups, Trogini and Nicagini, 
the latter being very near to the Ceratognathini in Lucanidae. 
So little is known as to the morphology and development of 
these groups that it is not possible to pronounce an opinion as 
to the validity of this, apparent alliance. Trox stridulates by 
rubbing two raised lines on the penultimate dorsal segment 
across two striate ribs on the inner face of the elytra ; Geotrupes, 
on the other hand, produces an audible sound by rubbing together 
a file on the posterior coxa and a fine ridge on the contiguous 
ventral segment. The larva in this genus has a different organ 

1 In this sub-family there are numerous forms iu which the elytra cover the 
pygidium, and in which the number of conspicuous ventral segments is reduced to 
five or even lour. We use the term Coprides as equivalent to the "Laparosticti" 
of Lacordaire (Gen. Col. iii. 1856) ; it thus includes the "Comlni" and " Glaphy- 
rini " of the Cutaloyus Coleopterorum, vol. iv. Munich, 1869. 



1 96 COLEOPTERA CHAP. 

for stridulatiou from the imago ; it is placed on the second and 
third pairs of legs, the latter pair being much reduced in size. 

The most interesting division of the Coprides is the group 
Scarabaeini. Xo member of this group inhabits the British 
islands, but in Southern Europe, and in still warmer lands, these 
Insects are well known from the curious habit many of the species 
have of rolling about balls of dung and earth. The long hind 
legs are chiefly used for this purpose, and it is on the peculiar 
structure of these limbs that the group has been established. 
Many of the stone Scarabaei found in Egyptian tombs represent 
some kind of Scarabaeini, and it has been said that the ancient 
Egyptians looked on these Insects as sacred because of their 
movements. These must certainly appear very strange to those 
who see them and are unacquainted with their object. It is 
stated that the dwellers in the vallev of the Xile thought the 

. 

actions of these Insects, when rolling their balls, were typical of 
the planetary and lunar revolutions : and that the sudden 
appearance of the beetles after a period of complete absence was 
emblematic of a future life. Many accounts have been o-iven 

c. 

of the habits of members of this group, but according to Fabre 
all are more or less erroneous ; and he has described the habits 
and life-history of Scarabaeus sacer (Fig. 89), as observed by him 
in Southern France. These Insects act the part of scavengers by 
breaking up and burying the droppings of cattle and other 
animals. The female Xcarabaeus detaches a portion of the dung 
and forms it iiito a ball, sometimes as large as the fist : this it 
rolls along by means of its hind legs, by pushing when necessary 
with its broad head, or by walking backwards and dragging the 
ball with its front legs. The strength and patience displayed by 
the creature in the execution of this task are admirable. Fre- 
quently the owner of this small spherical property is joined, so 
Fabre informs us, by a friend, who is usually of the same sex and 
assists her in pushing along the ball till a suitable place is reached. 
AVhen this is attained, the owner commences to excavate a chamber 
for the reception of the ball ; sometimes the false friend takes ad- 
vantage of the opportunity thus offered and carries off the ball 
for her own use. Should no disappointment of this sort occur, the 
Scarabaeus accomplishes the burying of the ball in its subterranean 
chamber, and accompanies it for the purpose of devouring it ; the 
feast is continued without intermission till the food is entirely 



LAMELLICORNIA COPRIDES 



197 



exhausted, when the *SV"/v//V/< //.< .-^eks a fresh store of provender 
u> lie treated in a similar manner. According to M. Fabre's 
account these events occur in the spring of the year, and when 
the hot weather sets in the ,SVv//w/,</< //.; passes through a period of 
quiescence, emerging again in the autumn to recommence its 
labours, which are now, however, directed immediately to the con- 
tinuance of the species : a larger subterranean chamber is formed. 
and to this retreat the beetle carries dung till it has accumulated 
a mass of the size of a 
moderate apple ; this mate- 
rial is carefully arranged, 
previous to the laying 
of the egg, in such a 
manner that the grub to 
be hatched from the egg 
shall find the softest and 
most nutritive portions 
close to it, while the 
d>arser and more innu- 
tritions parts are arranged 
- ts to be reached by the 
grub only after it has 
acquired some strength ; 
lastly, a still more deli- 
cate and nutritive paste 
is prepared by the mother 
tie for a first meal for 
the newly -hatched grub, 
by some of the food being 
submitted to a partial digestion in her organs ; finally, the - _ _ 
is deposited in the selected spot, and the chamber closed. Certain 
of the Coprides exhibit, according to Fabre, some extremely 
exceptional features in their life-histories. The mother, instead 
of dying after opposition, survives, and sees the growth of her 
vnung to the perfect state, and then produces another generation. 
Xo similar case can be pointed out in Insects, except in the Social 
kinds: but from these the Coprides observed by Fabiv differ pro- 
foundly, inasmuch as the number of eggs produced by the mother 
i- extremely small; Copris hispanus, for instance, producing in 
eaeh of its acts of oviposition only one. two, or three __- 




FIG. 89. Scambaeu-s sneer. Portueal. 



COLEOPTERA CHAP. 



ii. The MELOLONTHIDES are probably almost as numerous as the 
Coprides, some 4000 species being already known. The larvae 
are believed to feed chiefly on roots. Melolontha vulgar is, the 
common cockchafer, is very abundant in some parts of Europe, 
and owing to this and to the great damage it causes, has attracted 
much attention. The memoir by Straus-Durckheim l on its 
anatomy is one of the classical works of Entomology. This In- 
sect is so injurious in some parts of France that money is paid 
by the local authorities for its destruction. M. Eeiset informs 
us that under this arrangement 867,173,000 perfect cockchafers, 
and 647,000,000 larvae were destroyed in the Seine-inferieure 
in the four years from the middle of 1866 to 1870. Unlike 
the Coprides, the larval life in Melolonthides is prolonged, and 
that of the imago is of brief duration. In Central Europe the 
life-cycle of the individual in M. r//A/"/'/x occupies three years, 
though in dry periods it may be extended to four years. In 
Scandinavia the time occupied by the development appears to be 
usually five years. The fertile female enters the ground and 
deposits its eggs in two or three successive batches of about 
fifteen each. The eggs swell as the development of the embryo 
progresses ; the larva emerges about five weeks after the eggs 
have been deposited, and is of relatively large size. When 
young the larvae can straighten themselves out and crawl, but 
when older they lose this power, and when above ground rest 
helplessly on their sides. In the winter they descend deeply 
into the earth to protect themselves from frost. The pupa 
state lasts only a few days, but after the final transformation the 
perfect Insect may remain motionless for as much as eight 
months underground before commencing its active life in the air. 2 
In the perfect state the Insect is sometimes injurious from the 
large quantity of foliage it destroys. Schiodte 3 considered that 
these larvae (and those of numerous other Scarabaeidae) stridu- 
late by rubbing certain projections on the stipes of the maxilla 
against the under-surface of the mandible. These surfaces appear, 
however, but little adapted for the purpose of producing sound. 

iii. The EUTELIDES number about 1500 species : there ;ire many 



1 Considerations y/^'/v/A's .>/// I'anatomie comparde des animautc articulds, etc., 
Paris 1828, 4to. xix. and 435 pp., and Atlas of ten (xx.) plates, and 36 pp. 
- Raspail. Me, n. SIK-. zool. France, vi. 1893, pp. 202-213. 
:l Ann. soc. ent. /''m/tce, (v.) iv. 1874. p. 39. 



v LAMELLICORNIA DYNASTIDES CETONIIDES 199 

Insects of brilliant metallic colours amongst them, but very little 
is known as to their life-histories. The larvae are very much 
like those of Melolonthides. 

iv. The DYNASTIDES are the smallest division in number of 
species, there being scarcely 1000 known ; but amongst them 
we find in the genera Dynastes and Megasoma some of the 
bniM'st of existing Insects. The horns and projections on the 
heads and prothoraces of some of the males of these Insects are 
truly extraordinary, and it does not appear possible to explain 
their existence by any use they are to their possessors. These 
structures are but little used for fighting. Baron von Hiigel 
informs the writer that in Java he has observed large numbers 
of Xylotrupes gideon; he noticed that the males sometimes carry 
the females by the aid of their horns ; but this must be an excep- 
tional case, for the shape of these instruments, in the majority of 
Dynastides, would not allow of their being put to this use. The 
development of these horns varies greatly in most of the species, 
but he did not find that the females exhibited any preference for 
the highly armed males. The fact that the males are very much 
larger than the females, and that the armature is usually confined 
to them, suggests, however, that some sexual reason exists for these 
remarkable projections. Many Dynastides possess organs of stridu- 
lation, consisting of lines of sculpture placed so as to form one or 
two bands on the middle of the propygidium, and brought into 
play by being rubbed by the extremities of the wing-cases. This 
apparatus is of a less perfect nature than the structures for the 
same purpose found in numerous other beetles. "We have no 
member of this sub-family in Britain, and there are scarcely a 
dozen in all Europe. Decaying vegetable matter is believed to 
be the nutriment of Dynastides. The European Oryctes nasicofnis 
is sometimes found in numbers in spent tan. The growth a::d 
development of the individual is believed to be but slow. 

v. The CETONIIDES are renowned for the beauty of their colours 
and the elegance of their forms; hence they are a favourite 
group, and about 1600 species have been catalogued. They ;nv 
specially fond of warm regions, but it is a peculiarity of the 
sub-family that a large majority of the species are found in the 
Old World ; South America is inexplicably poor in these Insects, 
notwithstanding its extensive forests. In this sub-family the 
mode of night is peculiar ; the elytra do not extend down the 



20O COLEOPTERA CHAP. 

sides of the body, so that, if they are elevated a little, the wings 
can be protruded. This is the mode of flight adopted by most 
Cetoniides, but the members of the group Trichiini fly in the 
usual manner. In Britain we have only four kinds of Cetoniides ; 
they are called Kose-chafers. The larvae of C.fioricola and some 
other species live in ants' nests made of vegetable refuse, and it 
is said that they eat the ants' progeny. Two North American 
species of Euphoria have similar habits. The group Cremasto- 
chilini includes numerous peculiar Insects that apparently have 
still closer relations with ants. Most of them are very aberrant 
as well as rare forms, and it has been several times observed in 
N'nrth America that species of Cremastochilus not only live in 
the nests of the ants, but are forcibly detained therein by the 
owners, who clearly derive some kind of satisfaction from the 
companionship of the beetles. The species of the genus Lomap- 
ft'i-a stridulate in a peculiar manner, by rubbing the edges of the 
hind femora over a striate area on the ventral segments. 



Series II. Adephaga or Caraboidea. 

All tli,- tarsi five-jointed ; antennae filiform, or nearly so; -mouth - 
l>iirf* //////>/// ilcrelvpi'il, flic outer lolie of tin' nui.'-iUn /n <///// 
nl inni/x di ri<l <-<! into ii I ii-ti-jiiiiiti-il pa/juts ; *//j>j>u,'ts of the 
liiliinl palpi il< rt'liipnl ^.s joints of tlie palpi, and in so///' 
cases approximate at their bases. Alxlunn n n-itli tin' imposed 
* <j me nix inn more in number at the sides than a/u//</ the 
in 'nl /lie, the number Iciinj nxnall/i : fi re along the midill<, */./ 
nt iifh side.. 

THIS extensive series includes the tiger-beetles, ground-beetles, 
and true water -beetles ; it consists of six families, and forms 
a natural assemblage. It is sometimes called Garni vora or 
Filicornia. The exceptions to the characters we have mentioned 
are but few. The supports of the labial palpi are frequently 
covered by the mentum, and then the palpi appear three-jointed; 
but when the joint-like palpiger is not covered these palps appear 
four-jointed. As a rule, approximation of these supports is indica- 
tive of high development. In some of the lower forms the trophi 
remain at a lower stage of development than is usual. This is 
especially the case with the genus Amphizoa, which forms of 



v ADEPHAGA TIGER-BEETLES 2OI 

itself the family Amphizoidae. The Bombardier-beetles make an 
exception as regards the abdominal structure, for in some of them 
no less than eight segments are visible, either along the middle 
line or at the sides of the venter. In Hydroporides (one of the 
divisions of Dytiscidae) the front and middle feet have each only 
four joints. Many naturalists unite the Gyrinidae with the 
Adephaga, and a few also associate with them the Paussidae and 
Ehyssodidae ; but we think it better at present to exclude all 
these, though we believe that both Paussidae and Rhyssodidae 
will ultimately "be assigned to the series. The larvae are usually 
very active, and have a higher development of the legs than is 
usual in this Order. Their tarsi possess two claws. 

Fam. 4. Cicindelidae (Tiger -beetles). Clypeus extending 
laterally in front of the insertion of the antennae. Lower lip with 
the palpi usually yreatli/ developed, but with the liijula and pam- 
f/lossae very much reduced, often .sr^/w/// to be detected. M/i.i'illni' 
with the outer lobe form in*/ a tiro-jointed palp} the inner lobe 
elongate, furnished t the tip with a hook-like process, which is 
usually articulated by a joint ////// f//c lobe itself. The tiger-beetles 
are very active Insects, running with extreme speed, and some- 
times flying in a similar manner ; they are all predaceous, and 
amongst the most voracious and fierce of the carnivorous beetles, 
so that they well deserve their name. Bates, speaking of the 
Amazonian Megacephala, says " their powers of running exceed 
anything I have ever observed in this style of Insect locomotion ; 
they run in a serpentine course over the smooth sand, and when 
closely pursued by the hand they are apt to turn suddenly back 
and thus baffle the most practised hand and eye." He further 
says that the species he observed (being of diverse colours) 
a "'reed in colour with the general colours of the " locale thev 

O f 1 

inhabit." The larvae of Cicindelidae live in deep burrows, sink- 
ing more or less vertically into the ground, and in these they 
take up a peculiar position, for which their shape is specially 
adapted; the head and prothorax are broad, the rest of the body 
slender, the fifth segment of the abdomen is furnished on the 
back with a pair of strong hooks ; the ocelli on the sides of the 
head are very perfect. Supporting itself at the top of the burrow 
by means of these hooks and of its terminal tube, the larva blocks 
the mouth of the burrow with its large head and prothorax, and 

1 In Theratides this outer lobe is in a rudimentary state, like a seta. 



2O2 



COLEOPTERA 



CHAP. 



in this position waits for its prey. This consists of Insects that 
may alight on the spot or run over it. AVhen an Insect ventures 
within reach, the head of the larva is thrown back with a rapid 
jerk, the prey is seized by the long sharp mandibles, dragged to 
the bottom of the burrow and devoured. The burrows are often 
more than a foot deep, and are said to lie excavated by the larva 
itself, which carries up the earth on the shovel-like upper surface 
of its head. The female tiger-beetle is endowed with powerful 
and elongate excavating instruments at the termination of the 
body, and it is probable that when placing the egg in the earth 

she facilitates the future opera- 
tions of the larva by forming 
the outlines of the burrow. Ex- 
tremely few larvae of Cicin- 
delidae are known, but they all 
exhibit the type of structure 
mentioned above, and apparently 
have similar habits. Our little 
British Cicindd, most of which 
are so active on the wing, agree 
in these respects with the African 
species of Jbnificora, which are 
entirely apterous, and are the 
largest of the Cicindelid.ie. Per- 

Fio. 90. (Jicindela hybrida. Britain. 

A, larva (after Schiodte) ; B, imago, inguey found a breeding-ground 

of M, tultci'culata near Kimberley ; 

the larvae were living in the usual Cicindelid manner: but the 
ground was so hard that he was not able to investigate the 
burrows, and there were but few Insects that could serve as food 
in the neighbourhood. 

The Ciciudelidae, although one of the smaller families of Cole- 
optera, now number about 1400 species; of these about one-half 
belong to the great genus Cicindela, to which our four British 
representatives of the Cicindelidae are all assigned. There is no 
general work of much consequence on this important family, and 
its classification is not thoroughly established. 1 

Tiger-beetles display considerable variety of structure, especially 
iis regiirds the mouth, which exhibits very remarkable develop- 

1 Tin- first portion of a classification of (.'icindrliiUe by Dr. "\Valther Horn, 
J'i'i-isi"n ili' i' Cicindeliden, Berlin, 1898, lias appeared since this was written. 




CICINDELIDAE TIGER-BEETLES 



203 



iin'iits of the palpi and labrum (Fig. 91). The tiger-beetles, 
like most other Insects that capture living prey, do not consume 
their victims entire, but subsist chiefly on the juices they 
squeeze out of them ; the hard and innutritions parts are rejected 
after the victim has been thoroughly lacerated and squeezed: the 
mouth forms both trap and 
press ; the palpi spread out 
in order to facilitate the 
rapid engulfing of a victim, 
then close up under it and 
help to support it in the 
mouth ; while the labrum 
above closes the cavity in 
the other direction. The 
mouth itself is a large cavity 
communicating very freely 
with the exterior, but so 
completely shut off from the 
following parts of the ali- 
mentary canal that it is 
difficult to find the orifice of 
communication ; the labium 




FIG. 91. Mouth - parts of tiger - beetles. A. 
Profile of Po0Wioso!ffl sp. (Madagascar) : a, 
antenna ; b, labial palp ; c, maxillary palp ; 
d, palpifonn lobe of maxilla ; e, mandible ; /. 
labrum. B, Section of head of Manticora, 



heing 



(South Africa) : a, front of upper 
part of head-capsule ; b, gula ; c, teutorium ; 
(/, eye ; e, labrum ; /, left mandible ; g, max- 
illa ; /;, maxillary palp ; i, labial palp ; f,; 
support of this palp ; /, labium. 



much modified to 
form the posterior wall. 
For the capture of the prey, 
always living but of various 
kinds, a mechanism with great holding power and capable of 
rapid action is required. The mouth of the terrestrial Manticora 
(Fig. 91, B), exhibits great strength ; some of the chitinous parts 
are extremely thick, the mandibles are enormous, the palpi, how- 
ever, are comparatively low in development. In the arboreal 
genus Pogonostoma the palpary structures (Fig. 91, A) attain a 
development scarcely equalled elsewhere in the Insect world. The 
great majority of the Cicindelidae are inhabitants of the warmer, 
or of the tropical regions of the world, and very little is known as 
to their life-histories ; they show great diversity in their modes 
of hunting their prey. Some are wingless ; others are active on 
the wing ; and of both of these divisions there are forms that are 
found only on trees or bushes. Some, it is believed, frequent 
onlv the mounds of Termites. The characteristic feature common 



204 



COLEOPTERA 



CHAl'. 



to all is great activity and excessive wariness. The genus 
Pogonostoma, to which we have already alluded, is confined to 
Madagascar, where the species are numerous, but are rare in 
collections on account of the difficulty of securing them. Eaffray 
informs us that certain species frequent the trunks of trees, up 
which they run in a spiral manner on the least alarm. The only 
way he could obtain specimens was by the aid of an assistant ; 
the two approached a tree very quietly from opposite sides, and 
when near it, made a rush, and joined hands as high up the 
trunk as they could, so as to embrace the tree, when the Pogonos- 
toma fell to the ground and was captured. 

Fam. 5. Carabidae (Ground-beetles). Clypeus not extending 
lateral/// in front of the antennae. Maxillae iritli the outer lobe 
destitute of an articulated hook at the ///<. Antennae covered 
(except tin' band joints} with a minute pubescence. Hind legs not 
very different from the middle pair, formed for running, as ttst/al 
in beef fry. This is one of the largest and most important of the 

families of Coleoptera, in- 
cluding as it does 12,000 
or 13,000 described 
species. In this country 
( 'a ra b idae are nearly 
entirely terrestrial in 
habits, and are scarcely 
ever seen on the wing : 
many of the species indeed 
have merely rudimentary 
wings ; in the tropics 
there are, however, many 
arboreal forms that take 
wing with more or less 
alertness. The larvae (Fig. 
( .>2, A) are usually elon- 
gate in form and run 
freely : they may be known 
by their tarsi ending in two claws, by the exserted, sharp, calliper- 
like mandibles, by the body ending in two processes (sometimes 
jointed) and a tube of varying length projecting backwards. 
The pupae usually have the hind pair of legs so arranged that 
the tips of the tarsi project behind, beyond the extremity of the 





Sdiiodte) ; B, imago. 



A, Larva (after 
Britain. 



v ADEPHAGA CARABIDAE 2O5 

body. The Carabidae are carnivorous and predaceous both as 
larvae and perfect Insects ; they attack living Insects, worms, or 
other small, soft creatures, but do not disdain dead specimens. 
Some species of Carabus, found in North Africa where snails 
abound, are specially formed for attacking these molluscs, having 
the head long and slender so that it can be thrust into the shell 
of the snail. A few species have been detected eating growing corn, 
and even the young seeds of some Umbelliferae ; these belong chiefly 
to the genera Harpalus, Zabrus, and Amara. Some species of the 
abundant genera Pterostichus and Harpalus, are said to be fond of 
ripe strawberries. The most anomalous forms of Carabidae are the 
Pseudomorpliides, a sub-family almost peculiar to Australia, the 
members of which live under bark, and have but little resemblance 
to other Carabids owing to their compact forms and continuous 
outlines. The genus Mormolyce is one of the wonders of the Insect 
world on account of the extraordinary shape of its members ; the 
sides of the elytra form large crinkled expansions, and the head 
is unusually elongate. These Insects live on the underside of 
fallen trees in the Malay Archipelago and Peninsula ; no reason 
whatever can be at present assigned for their remarkable shape. 

There are a considerable number of blind members of this 
family : some of them live in caverns ; these belong chiefly to 
the genus Anophthcdmus, species of which have been detected in 
the caves of the Pyrenees, of Austria, and of North America. 
It has been shown that the optic nerves and lobes, as well as tin- 
external organs of vision, are entirely wanting in some of these 
cave Carabidae ; the tactile setae have, however, a larger develop- 
ment than usual, and the Insects are as skilful in running as if 
they possessed eyes. Anophthalmus is closely related to our 
British genus Trechvs, the species of which are very much given 
to living in deep crevices in the earth, or under large stones, and 
have some of them very small eyes. In addition to these 
cavernicolous Anophthalmus, other blind Carabidae have been 
discovered during recent years in various parts of the world, 
where they live under great stones deeply embedded in the 
earth : these blind lapidicolous Carabidae are of extremely 
minute size and of most sluggish habits ; the situations in 
which they are found suggest that many successive generations 
are probably passed under the same stone. Not a single 
specimen has ever been found above ground. The minute 



206 COLEOPTERA 



CHAP. 



Carabids of the genus ACJIKS, that pass a large part of their 
lives under stones below high-water mark (emerging only when 
the tide uncovers them), on the borders of the English Channel 
and elsewhere, are very closely allied to these blind Insects, and 
have themselves only very small eyes, which, moreover, according 
to Hammond and Miall, are covered in larger part by a peculiar 
shield. 1 A few Carabidae, of the genera 6V///>/V.s and Orthogonius, 
are believed to live in the nests of Termites. Savage found the 
larva of G. sculptilis in the nests of Tcrmes IcHicosus; it has 
been described by Horn, and is said to bear so great a resem- 
blance to young queens of the Termites as to have been mistaken 
for them. 2 Mr. Haviland found Rhopalomelus angusticoUis in 
Termites' nests in South Africa. Peringuey states that it emits 
a very strong and disagreeable odour. It is probable that it 
preys on the Termites, and this also is believed to be the habit 
of the Ceylonese Hdluodes taprobanae. Some species of the 
Mediterranean genus Siagona stridulate by means of a file on 
the under surface of the prothorax, rubbed by a striate area, 
adapted in form, on the anterior femora. 

A valuable memoir on the classification of this important 
family is due to the late Dr. (I. H. Horn ; 3 he arranges Carabidae 
in three sub-families ; we think it necessary to add a fourth for 



1. Middle coxal cavities enclosed externally by the junction of the nieso- and 

nieta-sternum ; neither epimeron nor episternum attaining tin- 

cavity. 
Head beneath, "with a deep groove on each side near the eye for 

the reception of the antennae or a part thereof. 

Sub-fam. 3. PSEUDOMORPHIDES. 
Head without antennal grooves. Sub-fam. 2. HARPALIDES. 

2. Middle coxal cavities attained on the outside by the tips of the episterna 

and epimera. Sub-fam. 4. MORMOLYCIDES. 

3. Middle coxal cavities attained on the outside by the tips of the epinirm, 

but not by those of the episterna. Sub-fain. 1. CARABIDES. 

These four sub-families are of extremely different extent and 
nature. The Harpalides are the dominant forms, and include 
upwards of 10,000 known species; while the various tribes 
into which the sub-family is divided include, as a rule, each many 



i/ of aquatic Insects, 1895, p. 376. 
2 Tr. Anur. cnt. Soc. xv. 1888, p. 18. 
<>j>. cit. v. 1881, p. 91 ; cf. Sharp, Tr. cut. Soc. London, 1S82, p. 61. 



ADEPHAGA AMPHIZOIDAE PELOBIIDAE 



207 



genera ; the Carabides are next in importance, with upwards of 
2000 species, but are divided into a comparatively large number of 
tribes, each of which averages a much smaller number of genera 
than do the tribes of Harpalides ; Pseudomorphides includes only 
about 100 species ; and Mormolycides consists of the single genus 
Mormolycc with three species. 

Fam. 6. Amphizoidae. Antennae destitute of pubescence : 
outer lobe of maxilla not jointed; metasternum with a short 
transverse impressed line on the middle behind. Hind legs slender, 
not formed for swimming. This family is limited to the genus 
Ampliizoa ; the species of 
which may be briefly de- 
scribed as lowly organised 
Carabidae that lead an 
aquatic life. The geo- 
graphical distribution is 
highly remarkable, there 
being but three species, two 
of which live in Western 
Itforth America, the third 
in Eastern Tibet. The 
habits of American Am- 
phizoa are known ; they pass 
a life of little activity in 
very cold, rapid streams ; 

FIG. 93. Amphizoa lecontei. North America, 
they do not SWllll, but A, Larva ; B, imago. 

cling to stones and timber. 

The larva was recently discovered in Utah by Messrs. Hubbard 
and Schwarz : : it has the same habits as the perfect Insect, 
and in general form resembles the larvae of the genus Cardbus ; 
but it has no terminal tube to the body, the abdomen consisting 
of eight segments and a pair of short terminal appendages ; the 
spiracles are obsolete, with the exception of a pair placed near to 
one another at the termination of the eighth abdominal segment. 
As regards the mouth this larva is Carabid, as regards the 
abdomen and stigmata Dytiscid of a primitive type. 

Fam. 7. Pelobiidae. Antennae destitute of pubescence : outer 
lobe of maxilla jointed, metasternum with a short transverse 
impressed line on the middle behind. Hind legs rather slender, 

1 P. ent. Soc. Washington, ii. 1892, p. 341. 




208 



COLEOPTERA 



CHAP. 



formed for swimming, the tarsi longer tlt/m lite tililae. This family 
is limited to the one genus Pelolnus (Hygrolna of some authors). 
Like Ani]i}ii~oa, to which it is in several respects analogous, it 
lias a singular geographical distribution ; there are only four 
known species, one lives in Britain and the Mediterranean region, 
one in Chinese Tibet, two in Australia, Pelobivs may be briefly 

t/ *' 

described as a Carabid adapted to a considerable extent for 
living in and swimming about in water ; differing thus from 




FIG. 94. Pelobius tardus. Britain. A, Young larva ; B, adult larva ; C, imago. 

(A ami B alter Scliiodte.) 

Amphizoa, which has no special adaptation for swimming. The 
larva of Pelobius is remarkable ; it breathes by means of 1 iranchial 
filaments on the under surface of the body, the spiracles being 
present, though those of the abdomen are very minute and the 
others small. The head is very large, the mandibles are not 
tube-like, the food being taken after the manner of the Carabidae ; 
the 8th abdominal segment ends in three long processes ; the 
small 9th segment is retracted beneath them. The adult 7V/W^Vs 
inriliix is remarkable for its loud stridulation. The sound is pro- 
duced by an apparatus described correctly by Charles Darwin: 1 

1 Descent of Mi in,, i. 1890, p. 338; The views of Landois aud Recker, Arch. f. 
Naturgesch. Ivii. 1, 1891, p. 101, are erroneous. 



V 



ADEPHAGA HALIPLIDAE 



209 



there is a file on the inside of the wing-cases, and the Insect 
turns up the tip of the abdomen and scrapes the tile therewith. 
The Insects are called squeakers in the Covent Garden market, 
where they are sold. 

Fam. 8. Haliplidae. Antennae hire, ten - jointed ; meta- 
sternum marked l>y a transverse line; posterior coxae 'prolonged 
as plates, covering a, large 
part of the lower surface 
of the abdomen ; the 
slender, hut clubbed, hind 
femora move between these 
plates and the abdomen. 
The Haliplidae are 
aquatic, and are all small, 
not exceeding four or five 
millimetres in length. 
The ventral plates are 
peculiar to the Insects of 
this family, but their func- 
tion is not known. The 
larvae are remarkable on 
account of the fleshy pro- 
cesses disposed on their 
bodies ; but they exhibit 
considerable variety in 
this respect ; their man- 
dibles are grooved so that 
they suck their prey. In 
the larva of Haliplus, 
according to Schiodte, 
there are eight pairs of 
abdominal spiracles, but 
in Cnemidotus (Fig. 95, 

-r, N , -, , FIG. 95. Cnemidotus caesus. England. A, Imago ; 

B), there are 110 spiracles, B , larva, highly magnified. (After Schiodte.) 

and air is obtained by 

means of a trachea traversing each of the long filaments. The 
Insects of these two genera are so similar in the imaginal instar 
that it is well worthy of note that their larvae should be distin- 
guished by such important characters. Haliplidae is a small 
family consisting of three genera, having about 100 species; 

VOL. VI P 




210 



COLEOPTERA 



CHAP. 



it is very widely distributed. We have 13 species in Britain, 
all the genera being represented. 

Fam. 9. Dytiscidae (Water-beetles). Antennae bare; hind 
legs formed for sunmming, not capable of ordinary walking : nieta- 
sternum without a- transverse line across it ; behind closely united 
with the extremely large coxae. Outer lobe of maxilla forming a 
two-jointed palpus. ' The Dytiscidae, or true water-beetles, are of 
interest because unlike the aquatic Neuroptera they exist in 

water in both the 
larval and imaginal 
instars ; nevertheless 
there is reason for sup- 
posing that they are 
modified terrestrial 
Insects : these reasons 
are (1) that in their 
general organisation 
they are similar to 
the Carabidae, and 
they drown more 
quickly than the 
majority of land 
beetles do ; (2) though 
the larvae are very 
different from the 
larvae of terrestrial 
beetles, yet the ima- 
ginal instars are much 
less profoundly 
changed, and are capable of existing perfectly w r ell on land, and 
of taking prolonged nights through the air ; (3) the pupa is, so 
far as known, always terrestrial. The larvae and images are 
perfectly at home in the water, except that they must come to 
the surface to get air. Some of them are capable, however, when 
quiescent, of living for hours together beneath the water, but 
there appears to be great diversity in this respect. 1 The hind 
pair of legs is the chief means of locomotion. These swiinming- 
legs (Fig. 97) are deserving of admiration on account of their 
mechanical perfection ; this, however, is exhibited in various 

1 Src J. Linn. Soc. Zool. xiii. 1876, p. 161. 




A 



FIG. 96. Cybister roesdi ( = laterima,rginalis De G.) 
Europe. A, Larva (after Sclriudte) ; B, <J imago. 



ADEPHAGA WATER-BEETLES 



2 I I 



B 



degrees, the legs in the genera Dytiscus and Hydroporus being 
but slender, while those of Cybister are so broad and powerful, that 
a single stroke propels the Insect for a considerable distance. 

The wing-cases fit perfectly to the body, except at the tip, so 
as to form an air-tight space between themselves and the back of 
the Insect ; this space is utilised as a reservoir for air. When 
the Dytiscus feels the necessity for air it rises to the surface and 
exposes the tip of the body exactly at the level of the water, 
separating at the same time the abdomen from the wing-cases 
so as to open a broad chink at the spot where the parts were, 
during the Insect's submersion, so well held together as to be air- 
and water - tight. The ter- 
minal two pairs of spiracles 
are much enlarged, and by 
curving the abdomen the 
beetle brings them into con- 
tact with the atmosphere ; 
respiration is effected by this 
means as well as by the store 
of air carried about under the 
wing - cases. The air that 
enters the space between the 
elytra and body is shut in 
there when the Insect closes 
the chink and again dives beneath the water. The enlargement 
of the terminal stigmata in Dytiscus is exceptional, and in forms 
more highly organised in other respects, such as Cybister, these 
spiracles remain minute ; the presumption being that in this 
case respiration is carried 011 almost entirely by means of the 
supply the Insect carries in the space between the elytra and the 
base of the abdomen. 1 The structure of the front foot of the male 
Dytiscus, and of many other water-beetles, is highly remarkable, 
the foot being dilated to form a palette or saucer, covered beneath 
by sucker-like structures of great delicacy and beauty ; by the aid 
of these the male is enabled to retain a position on the female for 
many hours, or even days, together. Lowne has shown that the 




' Hind- or swimming-leg of Cybister 
A, The whole leg detached ; 
B, the movable parts in the striking posi- 
tion, a, Coxa ; b, trochanter ; c, lemur ; 
d, tibia ; e, last joint of tarsus. 



1 For many particulars as to respiration of Dytiscus, and peculiarities of the 
larva see Miall, Aquatic Insects, 1895, pp. 39, etc. (In the figure given on p. (30 
the large stigma on the terminal segment of the abdomen is omitted, though it is 
referred to in the text.) 



2 1 2 COLEOPTERA CHAP. 

suckers communicate with a sac in the interior of the foot contain- 
ing fluid, which exudes under pressure. As the portions of the 
skeleton of the female on which these suckers are brought to 
bear is- frequently covered with pores, or minute pits, it is prob- 
able that some correlation between the two organisms is brought 
about by these structures. The females in many groups of 
Dytiscidae bear on the upper surface of the body a peculiar 
sculpture of various kinds, the exact use of which is unknown ; 
in many species there are two forms of the female, one possessing 
this peculiar sculpture, the other nearly, or quite, without it. 
The larvae of Dytiscidae differ from those of Carabidae chiefly 
by the structure of the mouth and of the abdomen. They are 
excessively rapacious, and are indeed almost constantly engaged 
in sucking the juices of soft and small aquatic animals, by no 
means excluding their own kind. The mode of suction is not 
thoroughly known, but so far as the details have been ascertained 
they are correctly described, in the work on aquatic Insects, by 
Professor Miall, we have previously referred to ; the mandibles 
are hollow, with a hole near the tip and another at the base, and 
being sharp at the tips are thrust into the body of a victim, and 
then by their closure the other parts of the mouth, which are 
very beautifully constructed for the purpose, are brought into 
fitting mechanical positions for completing the work of emptying 
the victim. Nagel states that the larva of Dytiscus injects a 
digestive fluid into the body of its victim, and that this fluid 
rapidly dissolves all the more solid parts of the prey, so that the 
rapacious larva can easily absorb all its victim except the 
insoluble outer skin. The abdomen consists of only eight 
segments, and a pair of terminal processes ; the stigmata are all 
more or less completely obsolete according to species with the 
exception of the pair on the eighth segment at the tip of the 
body ; the terminal segments are frequently fringed with hairs, 
that serve not only as means of locomotion, but also to float the 
pair of active stigmata at the surface when the creature rises to 
get air. Although the larvae of Dytiscidae are but little known, 
yet considerable diversity has already been found. Those of 
Hyphydrus and some species of Hydroporus have the front of the 
head produced into a horn, which is touched by the tips of the 
mandibles. 

Dytiscidae are peculiar inasmuch as they appear to flourish 



v POLYMORPHA CLAVICORNS, SERRICORNS, ETC. 2 I 3 

Lest in the cooler waters of the earth. Lapland is one of the 
parts of Europe richest in Dytiscidae, and the profusion of species 
in the tropics compared with those of Europe is not nearly so 
great as it is in the case of most of the other families of 
Coleoptera. About 1800 species are at present known, and we 
have rather more than 100 species in Britain. 1 

Series III. Polymorpha. 

Ante'nnae frequently either thicker at the tip (clavicorn) or serrate 
along their inner edge (serricorn) ; but these characters, as 
well as the number of joints in the feet and oilier points, are 
very variable. 

Upwards of fifty families are placed in this series ; many of 
these families are of very small extent, consisting of only a few 
species ; other families of the series are much larger, so 
that altogether about 40,000 species speaking broadly, about 
one-fourth of the Coleoptera are included in the series. We 
have already (p. 189) alluded to the fact that it is formed by 
certain conventional series, Clavicornia, Serricornia, etc, united, 
because it has hitherto proved impossible to define them. 

Fam. 10. Paussidae. Antennae of extraordinary form, usually 
two-jointed, sometimes six- or ten-jointed. Elytra elongate, but trun- 
cate behind, leaving the pygidium exposed. Tarsi five-jointed. The 
Paussidae have always been recognised as amongst the most 
remarkable of beetles, although they are of small size, the largest 
attaining scarcely half an inch in length. They are found only 
in two ways ; either in ants' nests, or on the wing at night. 
They apparently live exclusively in ants' nests, but migrate 
much. Paussidae usually live in the nests of terrestrial ants, but 
they have been found in nests of Cremastogaster in the spines of 
Acacia fistulosa. They have the power of discharging, in an 
explosive manner, a volatile caustic fluid from the anus, which is 
said by Loinan to contain free iodine. Their relations to the 
ants are at present unexplained, though much attention has been 
given to the subject. When observed in the nests they frequently 
appear as if asleep, and the ants do not take much notice of 
them. On other occasions the ants endeavour to drag them into 
the interior of the nest, as if desirous of retaining their company : 

1 For classification and structure see Sharp, "On Dytiscidae," Sci. Trans. H. 
Dublin Soc. (2) ii. 1882. 



214 



COLEOPTERA 



CHAP. 



the Paussus then makes no resistance to its hosts ; if, however, it 
be touched, even very slightly, by an observer, it immediately 
bombards : the ants, as may be imagined, do not approve of this, 
and run away. Nothing has ever been observed that would lead 
to the belief that the ants derive any benefit from the presence 
of the Paussi, except that these guests bear on some part of the 
body frequently the great impressions on thepronotum patches 
of the peculiar kind of pubescence that exists in many other kinds 
of ants'-nest beetles, and is known in some of them to secrete a 
substance the ants are fond of, and that the ants have been seen 

to lick the beetles. On the other hand, the 
Paussi have been observed to eat the eggs 
and larvae of the ants. The larva of Paussus 
is not known, 1 and E affray doubts whether 
it lives in the ants' nests. There are about 
200 species of Paussidae known, Africa, Asia 
and Australia being their chief countries ; 
one species, P. favieri, is not uncommon 
in the Iberian peninsula and South France, 
and a single species was formerly found in 
Brazil. The position the family should 
FIG. 98. Paussus cepha- occupy has been much discussed ; the only 
(After Raffray.) forms to which they make any real ap- 

proximation are Carabidae, of the group 

Ozaenides, a group of ground beetles that also crepitate. Bur- 
meister and others have therefore placed the Paussidae in the 
series Adephaga, but we follow Eaffray's view (he being the most 
recent authority on the family), 2 who concludes that this is 
an anomalous group not intimately connected with any other 
family of Coleoptera, though having more affinity to CarabkLie 
than to anything else. The recently discovered genus Proto- 
paussus has eleven joints to the antennae, and is said to come nearer 
to Carabidae than the previously known forms did, and we may an- 
ticipate that a more extensive knowledge will show that the family 
may find a natural place in the Adephaga. The description of 
the abdomen given by E affray is erroneous ; in a specimen of the 
genus Arthropterus the writer has dissected, lie finds that there 
1 Descriptions of larvae that may possibly be those of Paussicls have been pub- 
lished by Xambeu, Ann. Soe. Linn. Lyon, xxxix. 1892, p. 137, and Erich.son, Arch. 
Xatiirycsch. xiii. 1847, p. 275. 

- Arch. Mus. Paris (2), viii. and ix. 1SS7. 




POLYMORPHA WHIRLIGIG-BEETLES 



215 



are five ventral segments visible along the middle, six at the 
sides, as in the families of Adephaga generally. There is said 
to be a great difference in the nervous systems of Carabidae and 
Paussidae, but so little is known on this point that we cannot 
judge whether it is really of importance. 

Fam. 11. Gyrinidae (Whirligig beetles). --Antennae very 
short ; four eyes ; middle and hind legs forming short broad 
paddles ; abdomen with six segments visible along the middle, seven 
/ong each side. These Insects are known to all from their habit 
of floating lightly on the surface of water, and performing graceful 
complex curves round 
one another without 
colliding ; sometimes 
they may be met with 
in great congregations. 
They are admirably con- 
structed for this mode 
of life, which is com- 
paratively rare in the 
Insect world ; the 
Hydrometridae amongst 
the bugs, and a small 
number of different 
kinds of Diptera, being 
the only other Insects 
that are devoted to a 
life on the surface of the 
waters. Of all these, 
Gyrinidae are in their 
construction the most 
adapted for such a 
career. They are able 
to dive to escape danger, 




FIG. 99. A, Larva of Gyrinus (after Schiodte) ; B, 
under side of Gyrinus sp. (after Ganglbauer). 1, 
Prosternum ; 2, anterior coxal cavity ; 3. raeso- 
thoracic episternum ; 4, mesoepimeron ; 5, nieso- 
sterrmm ; 6, metathoracic episternum ; 7, middle 
coxal cavity ; 8, metasternum ; 9, hind coxa ; 10, 
ventral segments. [X.B. The first ventral segment 
really consists, at each side, of two segments united ; 
this may be distinctly seen in many Gyrinidae.] 



and they then carry with them a small supply of air, but do not 
stay long beneath the surface. Their two hind pairs of legs are 
beautifully constructed as paddles, expanding mechanically when 
moved in the backward direction, and collapsing into an extremely 
small space directly the resistance they meet with is in the other 
direction. The front legs of these Insects are articulated to the 
thorax in a peculiar direction so that their soles do not look 



2l6 COLEOPTERA CHAP. 

downwards but towards one another ; hence the sensitive ad- 
hesive surface used during coupling is placed on the side of the 
foot, forming thus a false sole : a remarkable modification other- 
wise unknown in Insects. They breathe chiefly by means of the 
very large metathoracic spiracles. 

The larvae (Fig. 99, A) are purely aquatic, and are highly 
modified for this life, being elongate creatures, with sharp, 
mandibles and nine abdominal segments, each segment bearing 
on each side a tracheal branchia ; these gills assist to some extent 
in locomotion. The stigmata are quite obsolete, but the terminal 
segment bears four processes, one pair of which may be looked on 
as cerci, the other as a pair of gills corresponding with the pair 
on each of the preceding segments. The mandibles are not 
suctorial, but, according to Meinert, possess an orifice for the 
discharge of the secretion of a mandibular gland. Gyrinidae 
are chiefly carnivorous in both the larval and imaginal instars. 
Fully 300 species are known; they are generally distributed, 
though wanting in most of the islands of the world except those 
of large size. The finest forms are the Brazilian Enliydrus and 
the Porrorhynchus of tropical Asia. 1 In Britain we have nine 
species, eight of Gyrinus, one of Orectochilus ; the latter form is 
rarely seen, as it hides during the day, and performs its rapid 
gyrations at night. 

The Gyrinidae are one of the most distinct of all the families 
of Coleoptera : by some they are associated in the Aclephagous 
series ; but they have little or no affinity with the other mem- 
bers thereof. Without them the Adephaga form a natural series 
of evidently allied families, and we consider it a mistake to force 
the Gyrinidae therein because an objection is felt by many tax- 
ouomists to the maintenance of isolated families. Surely if 
there are in nature some' families allied and others isolated, it is 
better for us to recognise the fact, though it makes our classifi- 
cations look less neat and precise, and increases the difficulty of 
constructing " tables." 

Fam. 12. Hydrophilidae.--T#m five-jointed, the first joint 
in many cases so small as to be scarcely evident: antennae 
short, of less than eleven joints, not filiform, but consisting of 

1 For classification and monograph of the family, see Regimbart, Ann. Svc. ent. 
France, 1882, 1883, and 1886. For a catalogue, Severin, Ann. Soe. cnt. Hclyiqi" , 
xxxiii. 1889. 



v POLYMORPHA HYDROPHILIDAE 2 I / 

three parts, a basal part of one or two elongate joints, an inter- 
mediate part of two or more small joints, and an apical part of 
lui-ger (or at any rate broader} joints, which are pubescent, the others 
being bare. Outer lobe of maxillae usually complex, but not at 
all palpiform, maxillary palpi of ten very long ; the parts of the 
labium much concealed behind the mentum, the labial palpi very 
widely separated. Hind coxae extending the width of the 
body, short, the lamina interior small in comparison with the 
lamina exterior. Abdomen of Jive visible segments. The Hydro- 
philidae are an extensive family of beetles, unattractive in colours 
and appearance, and much neglected by collectors. A large part 
of the family live in water, though most of them have only 
feeble powers of aquatic locomotion, and the beetles appear 
chiefly to devote their attention to economising the stock of air 
each individual carries about. The best known forms of the 
family are the species of Hydrophilus. They are, however, very 
exceptional in many respects, and are far more active and pre- 
daceous than most of the other forms. Much has been written 
about Hydrophilus piceus, one of the largest of British beetles. 
This Insect breathes in a most peculiar manner : the spiracles 
are placed near bands of delicate pubescence, forming tracts that 
extend the whole length of the body, and in this particular 
species cover most of the under surface of the body ; these 
velvety tracts retain a coating of air even when the Insect is 
submerged and moves quickly through the water. It would 
appear rather difficult to invent a mechanism to supply these 
tracts with fresh air without the Insect leaving the water ; but 
nevertheless such a mechanism is provided by the antennae of 
the beetle, the terminal joints of which form a pubescent scoop, 
made by some longer hairs into a funnel sufficiently large to 
convey a bubble of air. The Insect therefore rises to the sur- 
face, and by means of the antennae, which it exposes to the air, 
obtains a supply with which it surrounds a large part of its 
body ; for, according to Miall, it carries a supply on its back, 
under the elytra, as well as on its ventral surface. From the 
writer's own observations, made many years ago, he inclines to 
the opinion that the way in which the Hydrophilus uses the 
antennae to obtain air varies somewhat according to circumstances. 
Many of the members of the sub-family Hydrophilides con- 
struct egg-cocoons. In the case of Hydrophilus piceus, the boat- 



218 



COLEOPTERA 



CHAP. 



like structure is provided with a little mast, which is supposed by 
some to be for the purpose of securing air for the eggs. Helo- 
cliares and Spercheus (Fig. 100) carry the cocoon of eggs attached 

to their own bodies. Philydrus constructs, 
one after the other, a number of these 
egg-bags, each containing about fifteen 
eggs, and fixes each bag to the leaf of 
some aquatic plant ; the larvae as a rule 
hatch speedily, so that the advantage of 
the bag is somewhat problematic. 

The larvae of the aquatic division of 
the family have been to a certain extent 
studied by Schiodte and others ; those of 
the Sphaeridiides the terrestrial group 
of the family are but little known. All 
the larvae, seem to be predaceous and 
carnivorous, even when the imago is of 
vegetable -feeding habits ; and Dumi'ril 
FIG. 100. Spercheus emar- sta tes that in Hydrous caroboides the 

gmatus ?. Britain. A, 

Upper surface of beetle ; alimentary canal undergoes a great 
B, under surface of abdo- c ] lange at t j ie period of metamorphosis, 

nifiii. with the egg -sac 




B 



men. 



ruptured and some of the becoming very elongate in the adult, 
JS escaping. though in the larva "it was short, The 

legs are never so well developed as they are in the Adephaga, 
the tarsi being merely claw -like or altogether wanting; the 
mandibles are never suctorial. The respiratory arrangements 
show much diversity. In most of the Hydrophilides the process 
is carried on by a pair of terminal spiracles on the eighth 
abdominal segment, as in Dytiscidae, and these are either 
exposed or placed in a respiratory chamber. In Bcrosus the 
terminal stigmata are obsolete, and the sides of the body bear 
long branchial filaments. Cussac says that in fycrcliev.s (Fig. 
101) there are seven pairs of abdominal spiracles, and that the 
larva breathes by presenting these to the air ; l but Schiodte 
states that in this form there are neither thoracic nor abdominal 
spiracles, except a pair placed in a respiratory chamber on the 
eighth segment of the abdomen, after the manner described by 
Miall as existing in Hydrol'm*. No doubt Cussac was wrong in 
supposing the peculiar lateral abdominal processes to be stig- 

1 Ann. Soc. cut. France, xxi. 1852, p. 619. 



POLYMORPHA HYDROPHILIDAE PLATYPSYLLIDAE 2 I 9 




matiferous. In Berosus there are patches of aeriferous, minute 
pubescence on the body. The pupae of Hydrophilides repose on 
the dorsal surface, which is protected by 
spinous processes on the pronotum, and 
on the sides of the abdomen. 

We have already remarked that this 
is one of the most neglected of the 
families of Coleoptera, and its classifica- 
tion is not satisfactory. It is usually 
divided into Hydrophilides and Sphaer- 
idiides. The Sphaeridiid.es are in large 
part terrestrial, but their separation from 
the purely aquatic Hydrophilides cannot 
be maintained on any grounds yet pointed 
out. Altogether about 1000 species of 
Hydrophilidae are known, but this pro- 
bably is not a tenth part of those exist- FIG. 101. -Larva of Spercheus 
T T> -j. i i emarginatus. (After 

ing. In Britain we have nearly ninety Schiodte). 

species. Some taxononiists treat the 

family as a series with the name Palpicornia. The series Phil- 
hydrida of older authors included these Insects and the Parnidae 
and Heteroceridae. 

Fam. 13. Platypsyllidae.--This consists of a single species. 
It will be readily recognised from Fig. 102, attention being given 
to the peculiar antennae, and to the fact that the mentum is tri- 
lobed behind. This curious species has been found only on the 
beaver. It was first found by Ritsema on American beavers 
(Castor canadensis) in the Zoological Gardens at Amsterdam, but 
it has since been found on wild beavers in the Rhone in France ; 
in America it appears to be commonly distributed on these 
animals from Alaska to Texas. It is very remarkable that a 

t/ 

wingless parasite of this kind should be found in both hemi- 
spheres. The Insect was considered by Westwood to be a separate 
Order called Achreioptera, but there can be no doubt that it is a 
beetle. It is also admitted that it shows some points of resem- 
blance with Mallophaga, the habits of which are similar. Its 
Coleopterous nature is confirmed by the larva, which has been 
described by both Horn and Eiley. 1 Little is known as to the 
food and life-history. Horn states that the eggs are placed on 

1 Horn, Tr. Amer. cut. Soc. xv. 1888, p. 23 ; Riley, Insect Life, i. 1S89, p. 300. 



22O 



COLEOPTERA 



CHAP. 



the skin of the beaver amongst the densest hair ; the larvae move 
with a sinuous motion, like those of Staphylinidae. It has been 




B 



FIG. 102. Platypsyllus castoris. A, Upper side ; B, lower side, with legs of one side 
removed ; C, antenna. (After Westwood.) 

suggested that the Insect feeds on an Acarid, Schizocarpus 
mintjandi ; others have supposed that it eats scales of epithelium 

or hairs of the beaver. 

Fam. 14. Leptinidae. - - Antennae 
rather long, eleven-jointed, without club, bet 
a little thicker at the extremity. Eyes absent 
or imperfect. Tarsi five- jointed. Elytra 
quite, covering abdomen. Mentum with the 
posterior angles spinoiisly prolonged. A 
family of only two genera and two species. 
Their natural history is obscure, but is 
apparently of an anomalous nature ; the 
inference that may be drawn from the 
little that is known being that they are 
parasitic on mammals. There is little or 
nothing in their structure to indicate this, 

except the condition of blindness ; and 
FIG. 103. Lent inns testaceus. T i -,c i 

Britain until recently the Insects were classified 

amongst Silphidae. Leptinus testaceus 

(Fig. 103) is a British Insect, and besides occurring in Europe 
is well known in North America. In Europe it has been found 




v POLYMORPHA LEPTINIDAE SILPHIDAE 221 

in curious places, including the nests of mice and bumble-bees. 
In America it has been found on the mice themselves by Dr. 
Ryder, and by Riley in the nests of a common field-mouse, 
together with its larva, which, however, has not been described. 
The allied genus Leptinillus is said by Riley to live on the 
beaver, in company with Platypsyllus? It has been suggested 
that the natural home of the Leptinus is the bee's nest, and 
that perhaps the beetle merely makes use of the mouse as a 
means of getting from one nest of a bumble-bee to another. 

Fam. 15. Silphidae. The mentum is usually a transverse 
plate, having in front a membranous hypoglottis, which bears the 
exposed labial palpi, and immediately behind them the so-called 
bilobed ligula. The anterior coxae are conical and contiguous : 
prothoracic epimera and episterna not distinct. Visible abdomi- 
nal segments usually Jive, but sometimes only four, or as many as 
seven. Tarsi frequently Jive-jointed, but often with one joint less. 
Elytra usually covering the body and free at the tips, but occasion- 
ally shorter than the body, and even truncate behind so as to expose 
from one to four of the dorsal plates; but there are at least three 
dorsal plates in a membranous condition at the base of the abdomen. 
These beetles are extremely diverse in size and form, some being 
very minute, others upwards of an inch long, and there is also 
considerable range of structure. In, this family are included 
the burying-beetles (Necrophorus), so well known from their habit 
of making excavations under the corpses of small Vertebrates, 
so as to bury them. Besides these and Silpha, the roving 
carrion - beetles, the family includes many other very different 
forms, amongst them being the larger part of the cave-beetles 
of Europe and North America. These belong mostly to the 
genera Bathyscia in Europe, and Adelops in North America ; but 
of late years quite a crowd of these eyeless cave-beetles of the 
group Leptoderini have been discovered, so that the European 
catalogue now includes about 20 genera and 150 species. The 
species of the genus Catopomorphus are found in the nests of 
ants of the genus Aphaenogaster in the Mediterranean region. 
Scarcely anything is known as to the lives of either the cave- 
Silphidae or the myrmecophilous forms. 

The larvae of several of the larger forms of Silphidae are well 
known, but very little has been ascertained as to the smaller forms. 

1 Insect Life, i. 1889, pp. 200 and 306. 



222 



COLEOPTERA 



CHAP. 



Those of the burying-beetles have spiny plates on the back of the 
body, and do not resemble the other known forms of the family. 
The rule is that the three thoracic segments are well developed, 
and that ten abdominal segments are also distinct ; the ninth 
abdominal segment bears a pair of cerci, which are sometimes 
elongate. Often the dorsal plates are harder and better developed 
than is usual in Coleopterous larvae. This is especially the case 

with some that are en- 
dowed with great powers 
of locomotion, such as 8. 
obscura (Fig. 104). The 
food of the larvae is as a 
rule decomposing animal 
or vegetable matter, but 
some are predaceous, and 
attack living objects. 
The larger Silplia larvae 
live, like the Necro- 
plmrus, on decomposing 
animal matter, but run 

obscuni. Europe, ''bout to Seek it; hence 

lacrymosa, ma ny Specimens of SOme 
J 

ot these large larvae may 
sometimes be found amongst the bones of a very small dead bird. 
We have found the larva and imago of 8. tJwracica in birds' nests 
containing dead nestlings. 8. atrata and 8. laevigata make war 
on snails. S. lapponica enters the houses in Lapland and ravages 
the stores of animal provisions. 8. opaca departs in a very 
decided manner from the habits of its congeners, as it attacks 
beetroot and other similar crops in the growing state ; it is 
sometimes the cause of serious loss to the growers of beet. The 
larvae of the group Anisotomides are believed to be chiefly 
subterranean in habits ; that of A. cinnamomea feeds on the 
truffle, and the beetle is known as the truffle-beetle. 

The number of species of Silphidae known must be at present 
nearer 900 than 800. Of these an unusually large proportion lie- 
long to the European and North American regions ; Silphidae being 
apparently far from numerous in the tropics. Rather more than 
100 species are natives of Britain. The family reappears incon- 
siderable force in New Zealand, and is probably well represented 




FIG. 104. A, Larva of 
(After Schiodte). B, 
Australia. 



v SCYDMAENIDAE GNOSTIDAE PSELAPHIDAE 223 

in South Australia and Tasmania. The most remarkable form 
known is perhaps the Australian genus Ptomaphila (Fig. 104, B). 
The classification of the family is due to J)r. Horn. 1 The 
only change of importance that has since been suggested is 
the removal of Sphaerites from this family to Synteliidae. 
Anisotomidae and Clambidae have been considered distinct 
families, but are now included in Silphidae. 

Fam. 16. Scydmaenidae. Minute Insects allied to Silphidae, 
I nit with the hind coxae separated, and the facets of the eyes coarser ; 
the tarsi are five-jointed ; the number of visible abdominal segments is 
six. These small beetles are widely spread over the earth's surface, 
and about 700 species are now known, of which we have about a 
score in Britain ; many live in ants' nests, but probably usually 
rather as intruders than as guests that have friendly relations 
with their hosts. Nothing is known as to their life-histories, but 
the food of the imago, so far as is known, consists of Acari. Mastigus 
is a very aberrant form, found in moss and dead leaves in Southern 
Europe. By means of Brathinus the family is brought very near 
to Silphidae ; Casey, however, considers Brathinus to belong to 
Staphylinidae rather than to Scydmaenidae. The South European 
Leptomastax is remarkable on account of the slender, long, sickle- 
shaped mandibles. The Oriental genus Clidicus is the largest and 
most remarkable form of the family ; it has a very slender neck 
to its broad head, and is more than a quarter of an inch long. 

Fam. 17. Gnostidae. - - Minute Insects with three-jointed 
antennae,, five-jointed tarsi, and three apparent ventral segments, the 
first of which, however, is elongate, and consists of three united 
plates. Elytra entirely covering the after-body. The family con- 
sists of two species which have been found in the nests of ants, 
of the genus Cremastogaster, in Brazil. 2 

Fam. 18. Pselaphidae. Very small Insects ; the elytra much 
abbreviated, usually leaving as much as half the abdomen uncovered ; 
the maxillary palpi usually greatly developed, and of a variety of 
remarkable forms; the segments of the abdomen not more than 
seven in number, with little or no power of movement. T<t rxi ////// 
not more than three joints. These small Coleoptera mostly live in 
the nests of ants, and present a great diversity of extraordinary 

1 Tr. Amer. ent. Soc. viii. 1880, pp. 219-321. 

! Westwood, Tr. cni. Soc. London (X.S.) iii. 1855, p. 90; Wasmaim, Krit. 
Verzeichniss Myrmekoph. Arthropod. 1894, p. 121. 



224 COLEOPTERA CHAP. 

shapes, and very peculiar structures of the antennae and maxillary 
palpi. Owing to the consolidation of some of its segments, the 
abdomen frequently appears to have less than the usual number. 
In the curious sub-family Clavigerides, the antennae may have 
the joints reduced to two or even, to all appearance, to one ; the 
tarsi suffer a similar reduction. There are about 2500 species of 
Pselaphidae known ; many of them have never been found outside 
the ants' nests ; very little, however, is known as to their natural 
history. It is certain that some of them excrete, from little 
tufts of peculiar pubescence, a substance that the ants are fond 
of. The secretory patches are found 011 very different parts of 
the body and appendages. Claviger testaceus is fed by the ants 
in the same way as these social Insects feed one another ; the 
Claviger has also been seen to eat the larvae of the ants. They 
ride about on the backs of the ants when so inclined. The 
family is allied to Staphylinidae, but is easily distinguished by 
the rigid abdomen. Only one larva that of Chennium bituber- 
c'ulatum is known. It appears to be very similar to the larvae 
of Staphylinidae. The best account of classification and structure 
is that given by M. Achille Eaffray, 1 who has himself discovered 
and described a large part of the known species. 

Fam. 19. Staphylinidae. Elytra very short, leaving always 
some of the abdominal segments exposed, and covering usually only 
two of the segments. Abdomen usually elongate, with ten dorsal, an ft 
seven or eight ventral segments ; of the latter six or seven are usual/// 
exposed; the dorsal plates as hard as the ventral, except sometime 
in the case of the first two segments ; the segments very mobile, so 
lit at the abdomen can be curled upwards. The number of tar sal 
joints very variable, often Jive, but frequently as few as three, and 
not ahoays the same on all the feet. Staphylinidae (formerly called 
Brachelytra or Microptera) is one of the most extensive of even 
the great families of Coleoptera ; notwithstanding their diversity, 
they may in nearly all cases be recognised by the more than 
usually mobile and uncovered abdomen, combined with the fact 
that the parts of the mouth are of the kind we have mentioned 
in Silphidae. The present state of the classification of this 
family has been recently discussed by Ganglbauer. 2 At present 

1 Rev. cnt. franc, ix. 1890. 

Die Kdfer von Mitteleuropa : II. Familicnreihc, Staphylinoideu. Vienna, 1895 
and 1899. 



POLYMORPHA STAPHYLIXIDAE 



225 



a limit 9000 species are known, some of which are minute, while 
scarcely any attain a size of more than an inch in length, our 
cm union British black cock-tail, or "devil's coach-horse beetle," 
(_)i'!/l>nx oh us, being amongst the largest. Though the elytra 
are short, the wings in many forms are as large as those of 
the majority of beetles; indeed many Staphylinidae are more 
apt at taking flight than is usual with Coleoptera ; the wings 
when not in use are packed away under the short elytra, 
being transversely folded, and otherwise crumpled, in a com- 
plicated but orderly manner. It is thought that the power 
of curling up the abdomen is 
connected with the packing 
away of the wings after flight ; 
but this is not the case : for 
though the Insect sometimes 
experiences a difficulty in fold- 
ing the wings under the elytra 
after they have been expanded, 
yet it overcomes this difficulty 
by slight movements of the base 
of the abdomen, rather than 
by touching the wings with 
the tip. What the value of 
this exceptional condition of 
short elytra and corneous dorsal 
abdominal segments to the , 

FIG. 105. Staphylinidae. A, Larva of Pln- 
Insect may be IS at present lonthusnitidus. Britain. (After Schiddte.) 

I uite mysterious. The habits B - ^ M f o! ? ns > Brit / lil1 > , C ' ^ f aljd - 

1 . men, of 0. olens \vith stink-vessels. 

of the members of the family 

are very varied ; many run with great activity ; the food is 
very often small Insects, living or dead ; a great many are 
found in fungi of various kinds, and perhaps eat them. It is 
in this family that we meet with some of the most remarkable 
cases of symbiosis, i.e. lives of two kinds of creatures mutually 
accommodated with good will. The relations between the 
Staphylinidae of the genera Atemeles and Lomechusa, and certain 
ants, in the habitations of which they dwell, are very interesting. 
The beetles are never found out of the ants' nests, or at any rate 
not very far from them. The most friendly relations exist between 
them and the ants : they have patches of yellow hairs, and these 
VOL. vr Q 




226 COLEOPTERA 



CHAP. 



apparently secrete some substance with a flavour agreeable to the 
ants, which lick the beetles from time to time. On the other 
hand, the ants feed the beetles ; this they do by regurgitating 
food, at the request of the beetle, on to their lower lip, from which 
it is then taken by the beetle (Fig. 82). The beetles in many 
of their movements exactly resemble the ants, and their mode 
of requesting food, by stroking the ants in certain ways, is quite 
ant-like. So reciprocal is the friendship that if an ant is in 
want of food, the Lomecliusa will in its turn disgorge for the 
benefit of its host. The young of the beetles are reared in the 
nests by the ants, who attend to them as carefully as they do to 
their own young. The beetles have a great fondness for the 
ants, and prefer to sit amongst a crowd thereof; they are 
fond of the ants' larvae as food, and indeed eat them to a 
very large extent, even when their own young are receiving 
food from the ants. The larva of Lonn'di usa, as described by 
Wasmann (to whom we are indebted for most of our knowledge 
of this subject), 1 when not fully grown, is very similar to the 
larvae of the ants ; although it possesses legs it scarcely uses 
them : its development takes place with extraordinary rapidity, 
two days, at most, being occupied in the egg, and the larva 
completing its growth in fourteen days. Wasmann seems to be 
of opinion that the ants scarcely distinguish between the beetle- 
larvae and their own young ; one unfortunate result for the beetle 
follows from this, viz. that in the pupal state the treatment that 
is suitable for the ant -larvae does not agree with the beetle- 
larvae : the ants are in the habit of digging up their own kind 
and lifting them out and cleaning them during their meta- 
morphosis ; they also clo this with the beetle-larvae, with fatal 
results ; so that only those that have the good fortune to be 
forgotten by the ants complete their development. Thus from 
thirty Lomechusa larvae Wasmann obtained a single imago, and 
from fifty Ate //teles larvae not even one. 

Many other Staphylinidae are exclusively attached to ants' 
nests, but most of them are either robbers, at warfare with the 
ants as is the case with many species of Myrmedonia that lurk 
about the outskirts of the nests or are merely tolerated by the 
ants, not receiving any direct support from them. The most 

1 J'crylcichende Studien /r Amfiscngiistc, Xijhoff, 1890; and Tijdschr. cut. 
xxxiii. 1890, pp. 93, etc. ; T,wl. Ccntralbl. xv. 1895, p. 632. 



v STAPHYLTNIDAE SPHAERIIDAE TRICHOPTL-.RYGIDAE 22/ 



the 

The 



remarkable Staphylinidae yet discovered are some viviparous 
species, forming the genera Corotoca and Spiraclitlia, that h;i\i- 
very swollen abdomens, and live in the nests of Termites in 
Brazil: 1 very little is, however, known about them. A very 
l;irge and powerful Staphylinid, Velleius dilatatus, lives only in 
the nests of hornets and wasps. It has been supposed to be a 
defender of the Hymenoptera, but the recent observations of 
Janet and Wasmann make it clear that this is not the case : the 
Velleius has the power of making itself disagreeable to 
hornets by some odour, and they do not seriously attack it. 
Velleius finds its nutriment in larvae or 
pupae of the wasps that have fallen from 
their cells, or in other organic refuse. 

The larvae of Staphylinidae are very 
similar to those of Carabidae, but their 
legs are less perfect, and are terminated 
only by a single claw ; there is no dis- 
tinct labruni. The pupae of some are 
obtected, i.e. covered by a secondary exu- 
dation that glues all the appendages 
together, and forms a. hard coat, as in 
Lepidoptera. We have about 800 species 
of Staphylinidae in Britain, and it is prob- 
able that the family will prove one of the 
most extensive of the Order. It is prob- 
able that one hundred thousand species 
or even more are at present in existence. 

Fam. 20. Sphaeriidae. - - Very 
minute. Antennae eleven-jointed, clubbed. 
Tarsi three-jointed. Abdomen icith only 
three visible ventral segments. This family 
includes only three or four species of In- 
sects about 

very convex, and may be found walking 
on mud. 8. acaroides occurs in our fens. 
Mr. Matthews considers that they are 
most nearly allied to Hydrophilidae. 2 

Fam. 21. Trichopterygidae. Extremal;/ minute: aiit<:/in<i<- 




of an inch long. They are FlG - 106. TYicAopferys /*- 

cularis. Britain. A, Gut- 
line of perfect Insect ; B, 
part of upper surface ; C, 
larva from side ; D, from 
above ; E, pupa ; F, wiug ; 
G, natural size of imago. 



1 Schiodte, Ann. Sci. Nat. Zool. (4) v. 1857, p. 169. 
- Mol. Centr. Amer. Col. ii. pt. i. 1888, p. 156. 



228 



COLEOPTERA 



CHAP. 



clavicorn (bfisid and <q>icl joints thicker than middle joints} ; tarsi 
three-jointed; elytra so'inctinirs covering abdomen, in other cases 
leaving a variable number of segments exposed; 'icings fringed. 
Tliis family comprises the smallest Insects ; Nanosella fungi being 
only y-J-Q- of an inch long, while the largest Trj.choptej.ygid is 
only yV of an inch. The small size is not accompanied by any 
degeneration of structure, the minute, almost invisible forms, having 
as much anatomical complexity as the largest Insects. Very little 
is known as to the natural history. Probably these Insects exist in 
all parts of the world, for we have about eighty species in England, 
and Trichopterygidae are apparently numerous in the tropics. 1 

Fam. 22. Hydroscaphidae. Extremely minute aquatic Insects, 
t'-i/h chingnte abdomen. Antennae eight -jointed. The other 
characters are much the same as those we have mentioned for 
Trichopterygidae. The family is not likely to come before the 
student, as only three or four species from Southern Europe and 
North America are known." 

Fam. 23. Corylophidae. Minute beetles. Tarsi fonr-jn', ni> <1 . 
Imt tijtj>i'ii,-i ,i<i on/// three -jointed, oii'i/nj to the hind joint bring 
nnirrti/rd //// tlie < inn i'i/i n<itc (or notched^) second j<>i/if. tiix free 
ventral segments. Maxillae v.'ith only one lobe. Antfiiiuir of 

peculiar form. There are about 
200 species of these little Insects, 
but the family is apparently repre- 
sented all over the world, and will 
probably prove to be much more 
extensive. The peculiar larva of 
Ortlioperus brunnipes was found 
abundantly by Perris in thatch in 
France. Mr. Matthews proposes 
to separate the genus Aphanoceph- 
alus as a distinct family, Pseudocory- 
FIG. 107. A, Larva of Orthoperus lophidae. 3 In Corylophidae the 

brim ill ties (after Perris); B, 0. <'/"- r. ,-, 

marius, perfect Insect. Britain. wm g s are fringed With long hairs, 

as is the case in so many small 

Insects: the species of Aphanoceplialus are rather larger Insects, 
and the wings are not fringed ; the tarsi are only three-jointed. 

Monograph, Trichopterygia illustrata, by A. Matthews, London, 1872. 
2 For further information refer to Matthews, An Essay on Hydroscapha, London, 




B 



\ 



1S76, 20 pp. 1 pi. 



Hist. (5) xix. 1887, p. 115. 



POLYMORPRA SCAPHIDIIDAE SYNTELIIDAE 



229 



Fam. 24. Scaphidiidae. Front coxae small, conical; prothorax 

very closely applied to the after-body ; bind coxae transverse, witlcl;/ 
separated : abdomen with six or seven visible ventral plates ; 
antennae at the extremity with about Jive joints that become 
<l fa dually broader. Tarsi Jive-jointed. This family consists of a 
few beetles mat live in fungi, and run with extreme rapidity ; 
they are all small, and usually rare in collections. Some of the 
exotic forms are remarkable for the ex- 
treme tenuity and fragility of the long 
antennae, which bear fine hairs. The 
number of described species does not at 
present reach 200, but the family is very 
widely distributed. We have three or 
four species in Britain. All we know of 
the larvae is a description of 'that of 
Scaphisoma agaricinum by Ferris; 1 it is 




like the larva of Staphylinidae, there are 
nine abdominal segments in addition to a 



FIG. 108. Scaphisoma agari- 
cinum. Britain. A 
Larva (after Perris) ; B 
perfect Insect. 



very short, broad pseudopod, and very short 

cerci. This larva feeds on agarics ; 'it goes through its development 

in about three weeks ; unlike the adult it is not very active. 
Fam. 25. Synteliidae. Antennae davicorn, with very large 

club : labium, with hypoglottis and the parts beyond it, exposed. 

Front coxae transverse. Abdmni'ii with Jive visible ventral segments, 

and eight or nine dorsal, the b/ixidfour of which are semi-corneous. 

This family includes only five species; its clas- 
sification has given rise to much difference 
of opinion. We have, after consideration 
of all its characters, established it as a 
distinct family 2 allied to Silphidae. The 
perfect Insects live on the sap running from 
trees : but nothing else is known of their 
natural history. Like so many others of 
the very small families of aberrant Cole- 
optera, it has a very wide distribution ; 

PIG. \Q9.-Syntelia west- fyntelia being found in Eastern Asia and 
modi. Mexico. (From Mexico, while the sub-family Sphaeritides 
occurs, as a single species, in Europe and 




f 



Biol. Cent/: Amer.) 



North America. The earlier instars are unknown. 
1 Larres dc Coldopteres, 1878, p. 11. pi. i. - Biol. Centr. Amer. Col. ii. pt. i. p. 438. 



230 



COLEOPTERA 



CHAP. 



Fam. 26. Histeridae. Very compact beetles, with very hard 
integument, short, bent /<// //'<v, with a very compact club: no 
hypofjlottis. Elytra closely applied to body, but straight behind, 
leaving fc'<> xe/juients exposed. Abdomen with Ji re visible ventral 
segments ; vo'/A. seven <l<>rxl */>/i//ents, all liard. Front ce.'-<- 
strongly transverse, hind coxae widely separated. The extremely 
compact form, and hard integument, combined with the peculiar 
antennae consisting of a long basal joint, six or seven small 
joints, and then a very solid club of three joints covered with 

minute pubescence render 
these Insects unmistakable. The 
colour is usually shining black, 
but there are numerous depar- 
tures from this. The way in 
which these Insects are put 
together so as to leave no chink 
in their hard exterior armour 
when in repose, is very remark- 



able. The mouth - parts are 
rather highly developed, and 
the family is entitled to a high 
rank ; it consists at present of 
about 2000 species ; l in Britain 
we have about 40. The larvae 
are without ocelli or labrum, 
FIG. UQ.Platysoma depression. Europe, but have well-developed mancli- 

A, Larva (after Schiodte) ; B, perfect , , i T 

insect. oles, the second and third 

thoracic segments being short, 

the ninth segment of the abdomen terminal, with two distinctly 
jointed cerci. 2 Histeridae are common in dung, in carcases, decay- 
ing fungi, etc., and some live under bark these being, in the 
case of the genus Hololepta, very flat. Some are small cylinders, 
elaborately constructed, for entering the burrows of 'Insects in 
wood (Trypanaeus) ; a certain number are peculiar to ants' nests. 
Formerly it was supposed that the Insects were nourished on 
the decaying substances, but it is now believed, with good reason, 
that they are eminently predaceous, in both larval and imaginal 

The family was monographed by the Abbe de Marseul in Ann. Soc. cut. France, 
1853-1862, but great additions have been made since then. 

2 For characters of larvae of various genera, see Ferris, Larrcs, etc. p. 24. 




HISTERIDAE PHALACRIDAE NITIDULIDAE 



231 




instars, and devour the larvae of Diptera, etc. The relations of the 
ants'-nest forms to the ants is not made out, but it is highly probable 
that they eat the ants' larvae, and furnish the ants with some 
dainty relish. A few species live in company with Termites. 
Fam. 27. Phalacridae. Body very compact; elytra entirely 

covering it; apical joints of antennae rather 
broader, usually long ; front coxae globular ; 
posterior coxae contiguous ; abdomen with 
five visible ventral segments ; tarsi five- 
jointed, fourth joint usually small and 
obscure. This family consists entirely of 
small Insects : the tarsal structure is very 
aberrant, and is also diverse, so that the 
student may without careful observation 
FIG. ill. Olibrus bicolor. pass the Insects over as having only four- 

J illted toi ; theh> structure, so far _ as 
the front two pairs are concerned, being 
very nearly that of many Fhytophaga. The larvae live in the 
heads of flowers, especially of the flowers of Compositae. Having 
bored their way down the stems, they pupate in earthen cocoons. 
Heeger 1 says that he has observed in favourable seasons six 
generations ; but the larvae die readily in unfavourable seasons, 
and are destroyed in vast numbers when the meadows are 
mowed. Seven years ago very little was known as to the family, 
and the list of their species scarcely amounted to 100, but now 
probably 300 are described. They occur in all parts of the 
world ; we have fourteen in Britain. 

Fam. 28. Nitidulidae. Antennae with a three-jointed club ; 
all the coxae separated, and each with an external prolongation ; 
tarsi five-jointed, the fourth joint smaller than any of the others ; 
abdomen with Jive visible plates. These Insects are numerous, 
about 1600 species being at present known; in many of them 
the elytra nearly or qiiite cover the hind body, but in many 
others they are more or less abbreviated ; in this case the 
Insects may be distinguished from Staphylinidae by the form 
of their antennae, and the smaller number of visible ventral seg- 
ments. The habits are very varied, a great many are found on 
flowers, others are attracted by the sap of trees ; some live in 
"We have about 90 species in Britain ; several forms of 

1 SB. Ak. JFien. xxiv. 1857, p. 330. 



232 



COLEOPTERA 



CHAP. 




A 



the geneva, Meligethes and Epuracn are among the most abundant 
of our beetles. Most of what is known as to the larvae is 
due to Ferris ; several have been found living in flowers ; that 
of Pria haunts the flower of Xalaniim dulcamara at the junction 
of the stamens with the corolla; the larva of Meligethes aeneus 

sometimes occasions much loss by prevent- 
ing the formation of seed in cultivated 
Cruciferae, such as Eape. These floricolous 
larvae grow with great rapidity, and then 
leave the flowers to pupate in the ground. 
The larva of Nitidula lives in carcases, 
though it is not very different from that 
of Pria. The larva of tforuiiia. lives in 

FIG. 112. Pria dulcamarae. fermenting sap, and has four hooks curv- 
Britain. A, Larva (after i n g upwards at the extremity of the body. 

Ferris) ; B, perfect Insect. .,"' . /-/ 7 7 7 ", 

The curious genus Uybocephalus consists 

of some very small, extremely convex Insects that live in flowers 
in Southern Europe ; they have only four joints to the tarsi. 
The perfect Insects of the group Ipides are remarkable from 
having a stridulating organ on the front of the head. The classi- 
fication of the well-known genus Pihizophagus has given rise to 
much discussion ; although now usually placed in Mtidulidae, 
\vc think it undoubtedly belongs to Cucujidae. 

Fam. 29. Trogositidae. Differs from Nitidulidae in the struc- 
ture of tin' tarsi ; tliese appear f<> lefour- 
jointed, with tlie third joint similar in 
size and form to the jircccdi nij ; tliei/ 
are, however, reaUt/ five-jointed, an eX- 
fre/rtC/1/ sit oft liana/ joint lici/iif /'/'/'Sent. 
Hind COXae continuous. The chili of cacli 
antenna is hilat,-,-all// asymmetric, and 
the sensitive surface is confined to certain 
parts of the joints. There are some 400 
or 500 species of Trogositidae, but nearly 
all of them are exotic. The larvae (Fig. 
113, A), are predaceous, destroying other FIG. \\$.TemnocMa coerulea. 

Europe. A, Larva (after 

larvae in large numbers, and it is pro- p^-ris) . B, perfect insect, 
bable that the imagos do the same. 

The larva of Tcnelroides (better known as Troaoxita} -inauritanica 
is found in corn and flour, and is said to have sometimes been very 




1'oLVMORPHA TROGOSITIDAE COLYDIIDAE 



233 



injurious by eating the embryo of the corn, but it is ascertained 
that it also devours certain other larvae that live on the corn. 
This beetle has been carried about by commerce, and is now nearly 
cosmopolitan. Our three British species of Trogositidae repre- 
sent the three chief divisions of the family, viz. Xemosomides, 
Temnochilides, Peltides ; they are very dissimilar in form, the 
Peltides being oval, with retracted head. It is doubtful whether 
the members of the latter group are carnivorous in any of their 
stages ; it is more probable that they live on the fungi they 
frequent. Peltidae stand as a distinct family in many works. 1 

Fam. 30. Colydiidae. Antennae with a terminal club, tarsi 
four -jointed, none of the joints broad; front and middle coxae 
small, globose, embedded ; hind coxae transverse, either contiguous or 
separated ; Jive visible ventral segments, several of which have no 
movement. This is a family of interest, owing to the great diver- 
sity of form, to the extraordinary sculpture and clothing exhibited 
by many of its members, and to the fact that most of its members 
are attached to the primitive forests, and disappear entirely 
when these are destroyed. We have fifteen species in Britain, 
but about half of them are of the greatest rarity. There are 
about 600 species known at present; Xew Zealand has produced 

the greatest variety of forms ; the forests 
of Teneriffe are rich in the genus Tar- 
[iliius. The sedentary lives of many of 
these beetles are very remarkable ; the 
creatures concealing themselves in the 
crannies of fungus-covered wood, and 
scarcely ever leaving their retreats, so 
that it is the rarest circumstance to find 
them at any distance from their homes. 
Langelandia anophthalma, lives entirely 
underground and is quite blind, the 




FIG. 114. 



crenata. 



Britain. A, Larva (after optic lobes bei absent. Some Coly- 

erfect Insect. 



Ferris) ; B, perfect Insect. 



the 



. 

diidae are more active, and enter 

burrows of wood-boring Insects to destroy the larvae ((',> 
Few of the larvae are known ; but all appear to have the body 
terminated by peculiar hard corneous processes, as is the case 
with a great variety of Coleopterous larvae that live in wood. 2 

1 Catalogue of Trogositidae, by Leveille, in Ann. Soc. cnt. France, 1888, p. 429. 
- For classification, see Sharp, Biol. Ccntr. A mcr. Co 7 , ii. pt. i. 1894, p. 443. 



234 



COLEOPTERA 



CHAP. 



Fam. 31. Rhysodidae. Tarsi four-jointed ; mouth-parts 

covered, l>y the I a rye 'inentum ; front tibiae notched on the inner 
cd(i>\ This family consists only of a few species, but is found 
nc.-irly all over the world in the warm and temperate regions. 
In many of their characters they resemble the Adephaga, but are 
very different in appearance and in the mouth. The larvae are 
not known. Some authorities think these Insects should be 
placed in the series Aclephaga, 1 but it is more probable that -they 
will prove to be amongst the numerous aberrant forms of Coleop- 
tera that approach the various large natural series, without 
really belonging to them. The three families, Colydiidae, Cucu- 
jidae, and Rhysodidae, exhibit relations not only with other 
families of the Coleoptera Polymorpha, but also with most of the 
great series ; Adephaga, Rhynchophora, Fhytophaga, and Hetero- 
mera, being each closely approached. 

Fam. 32. Cucujidae. Tarsi Jive- or four-jointed, the first 
joint often short: anteniia.e sometimes r/nlilied, but more often 
<j_iiite tli i ii ut the tip; front cud midille coj.ae deeply embedded, 
globular, but n-ith an, an<inlar prolongation externally ; abdomen 
with Jive visible rrnfral segments, all movable. This family and 

the Cryptophagidae are amongst the 
most difficult families to define ; 

3=* 

indeed it is in this portion of the 
Clavicorns that an extended and 
thorough study is most urgently 
required. The Cucujidae include a 
great diversity of forms ; they are 
mostly found under the bark of trees, 
and many of them are very fl;ii. 
Many of the larvae are also very 

t/ t/ 

Fro. 115. Brontes plcmatus. Britain, flat, but Ferris says there is great 

A, Larva; B, pupa; C, perfect i- - f fV-pir ^rnr-tnrp thpv 

Insect. (A and B after Perils.) 

are probably chiefly carnivorous. 

There are about 400 species described ; we have nearly a score in 
Britain. 

The family Cupesidae of certain taxonomists must be at 
present associated with Cucujidae, though the first joint of the 
tarsus is elongate. 

1 See Ganglbauer, Kaf. Mitteleuropas, i. p. ">:;(). as well as Leconte and Horn 
Classification, etc., p. 130. 




\ 



POLYMORPHA CRYPTOPHAGIDAE HELOTIDAE 235 




A 



Fam. 33. Cryptophagidae. Front and middle coxae very 
small and deeply embedded; antennae with enlarged terminal joints ; 
lursi five- jointed, the posterior sometimes in the male only four - 
jointed; abdomen with five visible ventral segments, capable of 
in < > r*' at cut, the first much longer than any of the others. A small 
family composed of obscure forms of 
minute size, which apparently have 
mould-eating habits, though very little 
is known on this point, and several of 
the genera (Anther ophagus, Tclmatophi- 
lus) are found chiefly on growing plants, 
especially in flowers. Although the 
imago of Antherophagus lives in flowers, 
yet the larva has only been found in 
the nests of bumble-bees ; there is FIG. ne.Cryptophagvs denta- 
reason for believing that the imago 
makes use of the bee to transport 
it from the flowers it haunts to the nests in which it is to 
breed ; x this it does by catching hold of the bee with its 
mandibles when the bee visits the flower in which the beetle is 
concealed. It is strange the beetle should adopt such a mode of 
getting to its future home, for it has ample wings. We must 
presume that its senses and instinct permit it to recognise the 
bee, but do not suffice to enable it to find the bee's nest. Some 
of the larvae of the genus Cryptophagus are found abundantly in 
the nests of various wasps, where they are probably useful as 
scavengers, others occur in the nests of social caterpillars, and they 
are sometimes common in loose straw ; this being the habitat in 
which Penis found the one we figure. 

Fam. 34. Helotidae. Front and middle coxal cavities round, 
with scarcely any angular prolongation externally ; all the coxae 
widely separated ; five visible ventral segments, all mobile. 
The Insects of this family are closely allied to Trogositidae 
and Nitidulidae, and have the tarsal structure of the former 
family ; but the Helotidae are .^different in appearance from 
any members of either of these two families, and are readily 
distinguished by the coxal character. They are frequently 
classified with the Erotylidae, from which they differ by the 
differently shaped feet, especially by the diminished basal joint. 

1 Ferris, Larves. etc.. j>. 75. 



236 COLEOPTERA CHAP. 

There is lint one genns, and for a long time only two or three 
species were known, and were great rarities in collections : in 
the last few years the number has been raised to nearly forty. 1 
They are remarkable beetles with oblong form, and a somewhat 
metallic upper surface, which is much sculptured, and possesses 
four yellow, smooth spots on the elytra. According to Mr. George 
Lewis they are found feeding at the running sap of trees, but the 
larvae are not known. Helotidae are peculiar to the Indo-Malayan 
rr- inn (including Japan) with one species in Eastern Africa, 

Fam. 35. Thorictidae. Tarsi five-jointed, none ofthejoinf* 
hr od ; front coxae small, rather prominent, but not at oil fr /in- 
verse ; fire visible re/itn/J jila/rx, a/I mobile; mctasternn i very 
situ rt ; antennae xln>ff,irith a solid dub. This little family, con- 
sisting of the genus Thorirfnx, appears to be a distinct one, 
though the structure has only been very imperfectly studied. It 
is peculiar to the Mediterranean region, where the species live in 
ants' nests. They appear to be on terms of great intimacy with 
the ants ; a favourite position of the beetle is on the scape of the 
antenna of an ant ; here it hooks itself on firmly, and is carried 
about by the ant. Like so many other ants'-nest beetles, 
Thorictidae possess tufts of golden hair, which secrete some 
substance, the flavour of which is appreciated by the ants; these 
tufts in Thorictidae are situated either at the hind angles of the 
pronotmn, or on the under surface of the body on each side of the 
breast ; AVasmaim thinks that when the beetles are riding about, 

as above described, the ants have tln-n 
an opportunity of getting at the patches 
on the under surface. 

Fam. 36. Erotylidae.--ram five- 
jointed, but 'iritJi the fourth -usual/ // very 
small, the first three mure or less brood 
a/id i>id>e*ee/// beneath. Antennae xfro//>//f/ 
clubbed. Front and middle coxal aeetu- 
"\T/ ) bula round, without angulcw^prolongation 

FIG. in.-Tritoma Mpustulata. externally; fire risible ventral segments. 

Erotyiiiiae. Britain. A, Larva This is now a large and important family 
(after Ferris) ; B, perfect Insect. . ... i u 

of about 1800 species, but it is chiefly 
exotic and tropical, its members haunting the fungoid growths 

1 Ritsema, Catalogue of Helota, Xotcs Lcijdcn J/HS. xiii. 1891, p. 223, and xv. 
1S93, p. UK). 




EROTYLIDAE--MYCETOPHAGIDAE - COCCINELLIDAE 237 

in forests. "We have only six species in Britain, and the whole 
of Europe has only about two dozen, most of them insignificant 
(and in the case of the Dacnides aberrant, approaching the 
Cryptophagidae very closely). The sub-family Languriicles (quite 
wanting in Europe) consists of more elongate Insects, with front 
acetabula open behind ; they have different habits from Eroty- 
lides proper ; some are known to live as larvae in the stems of 
herbaceous plants. They possess a highly developed stridulating 
organ on the front of the head. The Clavicorn Polymorpha are 
very closely connected with the Phytophaga by Languriicles. 

Fam. 37. Mycetophagidae. Tarsi four-joint n], slender, the 
front fed of the male only three-jointed ; coxae oml, not deeply 
embedded; abdomen icif/i jin- rentral seg- 
ments, all movable. A small family, of in- 
terest chiefly because of the anomaly in the 
feet of the two sexes, for which it is im- 
possible to assign any reason. The species 
are small, uninteresting Insects that live 
chiefly on Cryptogams of various kinds, 
especially in connection with timber ; the 

larvae being also found there. There are - 

FIG. Jl8. Litargus bifas- 

about a dozen species in Britain, and ciatu*. Mycetophagidae. 




scarcely 100 are described from all the 
world. The DiphyUid.es, placed by Leconte 
and Horn in this family, seem to go better in Cryptophagidae. 

Fam. 38. Coccinellidae (Lady -birds}. Tarsi apparent!;/ three- 
jointed ; the jirst two joints pubescent ln/nntli; the third joint 
consisting really of tivo joints, the small true third joint being 
inserted near the base of the second joint, the upper surfun- of which 
is grooved to receive it. Head much concealed l>i/ the thorax. 
Antennae feebly clubbed. The lady-birds number fully 2000 
species. The structure of their feet distinguishes them from 
nearly all other Coleoptera- except Endomychidae, which are much 
less rotund in form, and have larger antennae. One genus of 
Endomychids Panomoea bears, however, a singular resemblance 
to lady-birds, both in form and style of coloration. Several 
species of Coccinellidae are remarkable on account of the 
numerous variations in colour they present. Coccinellidae 
frequently multiply to an enormous extent, and are of great 
value, as they destroy wholesale the plant -lice, scale - Insects, 



238 COLEOPTERA 



CHAP. 



and Acari thar are so injurious to cultivated plants. They 
also eat various other soft-bodied Insects that attack plants. As 
thev are excessivelv voracious, and are themselves sin<nilarlv 

* t, t 

free from enemies and multiply with great rapidity, all these 
features of their economy render them of inestimable value to the 
agriculturist and horticulturist. The species of the sub-faniily 
Epilachnides feed on plants, and one or two are occasionally in- 
jurious. The body-fluid of Coccinellidae has an unpleasant odour 
and taste. Many lady-birds have the power of exuding, when 
disturbed, small quantities of a yellow fluid. Lutz has shown that 
this is not a special secretion, but an exudation of the fluid of 
the body that takes place through a small orifice at the tip of 
the tibia, from pressure caused by contraction of the body and limb. 1 
The larvae are much more active than beetle-larvae usually 
are, and many of them are very conspicuous when running about 
on plants to hunt their prey. They usually cast their skins 
three times, and sometimes concomitantlv change a good deal in 



colour and form : the larval life does not usuallv exceed four or 



five weeks : at the end of which time the larva suspends itself 
by the posterior extremitv. which is slued bv a secretion to some 

* -L . * v 

object ; the larval skin is pushed back to the anal extremity, 
disclosing the pupa : this differs in several respects from the 
usual pupa of beetles : it is harder, and is coloured, frequently 
conspicuously spotted, and dehisces to allow the escape of the 
beetle, so that the metamorphosis is altogether more like that of 
Lepidoptera than that of Coleoptera. There is much variety in 
the larvae ; some of them bear large, complexly-spined, projections : 
those of the group Scymnites have small depressions on the 
surface, from which it has been ascertained that waxy secre- 
tions exude : but in Scymnus minimus no such excretions are 
formed. Certain species, when pupating, do not shuffle the skin to 
the extremitv of the bodv, but retain it as a covering for the 

e. t, ' O 

pupa. The larvae that feed on plants are much less active than 
the predaceous forms. "\Ve are well supplied with Coccinellidae 
in Britain, forty species being known here. 

The systematic position of Coccinellidae amongst the Coleoptera 
has been for long a moot point. Formerly they were associated 
with various other beetles having three-jointed, or apparently 
three-jointed, feet, as a series with the name Trimera, or 

1 Z<jol. virt:. sviii. 1895. p. 244. 



IXELLIDAE EXDOMYCHIDAE MYCETAEIDAE 2 

I'-'-udotrimera. But :..- _ -rally placed in the Clavicorn 

ar Endoniychidae, rhoeff has recently made con- 

morphological stu: the male genital organs of 

i, and as the result, he concludes that Coccinellidae 
radically from all other Coleoptera as regards these 
- _ ictures, and he therefore treats them as a distinct series or 
sub-order, termed Siphonophora. The genus Lithophilus has been 
red doubtfully a member of Coccinellidae, as the tarsi 
-sess only in a slight degree the shape characteristic of the 
.ilv : Verhoeff finds that they are truly Coccinellidae, forming 
a distinct division, Lithophilini: and our little species of Coccidula, 
which have somewhat the same appearance as Lithophilini, he 
its as another separate group, Coceidulini. 
Fam. 39. Kndomychidae. 1 - 

' tico joints broad, the terminal joint elongate ; at the base of 

terminal joint there is, however, a, very small joint, so that the 

<i are pseudotetramerous ; antennae rather large, with a large 

' ; labium not at all retracted behind the mentum; front and 

hlle coxae globose ; abdomen with five movable ventral segments, 

and, a sixth more or less visible at the tip. This family includes 

a considerable diversity of elegant Insects that frequent fungoid 

growths on wood. It comprises at present fully 500 species, but 

nearly the whole of them are exotic, and inhabit the tropical 

forests, "\Ve have only two British species, both of which are 

now rarities, but apparently were much commoner at the 

beginning of the century. The larvae are 

broader than is usual in Coleoptera : very 

few, however, are known. 

Fam. 40. Mycetaeidae. T 

'. the first two joints not very different 
third, usually slender; abdomen 
n-ve visible ventral segments, which are 
~le; front and middle coxae globular. A B 

The little Insects composing this familv are FIG. ii9.--J/y*ai 

.Britain. A. La: 

many placed as a division of Endomy- ^after Bii^ r. : B ? per- 
chidae, and Verhoeff is of opinion that the 
_ up is an altogether artificial one ;. but we think, with Duval, 
it makes matters simpler to separate them. There are only 

1 G- : - :. Monographie dcr E>id*>myehiden. Leipzig. -"- ". | .Since this 

work was published, the species known have been multiplied two or three times. 




24-O 



COLEOPTERA 



CHAP. 



some forty or fifty species, found chiefly in Europe and North 
America. We have three in Britain ; one of these, Myeetaea 
hi /-fa is very common, and may be found in abundance in cellars 
in the heart of London, as well as elsewhere ; it is said to have 
injured the corks of wine-bottles, and to have caused leakage 
of the wine, but we think that it perhaps only increases some 
previous deficiency in the corkage, for its natural food is fungoid 
matters. The larva is remarkable on account of the clubbed 
hairs at the sides of the body. 

Fam. 41. Latridiidae. Tarsi three-jointed; anterior 
cavities round, not prolonged externally; alxlomen with j! rr 
innl mobile ventral *</j inputs. Very small Insects, species of which 
are numerous in most parts of the world, the individuals being 
sometimes very abundant. The larvae (Fig. 120, A) are said by 
IVrris to have the mandibles replaced by fleshy appendages. The 
pupa of L*it r'nl i us is remarkable, on account of the numerous long 
hairs with heads instead of points; the larva of Cni-firm-ia is 
very like that of Lai rid ins, but some of the hairs are replaced by 
obconical projections. The suit-family Monotomides is by many 
treated as a distinct family ; they have the elytra truncate behind, 
exposing the pygidium, and the coxae are very small and very 
deeply embedded. Most of the Latridiidae are believed to live on 
fungoid matters ; species of Monotoma 
live in ants' nests, but probably have 
no relations with the ants. A few 
species of Latridiides proper also main- 
tain a similar life; Coluocera formicaria 
is said to be fond of the stores laid up 
by ApJiaenogaster structor in its nests. 
About 700 species are now known; 
scarcely any of the individuals are more 

than one-tenth of an inch long. We FIG. 120. Lutrui.ius mi nut n^. 
have about 40 species in Britain. The 
North American genus Stephostethus has 
the prosternum constructed behind the coxae, somewhat in the 
same manner as it is in the Ehynchophorous series of Colenptem. 

Fam. 42. Adimeridae. Tarsi apjix/ri/iir <>/i/// tico-jointed, 
I basal joint and an chif/afe claw-bearing joint, but between 




tin* two tin-re are two very small joints. This family consists 
only of the American genus Adi/ncrus; nothing is known of 



POLYMORPHA DERMESTIDAE 



241 




the life -history of these small Insects. They are of some 
interest, as this structure of the foot 
is not found in any other beetles. 

Fam. 43. Dermestidae. Tarsi 
jive-jointed; antennae usually short, 
with the club frequently very large 
in proportion, and with the. under 
side of the thorax hearing a. hollow 
for its reception. Front coxae rather 
long, oblique: hind coxa formed to 
receive the femur when in repose. 
A family of 300 or 400 species of 
small or moderate-sized beetles ; the FlG - 121. Adimerus setosvs. Adi- 

,, 1-1 meridae. A, the Insect ; B, one 

SUrlace, usually Covered With hlie foot more enlarged. Mexico. From 

hair, forming a pattern, or with BioL Centr - Amer - CoL ij - P*- * 
scales. By turns, the position of which has long been disputed, 
has now been placed in this family ; it has a more imperfectly 
formed prosternum, and the third and fourth joints of the tarsi 
are prolonged as membranous lobes beneath ; the hind coxae leave 
the femora quite free. Dermestidae in the larval state nearly all 
live on dried animal matter, and are sometimes very destructive ; 
some of them totally destroy zoological collections. They are 

very remarkable on account of the complex 
clothing of hairs they bear ; they have good 
powers of locomotion, and many of them 
have a peculiar gait, running for a short 
distance, then stopping and vibrating some 
of their hairs with extreme rapidity. They 
exhibit great variety of form. Many of 
them are capable of supporting life for long 
periods on little or no food, and in such 
cases moult an increased number of times : 
pupation takes place in the larval skin. 
Anthrenus fasciatus has been reared in large 
numbers on a diet of dried horse-hair in 
furniture. The young larva of this species 
observed by the writer did not possess 
the remarkable, complex arrangement of hairs that appeared 
when it was further grown. The most curious of Dermestid 
larvae is that of Tiresias serra, which lives amongst cobwebs in 

VOL. VI R 




FIG. 122. Tiresias serra 
Larva. New Forest. 



242 COLEOPTERA CHAP. 

old wood, and probably feeds on the remains of Insects therein, 
perhaps not disdaining the cobwebs themselves. Attention has 
been frequently called to the hairs of the larvae of these Insects, 
but they have never been adequately discussed, and their function 
is quite unknown. 

Fam. 44. Byrrhidae (Pill-beetles). Oval or round, convex beetles; 
tarsi five-jointed, front coxae not exserted, transverse ; hind coxa 
shic/di/iif the retracted femur. The ivhole of the appendages 
capable of a complete apposition to the body. Although a small 
family of only 200 or 300 species, Byrrhidae are so hetero- 
geneous that no characteristic definition that will apply to 
all the sub-families can be framed. Very little is known as to 
their life -histories. Byrrhus pilula is one of our commonest 
beetles, and may be found crawling on paths in early spring 
even in towns ; it moves very slowly, and when disturbed, at 
once contracts the limbs so completely that it looks like an 
inanimate object. The larva is cylindrical, soft ; the prothoracic 
and last two abdominal segments are larger than the others, 
the last bearing two pseudopods ; its habits are unknown, and 
110 good figure exists of it. 

The chief groups of Byrrhidae are Nosodendrides, Byrrhides 
(including Amphicyrtides), Limnichides, and Chelonariides. The 
first consists of species frequenting the exuding sap of trees ; 
they have an unusually large mentuni, abruptly clubbed antennae, 
and the head cannot be retracted and concealed. The genus 
yosw/,'>it/i'i'ii serins to be distributed over a large part of the 
world. The Byrrhides have the antennae gradually thicker 
towards the tip, the mentum small, and the head and thorax so 
formed that the former can be perfectly retracted. The species 
are rather numerous, and are found in the northern and anti- 
] MX leal regions, being nearly completely absent from the tropics. 
The Limnichides are minute Insects living in very moist places : 
they have small delicate antennae, which are imperfectly clubbed. 
The group is very widely distributed. 

The Chelonariides are a very peculiar form of Coleoptera : 
oval Insects of small size with the prothorax so formed that the 
head can be withdrawn under (rather than into) it, and then 
abruptly inflexed, so that the face then forms part of the under 
surface : the antennae have the basal three joints thicker than 
the others ; these being not in the least clubbed, but having the 



POLYMORPHA 243 



joints so delicately connected that the organs are rarely un- 
mutilated. The modifications of the head and prothorax are 
quite unlike those of other Byrrhidae, and if the Chelonariides 
do not form a distinct family, they should be associated with 
Dascillidae. -Nothing is known as to the earlier stages. They 
are chiefly tropical Insects, though one species is found in North 
America. 

Fam. 45. Cyathoceridae Minute Insects of "broad form ; parts 
of the mouth concealed ; antennae four-jointed ; tarsi not divided 
into joints ; prosternum small. The only species of this aberrant 
family, Cyathocerus horni, has been found in Central America. 
Nothing is known as to its life-history. 

Fam. 46. Georyssidae. -Antennae short, dulled ; tarsi four- 
jointcd; prosternum very small; front coxae exserted, but not 
contiguous. There are about two dozen species of these small 
beetles known. Our British Georyssus pygmaeus lives in extremely 
wet places, and covers itself with a coating of mud or fine sand 
so that it can only be detected when in movement. Nothing 
further is known as to its life-history or habits. Members of 
the genus have been detected in widely- separated parts of the 
globe. 

Fam. 47. Heteroceridae. Labrum and mandibles projecting 
forwards; antennae short, the terminal seven joints broad and short, 
forming a sort of broad serrate club ; legs armed u'ith stout spines ; 
fi/rsi four-jointed. The Heteroceridae are small beetles covered 
with very dense but minute pubescence ; they live in burrows 
among mud or sand in wet places, and are found in many parts 
of the world. They possess a stridulating organ in the form 
of a slightly elevated curved line on each side of the base of the 
abdomen, rubbed by the posterior femur. The larvae live in the 
same places as the beetles ; they have well-developed thoracic- 
legs, the mandibles are porrect, the three thoracic segments 
rather large, and the body behind these becomes gradually 
narrower ; they are believed to eat the mud amongst which 
they burrow. "We have seven British species of Hetero- 
ceridae. 

Fam. 48. Parnidae. Prosternum. distinct in front of the coxae, 
iixinillil i-lniii'iate, behind forming a process receirnJ into a definite, 
cavity on the mesosternum ; head retractile, 1 1/ c ///<///// protected 
by the prosternum. Tarsi Jive -jointed, term i /ml joint long. 



244 COLEOPTERA CHAP. 

Although the characters of these Insects are not very different 
from those of Byrrhidae, of Dascillidae, and even of certain 
Elateridae, there is practically but little difficulty in distinguish- 
ing Parnidae. They are of aquatic habits, though many, in the 
perfect state, frequently desert the waters. There are about 
300 or 400 species known, but the family is doubtless more 
extensive, as these small beetles attract but little notice. There 
are two groups: 1. Parnides, in which the front coxae have a con- 
siderable transverse extension, the antennae are frequently short 
and of peculiar structure, and the body is usually clothed with 
a peculiar, dense pubescence. 2. Elmides, with round front coxae, 
a bare, or feebly pubescent body, and simple antennae. Parnus 
is a genus commonly met with in Europe, and is less aquatic in 
habits than its congeners ; it is said to enter the water carrying 
with it a coating of air attached to its pubescence. Its larvae 
are not well known ; they live in damp earth near streams, and 
are said to much resemble the larvae of Elateridae. Potamophilus 
urii niinatus has a very interesting larva, described by Dufour ; it 
lives on decaying wood in the Adour. It is remarkable from 
the ocelli being arranged so as to form an almost true eye on 
each side of the head; there are eight pairs of abdominal 
spiracles, and also a pair on the mesothorax, though there are 
none on the pro- or meta-thorax ; each of the stigmata has four 
elongate sacs between it and the main trachea! tube ; the body 
is terminated by a process from which there can be protruded 
bunches of filamentous branchiae. The larvae of Macronychus 
quadrituberculatus is somewhat similar, though the features of its 
external structure are less remarkable. The Elmides live attached 
to stones in streams ; the larva is rather broad, fringed at the 
sides of the body, and bears behind three elegant sets of fine 
filamentous branchiae. The North American genus Psephrm/x 
is placed in Parnidae, though instead of five, the male has 
seven, the female six, visible ventral segments ; the larva is 
elliptical, with dilated margins to the body. Friederich, has 
given, 1 without mentioning any names, a detailed account 
of Brazilian Parnid larvae, that may perhaps be allied to 
Psephenus. 

Fam. 49. Derodontidae. Ta/'si //Vr-/W///<v/, x/mJer, fourth 
joint rather small ; front cover prominent ami transversely pro- 

1 Stettin, cnt. Zeit. xlii. 1881, pp. 104-112. 



POLYMORPHA DERODONTIDAE CIOIDAE 



245 



lonycd ; middle coxae small; abdomen with five visible segments, 
all mobile , the first not elongated. One of the smallest and least 
known of the families of Coleoptera ; it 
consists of four or five species of small 
Insects of the genera Derodontus and 
Peltasticta, found in Xorth America, 
Europe, and Japan. The distinction of the 
family from Cleridae is by no means cer- 
tain ; our European Laricobius apparently 
possessing characters hut little different. 
Nothing is known as to the life-histories. 
Fam. 50. Cioidae. Small or minute 
beetles; antennae short, terminal joints 
thicker; tarsi short, four-jointed ; anterior 
and middle coxae small, oval, deeply em- 




c 



FIG. 123. Derodontus macu- 

bedded ; abdomen with Jim ventral sey- i atus _ North America. 

ments, all mobile. The position of these 

obscure little Insects seems to be near Colydiidae and Crypto- 

phagidae, though they are usually 
placed near Bostrichidae. So far as 
known, they all live in fungi, or in 
wood penetrated by fungoid growths. 
The cylindrical larvae live also in 
similar matter ; they usually have the 
body terminated behind by one or two 
huoks curved upwards; that of Cis 
nielliei (Fig. 124) has, instead of these 
hooks, a curious chitinous tube. About 
300 species of the family are now 
known ; a score, or so, occurring in 
Britain. The Hawaiian Islands have 
a remarkably rich and varied fauna of 

FIG. l-2-i.Cis melliei. Martin- Cioiclae. 

ique. A, Perfect Insect ; B j. 51 SpMndidae. This family 

pupa ; C, larva ; D, terminal 

portion of body of larva, of half a dozen species of rare and 

(After Coquerel.) gmdl Ingects> Differs f rom Cioidae by 

the tarsi being five-jointed at any rate on the front and middle 
feet, opinions differing as to whether the number of joints of the 
hind tarsi is four or five. These Insects live in fungi growing 
in wood, e.g. Reticularia hortensis, that are at first pulpy and 




246 



COLEOPTERA 



CHAP. 



afterwards become powder. The larvae of both of our British 
genera, Sphindus and Aspidipliorus, have been described by 
Ferris, who considers them allied to the fungivorous Silphidae 
and Latridiidae. The systematic position of these Insects has 
been the subject of doubt since the days of Latreille. 

Fam. 52. Bostrichidae (Apatidae of some authors). Tarsi 
five-jointed, but the first joint very short and imperfectly separated 

from the second; front coxae prominent, con- 
tiguous, very little extended transversely ; five 
visible ventral segments. The Bostrichidae 
attack dry wood, and sometimes in such 
large numbers that timber is entirely 
destroyed by them ; most of them make 
cylindrical burrows into the wood. The 
larvae have the posterior part of the 
body incurved, and resemble the wood- 
boring larvae of Anobiidae rather than 
the predaceous larvae of Cleridae. We 
follow Leconte and Horn in placing 
Lyctides as a division of Bostrichidae ; 
although differing very much in appear- 

FIG. 125.-Apate capudna. ance > the 7 have similar habits and larvae. 
Europe. A, Larva (after The typical Bostiichides are remarkable 

Ferris) ; B, pt-rt'ect III- n , i , n 

sect . tor their variety of sculpture and for the 

shapes of the posterior part of the body : 

this part is more or less conspicuously truncate, and furnished 
with small prominences. Dinapate u-riyhtii, found in the stems 
of a species of Yucca in the Mojave desert of California, attains 
a length of nearly two inches ; its larva is extremely similar to 
that of A. capudna, Some of the forms (Plionapate} stridulate 
in a manner peculiar to themselves, by rubbing the front leg 
against some projections at the hind angle of the prothorax. Up- 
wards of 200 species of the family are known. In Britain we 
have only four small and aberrant forms. 

Fam. 53. Ptinidae. Tarsi five-jointed, first joint not reduced 
in size, often longer than second ; front and middle coxae small, 
not transversely extended, the former sliglitly prominent; five 
visible ventral segments; prostenuuii very short. Here are in- 
cluded two sub - families, Ptinides and Anobiides ; they are 
considered as distinct families by many authors, but in the 




POLYMORPHA PTINIDAE 



247 




FIG. 126." Biscuit-weevil." 
A nobium pan iceum. 



present imperfect state of knowledge l it is not necessary to treat 
them separately. Ptinidae are sometimes very destructive to dried 
animal matter, and attack specimens in 
museums ; Anobiides bore into wood, and 
apparently emerge as perfect Insects only 
for a very brief period; Anobiutn ($ito- 
il 1-1 />*) paniceum is, however, by no means 
restricted in its tastes ; it must possess 
extraordinary powers of digestion, as we 
have known it to pass several consecu- 
tive generations on a diet of opium ; it 
has also been reported to thrive on 
tablets of dried compressed meat ; in 
India it is said to disintegrate books ; a 
more usual food of the Insect is, how- 
ever, hard biscuits ; weevilly biscuits are 
known to every sailor, and the so-called " weevil " is usually 
the larva of A. paniceum (Fig. 127, B). In the case of this Insect 
we have not detected more than one spiracle (situate on the 
first thoracic segment) ; the other known larvae of Anobiides are 
said to possess eight abdominal spiracles. The skeleton in some 



FIG. 127. Early 
stages of Anobium 
paniceum. A, Eggs, 
variable in form ; 
B, larva ; C, pupa ; 
D, asymmetrical 
processes terminat- 
ing body of pupa. 
[This larva is pro- 
bably the "book- 
worm " of librar- 
ians]. 



of this sub-family is extremely modified, so as to allow the 
Insects to pack themselves up in repose ; the head is folded in 
over the chest, and a cavity existing on the breast is thus closed 
by the head ; in this cavity the antennae and the prominent 
mouth-parts are received and protected ; the legs shut together 

1 It is probable that we do not know more than the fiftieth part of the existing 
species, most of which lead lives that render them very difficult to find. 






248 COLEOPTERA CHAP. 

in an equally perfect manner, so that no roughness or chink 
remains, and the creature looks like a little hard seed. Anobium 
striatuin is a common Insect in houses, and makes little round 
holes in furniture, which is then said to be " worm-eaten." A. 
(Xestoln.tnn*) tessellatum , a much larger Insect, has proved very 
destructive to beams in churches, libraries, etc. These species 

are the "death-watches" or "greater 
death-watches " that have been associated 
with the most ridiculous superstitions 
(as we have mentioned in Volume V., 
when speaking of the lesser death- 
watches, or Psocidae). The ticking of 
these Insects is really connected with sex, 
and is made by striking the head rapidly 
against the wood on which the Insect 

O 

is standing. 

The very anomalous genus Ectreplies 
FIG. 128. Ectrepkes kingi. J 

West Australia. (After (Fig. 128) is found ill ants' liests ill 

Westwood.) Australia. Westwood placed it in Tti- 

nidae. Wasmann has recently treated it as a distinct family, 
Ectrephidae, associating it with Polyplocotes and Diplocotes, and 
treating them as allied to Scydmaenidae. 

Fam. 54. Malacodermidae. Kecen (or even, eiglif) visible ven- 
tral segments, tlie lasal one not co-adapted in form with the coxae ; 
tarsi Jive-jointed. Integument softer than usual, the ^orfe of tJie 
body not accurately co-ad xpti-d. This important family includes 
a variety of forms: viz. Lycides, Drilides, Lampyrides, Telepho- 
rides ; though they are very different in appearance, classifiers 
have not yet agreed on separating them as families. Of these 
the Lampyrides, or glow-worms, are of special interest, as most 
of their members give off a phosphorescent light when alive ; in 
many of them the female is apterous and like a larva, and then 
the light it gives is usually conspicuous, frequently much more 
so than that of its mate ; in other cases the males are the most 
brilliant. The exact importance of these characters in the crea- 
tures' lives is not yet clear, but it appears probable that in the 
first class of cases the light of the female serves as an attraction 
to the male, while in the second class the very brilliant lights of 
the male serve as an amusement, or as an incitement to rivalry 
amongst the individuals of this sex. The well-known fire-Hies 



V 



MALACODERMIDAE FIRE-FLIES GLOW-WORMS 



249 



(Ludol(C) of Southern Europe are an example of the latter con- 
dition. They are gregarious, and on calm, warm nights crowds 
of them may be seen moving and sparkling in a charm- 
ing manner. These individuals are all, or nearly all, males ; 
so rare indeed is the female that few entomologists have even 
noticed it. The writer once assisted in a large gathering of 
Luciola italica in the Yal Anzasca, which consisted of many 
hundreds of specimens ; all of those he caught, either on the 
wing or displaying their lights on the bushes, were males, but 
he found a solitary female on the ground. This sex possesses 
ordinary, small eyes instead of the 
large, convex organs of the male, 
and its antennae and legs are 
much more feeble, so that though 
provided with elytra and wings 
it is altogether a more imper- 
fect creature. Emery has given 
an account of his observations 
and experiments on this Insect, 
but they do not give any clear 
idea as to the exact function of 
the light. 1 In our British glow- 
worm the female is entirely apter- 
ous hence the name gloW-WOrm FIG. 129. Phengodes hieronymi. Cor- 
doba, South America. (After Haase.) 
A. Male ; B, female. I, I, Positions 
of luminous spots ; Is, spiracles. About 
x 3 
at night into lighted apartments. 




-but the male has elytra and 
ample wings, and frequently flies 



Although so little has been ascertained as to the light of Lampy- 
ridae, there are two facts that justify us in supposing that it is in 
some way of importance to the species. These are: (1) that in a great 
many species the eyes have a magnificent and unusual develop- 
ment ; (2) that the habits of the creatures are in nearly all cases 
nocturnal. It is true that the little Phosphaenus liemipterus is 
said to be diurnal in habits, but it is altogether an exceptional 
form, being destitute of wings in both sexes, and possessed of 
only very feeble light -giving powers, and we have, moreover, 
very little real knowledge as to its natural history : it is said 

1 Bull. cnt. Hal. 1886, p. 406, and Ent. Zeit. Stettin, xliii. 1887, pp. 201-206. 
Kmrry does not mention the name of the species, but \ve presume it to be the 
common Italian fire-fly, Luciola italica. 



250 COLEOPTERA CHAP. 

that the female is of the utmost rarity, though the male is not 
uncommon. 

The nature of the luminosity of Lampyris has given rise 
to many contradictory statements ; the light looks somewhat 
like that given off by phosphorus, and is frequently spoken of 
as phosphorescence ; but luminescence is a better term. The 
egg, larva, pupa, and male are luminous as well as the female 
(at any rate in L. noctilma^) ; the luminescence is, however, most 
marked in the female imago, in which it is concentrated near 
the extremity of the abdomen ; here there are t\vo strata of 
cells, and many fine capillary tracheae are scattered through the 
luminous substance. Wielowiejski concludes that the light- 
producing power is inherent in the cells of the luminous organ, 
and is produced by the slow oxidation of a substance formed 
under the influence of the nervous system. The cells are 
considered to be essentially similar to those of the fat-body. 1 
The luminescence of Lampyridae is very intermittent, that is to 
say, it is subject to rapid diminutions and increases of its 
brilliancy ; various reasons have been assigned for this, but all 
are guesses, and all that can be said is that the changes are 
possibly due to diminution or increase of the air-supply in the 
luminous organ, but of the way in which this is controlled there 
seems to be no evidence. Considerable difference of opinion has 
existed as to the luminescence of the eggs of Lampi/ris. If it exist 
in the matter contained in the egg, it is evident that it is 
independent of the existence of tracheae or of a nervous system. 
Newport and others believed that the light given by the egg 
depended merely on matter on its exterior. The observations of 
Dubois ' 2 show, however, that it exists in the matter in the egg ; 
he has even found it in the interior of eggs that had been 
deposited unfertilised. 

From time to time, since the commencement of the nineteenth 
century, there have appeared imperfect accounts of extraordinary 
light-giving larvae found in South America, of various sizes, but 
attaining in some cases a length, it is said, of three inches ; they 
are reported as giving a strong red light from the two extremities 
of the body, and a green light from numerous points along the 

1 Zcitschr. iviss. Zool. xxxvii. 1882, p. 354 ; also Emery, op. cit. xl. 1S84, 
p. 338. For another theory as to the luminescence, see p. 259. 

2 Bull. Soc. Zool. France, xii. 1887, p. 137, postea. 



v POLYMORPHA GLOW-WORMS 251 

sides of the body, and hence are called, it is said, in Paraguay 
the railway-beetle. We may refer the reader to Haase's paper 1 
on the subject of these " larvae," as we can here only say that it 
appears probable that most of these creatures may prove to be 
adult females of the extraordinary group Phengodini, in which it 
would appear that the imago of the female sex is in a more larva- 
like state than it is in any other Insects. The males, however, 
are well-developed beetles ; unlike the males of Lampyrides, in 
general they have not peculiar eyes, but on the other hand they 
possess antennae which are amongst the most highly developed 
known, the joints being furnished on each side with a long 
appendage densely covered with pubescence of a remarkable 
character. There is no reason to doubt that Haase was correct 
in treating the Insect we figure (Fig. 129, B) as a perfect Insect ; 
he is, indeed, corroborated by Eiley. 2 The distinctions between the 
larva and female imago are that the latter has two claws on the 
feet instead of one, a greater number of joints in the antennae, 
and less imperfect eyes ; the female is in fact a larva, making a 
slightly greater change at the last ecdysis, than at those previous. 
It is much to be regretted that we have so very small a know- 
ledge of these most interesting Insects. Malacodermidae are 

O O 

probably the most imperfect or primitive of all beetles, and it 
is a point of some interest to find that in one of them the 
phenomena of metamorphosis are reduced in one sex to a 
minimum, while in the other they are presumably at least 
normal in character. 

Numerous larvae of most extraordinary, though diverse, 
shapes, bearing long processes at the sides of the body, and 
having a head capable of complete withdrawal into a slender 
cavity of the thorax, have long been known in several parts of 
the world, and Dr. AVilley recently found in Xew Britain a 
species having these body-processes articulated. Though they 
are doubtless larvae of Lampyrides, none of them have ever been 
reared or exactly identified. 

A very remarkable Ceylouese Insect, Dioptoma adamsi 
Pascoe, is placed in Lampyrides, but can scarcely belong there, as 
apparently it has but five or six visible ventral segments ; this 
Insect has two pairs of eyes, a large pair, with coarse facets on 

1 Deutsche ent. Zeitsclir. xxxii. 1888, pp. 145-167. 
- Ent. Mag. xxiv. 1887, p. 148. 



252 



COLEOPTERA 



CHAP. 



the under side of the head, and a moderate-sized pair with fine 
facets on the upper side. Nothing is known as to the habits of 
this curiosity, not even whether it is luminous in one or both sexes. 
It is believed that the perfect instar of Lampyrides takes no 
food at all. The larvae were formerly supposed to be vegetarian, 
but it appears probable that nearly all are carnivorous, the chief 
food being Mollusca either living or dead. The larvae are active, 
and in many species look almost as much like perfect Insects as 
do the imagos. 

The other divisions of Malacodermidae Lycides, Drilides, 
Telephorides also have predaceous, carnivorous larvae. All these 
groups are extensive. Though much neglected by collectors and 
naturalists, some 1500 species of the family Malacodermidae 
have been detected. We have about 50 in Britain, and many 
of them are amongst the most widely distributed and abundant 
of our native Insects. Thus, however near they may be to the 
primitive condition of Coleoptera, it is highly probable that they 
will continue to exist alongside of the primitive Cockroaches and 
Aptera, long after the more highly endowed forms of Insect-life 
have been extinguished wholesale by the operations of mankind 
on the face of the earth. 

Fam. 55. Melyridae (or Malachiidae). Six visible and move- 
able ventral abdominal segments; the basal part more or less distinct I;/ 

co-ailiijit, <i with the coxae. These 
Insects are extremely numerous, 
but have been very little studied. 
In many works they are classified 
with Malacodermidae, but \vere 
correctly separated by Leconte 
and Horn, and this view is also 
taken by Dr. Yerhoelf, the latest 
investigator. The smaller num- 
ber of visible ventral segments 
appears to be due to a change 
at the base correlative with an 
adaptation between the base of 
Britain, the abdomen and the hind 
The characters are singu- 
with those of 




FIG. 130. Malacliius aeneus. 

A, Larva (after Ferris) ; B, female 
imago. 



coxae. 

larly parallel 
Silphidae ; but in Melyridae the antennae are filiform or serrate, 



v POLYMORPHA MELYRIDAE CLERIDAE 253 

not clavate. The habits in the two families are different, as the 
Melyridae are frequenters of flowers. Many of the Melyridae 
have the integument soft, but in the forms placed at the end 
of the family e.g. Zyyia they are much firmer. Thus these 
Insects establish a transition from the Malacodermidae to ordi- 
nary Coleoptera. Although the images are believed to consume 
some products of the flowers they frequent, yet very little is 
really known, and it is not improbable that they are to some 
extent carnivorous. This is the case with the larvae that are known 
(Fig. 130, larva of Malachius aeneus}. These are said by Ferris 
to bear a great resemblance to those of the genus Telephorus, 
belonging to the Malacodermidae. 

Fam. 56. Cleridae. Tarsi five-jointed ; but the basal joint of 
the posterior very indistinct, usually very small above, and closely 
united with the second by an oblique splice ; the apices of joints 
two to four usually prolonged as membranous flaps ; anterior coxae 
prominent, usually contiguous, rather large, but their cavities not 
prolonged externally ; labial palpi usually with large hatchet- 
shaped terminal joint ; ventral segments fire or six, very mobile. 
The Cleridae are very varied in form and colours ; the antennae 
are usually more or less clubbed at the tip, and not at all serrate, 
but in Cylidrus and a few others they are not clubbed, and in 
Cylidrus have seven flattened joints. The student should be 
very cautious in deciding as to the number of joints in the feet 
in this family, as the small basal joint is often scarcely dis- 
tinguishable, owing to the obliteration of its suture with the 
second joint. The little Alpine Laricobius has the anterior coxal 
cavities prolonged externally, and the coxae receive the femora to 
some extent, so that it connects Cleridae and Derodontidae. The 
Cleridae are predaceous, and their larvae are very active ; they 
are specially fond of wood-boring Insects ; that of Tillvs elon- 
gatus (Fig. 131) enters the burrows of Ptilinus pectinicornis in 
search of the larva. The members of the group Corynetides 
frequent animal matter, carcases, bones, etc., and, it is said, feed 
thereon, but Perris's recent investigations * make it probable that 
the larvae really eat the innumerable Dipterous larvae found in 
such refuse ; it is also said that the larvae of Cleridae spin 
cocoons for their metamorphosis ; but Ferris has also shown 
that the larvae of Nccrobia ruficollis really use the puparia formed 

1 Larvcs des CoUopteres, 1878, p. 208. 



254 



COLEOPTERA 



CHAP. 



by Diptera. Some of the species of Necrobia have been spread 
by commercial intercourse, and N. rufipes appears to be now one 
of the most cosmopolitan of Insects. The beautifully coloured 
Corynctcs cor rule us is often found in our houses, and is useful, as 
it destroys the death-watches (Anolium} that are sometimes very 
injurious. T'i'icho<les apiarius, a very lively -coloured red and 
blue beetle, destroys the larvae of the honey-bee, and Lampert 

has reared TricJiodes a/reariits 
from the nests of Chalicodoma 
tiiuraria, a mason-bee; he re- 
cords that one of its larvae, 
after being full grown, remained 
twenty - two months quiescent 
and then transformed to a pupa. 
Still more remarkable is a case 
of fasting of the larva of Tri- 
cJiodes ammios recorded by 
Mayet: 1 this Insect, in its 

> 




* 

ew Forest). 



w ;'Tf' 5 ' 

A, Head ; B, front leg ; 



C, termination of the body, more mag- immature form, destrOVS Ad'l- 

ft -t ** 



ft -t 

dium maroccanum ; a larva sent 

from Algeria to M. Mayet refused such food as was offered to 
it for a period of two and a half years, and then accepted 
mutton and beef as food ; after being fed for about a year and a 
half thereon, it died. Some Cleridae bear a great resemblance 
to Insects of other families, and it appears probable that they 
resemble in one or more points the Insects on which they feed. 
The species are now very numerous, about 1000 being known, 
but they are rare in collections; in Britain we have only nine 
species, and some of them are now scarcely ever met with. 

Fam. 57. Lymexylonidae. Elongate 1)eef/>-*, v/7/6 soft integu- 



ments, front and middle coxae exserted, longitudinal in 
hirxi slender, Jive-jointed ; (t/ife/m/tr *////, serrate, Imt I'nfli, r Iroad. 
Although there are only twenty or thirty species of this family, 
they occur in most parts of the world, and are remarkable on 
account of their habit of drilling cylindrical holes in hard wood, 
after the manner of Anobiidae. The larva of Lymex'ijlon navale was 
formerly very injurious to timber used for constructing ships, but 
of late years its ravages appear to have been of little importance. 
The genus Atractocerus consists of a few species of very abnormal 

1 Ann. Sue. cut. France, 1894, p. 7. 



POLYMORPHA DASCILLIDAE 



255 



Coleoptera, the body being elongate and vermiform, the elytra 
reduced to small, functionless appendages, while the wings are 
ample, not folded, but traversed by strong longitudinal nervures, 
and with only one or two transverse nervures. Owing to the 
destruction of our forests the two British Lymexylonidae L. 
navale and Hylecoetus dermestoides are now very rarely met 
with. 

Fam. 58. Dascillidae.- Small or moderate-sized beetles, with 
I'dtlier flimsy integuments, antennae either serrate, filiform, or 
even made flabellate by long appen- 
dages ; front coxae elongate, greatly 
('.'sc/'ted ; abdomen ivith jive mobile 
1-i'ntral segments ; tarsi Jive-jointed. 
This is one of the most neglected 
and least known of all the families 
of Coleoptera, and one of the most 
difficult to classify ; though always 
placed amongst the Serricornia, it 
is more nearly allied to Paruidae 
and Byrrhidae, that are placed in 

Clavicornia, than it is to any of the FlG _ 132 . __ H y drocyphou deflexions. 
ordinary families of Serricornia. It 
is probable that careful study will 
show that it is not natural as at present constituted, and that the 
old families, Dascillidae and Cyphonidae, now comprised in it, will 
have to be separated. Only about 400 species are at present 
known; but as nearly 100 of these have been detected in New 
Zealand, and 17 in Britain, doubtless the numbers in other 
parts of the world will prove very considerable, these Insects 
having been neglected on account of their unattractive exterior, 
;ind fragile structure. The few larvae known are of three or 
four kinds. That of Dascillus cervinus is subterranean, and is 
believed to live on roots ; in form it is somewhat like a Lamelli- 
corn larva, but is straight, and has a large head. Those of the 
< 'yphonides are aquatic, and are remarkable for possessing antennae 
consisting of a great many joints (Fig. 132, A). Tournier 
describes the larva of Helodes as possessing abdominal but not 
thoracic spiracles, and as breathing by coming to the surface of 
the water and carrying down a bubble of air adhering to the 
posterior part of the body; the larva of Hydrocyplion (Fig. 132, A) 




Britain. A, Larva (after Tournier) ; 
B, imago. 



256 



COLEOPTERA 



CHAP. 



possesses several finger-like pouches that can be exstulpated at 
the end of the body. It is probable that these larvae are carni- 
vorous. The imago of this Insect abounds on the bushes along 
the banks of some of the rapid waters of Scotland ; according to 
Touriiier, when alarmed, it enters the water and goes beneath it 
for shelter. The third form of larva belongs to the genus Euci- 
netus, it lives on fungoid matter on wood, and has ordinary 
antennae of only four joints. 1 It is very doubtful whether 
Eucinetas is related to other Dascyllidae ; some authorities indeed 
place it in Silphidae. 

Fam. 59. Rhipiceridae.--^?^ fire -jointed, furnished u-ith 
a robust onychium (a straight chitinons process bearing hairs) 

between th e claws ; a ntennae of 
the male bearing long processes, 
and sometimes consisting of a 
large manlier of joints. 3fan- 
dibles robust, strongly curved, 
and almost calliper -like in 
form. This small family of 
less than 100 species is widely 
distributed, though confined to 
the warmer regions of the 
earth, a single species occur- 
ring in the extreme south of 
Eastern Europe. Very little 
is known as to the natural 
history. The larva of Calli- 
rhipis dcjcani (Fig. 133, A) is 
described by Schiodte as hard, 
cylindrical in form, and peculi- 
truncate behind, so that 




FIG. 133. A, Larva of GaUirhqris dejeani 

(after Schiiidte) ; B, Rliipicrra mystacina there appear to be Ollly eight 
male, Australia ; C, under side of its hind , , i 

foot. abdominal segments, trie ninth 

segment being so short as to 

look like an operculum at the extremity of the body. It lives 
in wood. 

Fam. 60. Elateridae (Click-beetles]. Antennae more or less 
serrate along the inner margin, frequently pei'fi /i/<\ rarely 
filiform. Front coxae small, spherical. Thorax -usually with 

1 Perris, Ann. >SW. ent. France (2) ix. 1851, p. 48. 



CLICK-BEETLES 



257 




FIG. 13-1. Athous rhombeus. New 
Forest. A, Larva ; B, female 
imago. 



J/ i /if? nn riles more or less prolonged backwards; with a prosternal 
process that can he received in, and usually can move in, a 
mesosternal cavity. Hind coxa with a plate, above which the femur 
can lie received. Visible ventral segments 'usually Jive, only the 
if i- initial one being mobile. Tarsi Jive-jointed. This large family 
of Coleoptera comprises about 7000 
species. Most of them are readily 
known by their peculiar shape, and 
by their faculty of resting on the 
back, stretching themselves out 
flat, and then suddenly o-oing off 

t/ O O 

with a click, and thus jerking 
themselves into the air. Some, 
however, do not possess this faculty, 
and certain of these are extremely 
difficult to recognise from a defi- 
nition of the family. According 
to Bertkau 1 our British Lacon 
murinus is provided near the 
tip of the upper side of the ab- 
domen with a pair of eversible 

glands, comparable with those that are better known in Lepi- 
dopterous larvae. He states that this Insect does not try 
to escape by leaping, but shams death and " stinks away ' its 
enemy. The glands, it would appear, become exhausted after the 
operation has been repeated many times. The extent of the leap 
executed by click -beetles differs greatly ; in some species it is 
very slight, and only just sufficient to turn the Insect right side 
up when it has been placed on its back. In some cases the 
Insects go through the clicking movements with little or no 
appreciable result in the way of consequent propulsion. Although 
it is difficult to look on this clicking power as of very great value 
to the Elateridae, yet their organisation is profoundly modified so 
us to permit its accomplishment. The junction of the prothorax 
with the after-body involves a large number of pieces which an- 
all more or less changed, so that the joint is endowed with greater 
mobility than usual; while in the position of repose, on the other 
hand, the two parts are firmly locked together.' The thoracic 
stigma is of a highly remarkable nature, and the extensive 

1 Arch. Naturgesch. xlviii. 1, 1882, p. 371. 
VOL. VI S 



258 COLEOPTERA 



CHAP. 



membrane in which it is placed appears to be elastic. Although 
the mechanics of the act of leaping are still obscure, yet certain 
points are clear ; the prosternal process possesses a projection, or 
notch, on its upper surface near the tip ; as a preliminary to 
leaping, this projection catches against the edge of the meso- 
sterual cavity, and as long as this position is maintained the 
Insect is quiescent ; suddenly, however, the projection slips over 
the catch, and the prosternal process is driven with force and 
rapidity into the mesosternal cavity pressing against the front 
wall thereof, and so giving rise to the leap. 

Several larvae are well known ; indeed the " wire-worms " 
that are sometimes so abundant in cultivated places are larvae 
of Elateridae. In this instar the form is usually elongate and 
nearly cylindrical ; the thoracic segments differ but little from 
the others except that they bear rather short legs ; the skin 
is rather hard, and usually bears punctuation or sculpture ; 
the body frequently terminates in a very hard process, of 
irregular shape and bearing peculiar sculpture on its upper 
surface, while beneath it the prominent anal orifice is placed : this 
is sometimes furnished with hooks, the function of which has 
not yet been observed. The majority of these larvae live in 
decaying wood, but some are found in the earth ; as a rule the 
growth is extremely slow, and the life of the larva may extend 
over two or more years. Some obscurity has prevailed as to their 
food ; it is now considered to be chiefly flesh, though some species 
probably attack decaying roots ; and it is understood that wire- 
worms destroy the living roots, or underground stems, of the crops 
they damage. Various kinds of Myriapods (see Vol. Y. p. 29) are 
often called "wire-worm," but they may be recognised by possessing 
more than six legs. The larvae of the genus Cardiophorus are very 
different, being remarkably elongate without the peculiar terminal 
structure, but apparently composed of twenty-three segments. 

The genus Pyrophorus includes some of the most remarkable 
of light-giving Insects. There are upwards of 100 species, 
exhibiting much diversity as to the luminous organs ; some 
are not luminous at all ; but all are peculiar to the Xew "\Vorld, 
with the exception that there may possibly be luminous species, 
allied to the American forms, in the Fiji Islands and the New 
Hebrides. In the tropics of America the Pyrophorits, or Cucujos, 
form one of the most remarkable of the natural phenomena. 



v ELATERIDAE FIRE-FLIES 259 

The earliest European travellers in the New World were so im- 
pressed by these Insects that descriptions of their wondrous display 
occupy a prominent position in the accounts of writers like Oviedo, 
whose works are nearly 400 years old. Only one of the species 
has, however, been investigated. P. noctiliicus is one of the most 
abundant and largest of the Pyroplnirux, and possesses 011 each side 
of the thorax a round polished space from which light is given 
forth ; these are the organs called eyes by the older writers. 
Besides these two eye-like lamps the Insect possesses a third 
source of light situate at the base of the ventral surface of the 
abdomen ; there is no trace of this latter lamp when the Insect 
is in repose ; but when on the wing the abdomen is bent away from 
the breast, and then this source of light is exposed ; hence, when 
Hying, this central luminous body can be alternately displayed 
and concealed by means of slight movements of the abdomen. 
The young larva of P. noctilucus is luminous, having a light- 
giving centre at the junction of the head and thorax ; the older 
larva has also numerous luminous points along the sides of the 
body near the spiracles. It is remarkable that there should 
be three successive seats of luminescence in the life of the same 
individual. The eggs too are said to be luminous. The light 
given off by these Insects is 'extremely pleasing, and is used by 
the natives on nocturnal excursions, and by the women for orna- 
ment. The structure of the light-organs is essentially similar 
to that of the Lampyridae. The light is said to be the most 
economical known ; all the energy that is used being converted 
into light, without any waste by the formation of heat or 
chemical rays. The subject has been investigated by Dubois, 1 who 
comes, however, to conclusions as to the physiology of the 
luminous processes different from those that have been reached 
by Wielowiejski and others in their investigations 011 Glow- 
worms. He considers that the light is produced by the reactions 
of two special substances, luciferase and luciferine. Lucifernse 
is of the nature of an enzyme, and exists only in the luminous 
organs, in the form, it is supposed, of extremely minute granules. 
Luciferine exists in the blood ; and the light is actually evoked 
by the entry of blood into the luminous organ. 

We have given to this family the extension assigned to it by 

1 " Les Elateridea hnniueux," Bull. Soc. Zool. France, xi. 1SS6 ; also Lemons 
dc I'/ii/xin/tii/i'i' iji'ngralc, Paris. 1898, and C.R. Ac. Sci. cxxiii. 1S96, p. 653. 



260 



COLEOPTERA 



CHAP. 



Schiodte. Leconte and Horn also adopt this view, except that 
they treat Throscides as a distinct family. By most authors 
Eucnemides, Throscides, and Cebrionides are all considered dis- 
tinct families, but at present it is almost impossible to separate 
them on satisfactory lines. The following table from Leconte 
and Horn exhibits the characters of the divisions so far as the 
imago is concerned : 

Posterior coxae laminate ; trochanters small. 

Labrum concealed ; antennae somewhat distant from the eyes, their 



insertion narrowing the front 



EUCXEMIDES. 



Labnuu visible, free; antennae arising near the eyes under the frontal 
margin ..... ELATERIDES. 

Lalirum transverse, connate with the front. 

Ventral segments six ; claws simple ; tibial spurs well developed. 

( 'EBRIOXIDES. 
Ventral segments five ; claws serrate ; tibial spurs moderate. 

PEROTHOPIDES. 

Posterior coxae not laminate ; trochanters of middle and posterior legs very 
long . . ... CEROPHYTIDES. 

Throscides are considered to be distinguished by the 

mesosternum being impressed on each side 
in front for the accommodation of the 
posterior face of the front coxae. The 
genus Throscus has the antennae clavate. 
The classification of the Elaterides and 
these forms is a matter of the greatest 
difficulty, and, if the larvae are also 
considered, becomes even more complex. 
Cebrionid larvae are different from 
those of any of the other divisions, and 
possess laminate, not calliper-like, man- 
dibles. The larvae of Eucnemides (Fig. 
135) are very little known, 1 >ut are 
highly remarkable, inasmuch as it is 

FIG. 135. -Larva of Forna.c difficult to lilld any mouth-opeiling 

n. sp. Hawaii. A, Upper 

side; B, under side: s *, in some of them, and they have no legs. 
Ei^ore Srjd \ g The other divisions possess very few species 
under side of terminal seg- compared with Elaterides. Iii Britain 

ment ; a. anus. i > p T-II 

we have about sixty species ot Elate- 
rides, four of Throscides and three of Eucnemides ; Cerophytum 
was probably a native many years ago. Neither Perothopides 




POLYMORPHA BUPRESTIDAE 



26l 



nor Cebrionides are represented in our fauna ; the former of these 
two groups consists only of four or five North American species, 
and the Cerophytides are scarcely more numerous. 

Fam. 61. Buprestidae. Antennae serrate, never elongate ; 
prothorax fitting closely to the after-body, with a process received 
into a cavity of the mesosternum so as to permit of no movements 
of nutation. Five visible ventral segments, the first usually 
elongate, closely 'united with, t/ic second, the others mobile. Tarsi 
five-jointed, the first four joints usually with membranous pads 
beneath. This family is also of large extent, about 5000 species 
being known. Many of them are remarkable for the magnificence 
of their colour, which is usually metallic, and often of the greatest 
brilliancy ; hence their wing-cases are 
used by our own species for adorn- 
ment. The elytra of the eastern kinds 
of the genus Sternocera are of a very 
brilliant green colour, and are used 
extensively as embroidery for the 
dresses of ladies ; the bronze elytra 
of Buprestis (Euchroma) gigantea 
were used by the native chieftains in 
South America as leg -ornaments, a 
large number being strung so as to 
form a circlet. The integument of 
the Buprestidae is very thick and hard, 
so as to increase the resemblance to 
metal. The dorsal plates of the abdo- 
men are usually soft and colourless in 
beetles, but in Buprestidae they are 
often extremely brilliant. The metallic colour in these Insects is 
not due to pigment, but to the nature of the surface. Buprestidae 
appear to enjoy the hottest sunshine, and are found only where there 
is much summer heat. Australia and Madagascar are very rich 
in species and in remarkable forms of the family, while in Britain 
we possess only ten species, all of which are of small size, and 
nearly all are excessively rare. The family is remarkably rich in 
fossil forms; no less than 28 percent of the Mesozoic beetles 
found by Heer in Switzerland are referred to Buprestidae. 

The larvae (Fig. 136, A) find nourishment in living vegetable 
matter, the rule being that they form galleries in or under the 




FIG. 136. A, Larva of Euchroma 
(joliath (after Schiodte) ; B, imago 
of MdanopMla decostiyma. 
Europe. 



262 COLEOPTERA 



CHAP. 



bark of trees and bushes, or in roots thereof; some inhabit the 
stems of herbaceous plants and one or two of the smaller forms have 
been discovered to live in the parenchyma of leaves. A few are said 
to inhabit dead wood, and in Australia species of Etlion dwell in 
galls on various plants. Buprestid larvae axe of very remarkable 
shape, the small head being almost entirely withdrawn into the 
very broad thorax, while the abdomen is slender. 1 A few, how- 
ever, depart from this shape, and have the thoracic region but 
little or not at all broader than the other parts. The larvae of 
Julodis a genus that inhabits desert or arid regions are 
covered with hair ; they have a great development of the 
mandibles ; it is believed that they are of subterranean habits, 
and that the mandibles are used for burrowing in the earth. 
Only the newly hatched larva is, however, known. 



Series IV. Heteromera. 

Tarsi of the front and middle legs with five, tJiOse of the hind 

legs with four, joints. 

This series consists of some 14,000 or 15,000 species. 
Twelve or more families are recognised in it, but the majority of 
the species are placed in the one great family, Tenebrionidae. 
The number of visible ventral segments is nearly always live. 
Several of the families of the series are of doubtful validity ; 
indeed beyond that of Tenebrioiiidae the taxonomy of this series 
is scarcely more than a convention. The larvae may be con- 
sidered as belonging to three classes : one in which the body is 

o o */ 

cylindrical and smooth and the integument harder than usual in 
larvae ; a second in which it is softer, and frequently possesses 
more or less distinct pseudopods, in addition to the six thoracic 
legs; and a third group in which hypermetamorphosis prevails, 
the young larvae being the creatures long known as Triungulins, 
and living temporarily on the bodies of other Insects, so that 
they were formerly supposed to be parasites. 

1 It seems impossible to understand the morphology of the anterior segments by 
mere inspection ; the anterior spiracle being seated on the segment behind the 
broad thorax. Considerable difference of opinion has prevailed as to what is head, 
what thorax ; the aid of embryology is necessary to settle the point. The larva 
described by AVestwood (Mod. Classif. i. 1839, p. 229), and figured as probably 
Buprestis atttximf". is doubtless a Passalid. 



V 



HETEROMERA 



263 



3 

1 




h-h mditor. Europe, etc. A, 

Larva (meal-worm); B, pupa (after 

Schiodte) ; c, imago. 



Fam. 62. Tenebrionidae. Front coxae short, not projecting 
from the cavities, enclosed InhimJ. Feet destitute of lobed joints. 
Clmcs smooth. This is one of the largest families of Coleoptera, 
about 10,000 species being already known. A very large portion 
of the Tenebrionidae are entirely terrestrial, wings suitable for flight 
being absent, and the elytra frequently more or less soldered. Such 
forms are described in systematic works as apterous. Unfortunately 
no comprehensive study has 
ever been made of the wings 
or their rudiments in these 
" apterous forms." It is prob- 
able that the wings, or their 
rudiments or vestiges, always 
exist, but in various degrees of 
development according to the 
species, and that they are never A y 
used by the great majority of the 
terrestrial forms. Man v of the Flc - 137 - 

, ,. m . . . .. 

wood - feeding Tenebrionidae, 
and the genera usually placed 

at the end of the family, possess wings well adapted for flight. 
The apterous forms are chiefly ground -beetles, living in dry 
places ; they are very numerous in Africa, California, and 
North Mexico. Their colour is nearly always black, and this is 
probably of some physiological importance ; the integuments are 
thick and hard, and if the wing-cases are taken off, it will be 
found that they are usually more or less yellow on the inner face, 
even when jet-black externally ; the external skeleton is very closely 
fitted together, the parts that are covered consisting of very delicate 
membrane ; the transition between the hard and the membranous 
portions of the external skeleton is remarkably abrupt. These 
ground-Tenebrionidae form a very interesting study, though, on 
account of their unattractive appearance, they have not received 
the attention they deserve. 

Many of the Tenebrionidae, notwithstanding their dark 

1 Casey has examined the wings in the genus Blapstinus (an "apterous" 
genus), and found that the wings are extremely varied in development, according 
to the species ; in no case, however, did they appear to be capable of giving more 
than a laboured and feeble flight. Ann. Neio York Ac. v. 1890, p. 416. 

In Eleodcs, though the meso- and meta-notum are formed of delicate membrane, 
the wings exist as minute flaps, requiring some examination for their detection. 



264 COLEOPTERA CHAP. 

colours, are diurnal in habits, and some of them run 
with extreme velocity in places so bare and desert that the 
means of existence of the Insects is a mystery. Most of the 
Tenebrionidae, however, shun the light. The food is usually 
vegetable matter, and it is apparently preferred in a very dry 
state. Mr. Gahan has recently recorded that in Praogena 
the under surface of the head has the gular region striate for 
stridulating purposes. This is the only instance known of a 
voice -organ in this situation, and moreover is the only case 
in all the Tenebrionidae in which any sound-producing organ 
has been discovered. The larvae exhibit but little variety, they 
are elongate and cylindrical, with harder integument than is usual 
in Coleopterous larvae ; they have six thoracic legs, and at the 
under side of the posterior extremity the anus serves as a very 
short pseudopod. The resemblance of these larvae to those of 
Elateridae is considerable : but though the body is terminated by 
one or two small processes, these never attain the complexity of 
the terminal segment of Elateridae. The common meal-worm 
i.e. the larva of Tenebrio molitor is a very characteristic example 
of the group. The pupae are remarkable on account of peculiar 
projections, of varied and irregular form, that exist on the sides 
of the abdominal segments. Britain is very poor in these Insects ; 
our list of them scarcely attains the number of thirty species. 

Fam. 63. Cistelidae. Claws comb-like. The very obscure 
beetles forming this family are only separated from Tenebrionidae 
on account of their pectinate claws. About 500 species of Cis- 
telidae are recorded ; the early instars, so far as known, do not 
differ from those of Tenebrionidae ; the larvae are believed to live 
on dead wood. 

Fam. 64. Lagriidae. Anterior coxal cavities dosed, tips of 
the front coxae free, claws smooth, penultimate joint of the tarti 
broader, j>t>l>rxecnt beneath. This family has very little to dis- 
tinguish it from Tenebrionidae, and the group Heterotarsini 
appears to connect the two. It is a small family of about 200 
species, widely distributed, and represented in Britain by one 
species, Lai/fia hirta. The early instars are similar to those of 
the Tenebrionidae, except that the larva is less retiring in its 
habits and wanders about on foliage: it is of broader form than 
that of most of the Tenebrionidae. The pupa has long projections 
at the sides of the abdominal segments. 



HETEROMERA 265 



Fam. 65. Othniidae. Only about ten species are known of this 
dubious family. They are small Insects with weak integument, 
and are said by Leconte and Horn to be distinguished from 
" degraded Tenebrionidae " by the more mobile abdominal seg- 
ments, the hind-margins of which are semi -membranous. The 
antennae are of the clubbed shape, characteristic of " Clavicornia," 
but this also occurs in numerous undoubted Tenebrionidae. 
Species of Otlinius have been found in Japan and Borneo, as 
well as in North America. Nothing is known as to their 
metamorphoses. 

Fam. 66. ^Egialitidae. All the coxae very ividely separated ; 
mi co-adaptation Between the sides of the abdomen and the edges of 
the wing-cases ; five ventral segments and tip of a sixth visible. 
Two minute and rare Insects from North-West America constitute 
this family. It is distinguished from Pythidae by the minute 
front coxae, widely separated, completely closed in, and deeply 
embedded in the prosternum. 

Fam. 67. Monommidae.--This is a small family of less than 
100 species, the members of which have the details of their 
external structure much modified, permitting the Insect to pack 
itself up in repose in a very perfect manner. * They are of small 
size and oval form ; and are absent from Europe and the Anti- 
podes. Nothing appears to be known as to the metamorphosis. 

Fam. 68. Nilionidae. Broad, circular Heteromera, of moderate 
size, with the front coxae but little separated, and the anterior 
acetabula closed, though having the appearance of being open in 
consequence of the tips of the epimera being free. The infle,<-/ 
portion of the wing-cases remarkably broad. A small family of 
less than fifty species, found on fungi, chiefly in South America. 
The metamorphoses are not known. It is of very doubtful 
validity. 

Fam. 69. Melandryidae. Head not constricted, behind the 
eyes ; anterior acetabula not closed ; claws smooth. Prothorax broad 
behind. These are loosely -fitted- together Insects, of moderate 
or small size, frequenting dry wood or fungi. About 200 species 
are known, found chiefly in temperate regions. The few described 
larvae are rather varied in their details and cannot be generalised 
at present. The characters of the members of this family require 
fresh investigation. 

Fam. 70. Pythidae. Distinguished from Melandryidae by the 



266 COLEOPTERA 



CHAP. 



prothorax being narrow behind. This is a small family of about 
100 species, found in temperate regions in connection with 
timber. The species of Iihi/msum/s have the head prolonged in 
front of the antennae so as to form a beak. The larva of Pytho 
</,'ji,'i'xsi/s is flat and has parallel sides ; the body is terminated by 
two widely-separated sharp processes. It is found occasionally 
under the bark of firs in Scotland. 

Fam. 71. Pyrochroidae. Differs from Melandryidae by the 
head forming a very narrow neck behind, and by the penultimate 
tarsal joints being broad. They are feeble Insects, though active 
on the wing. They are destitute of any of the various remark- 
able structures found in Mordellidae. Only about forty species 
are known, and the family is confined to the north temperate 
region, being best represented in Japan. Pyrocliroa rubens is 
common in some parts of England ; the larva is found under 
the bark of tree-stumps ; it is remarkably flat, and has the eighth 
abdominal segment unusually long, while the ninth terminates 
the body in the form of two long sharp processes. 

Fam. 72. Anthicidae. Head v:ith an abrupt narrow neck ; 
'jirafhorax narrower than the elytra. Middle and hind coxae placed 
in definite aeetalnda. Cla-ics single. These little Insects are 
numerous in species ; they have little resemblance to Pyrochroidae, 
though the characters of the two families cause us to place 
them in proximity. There are about 1000 species known; 
though we have only about 12 in Britain, they are very 
numerous in the Mediterranean region. The family Pedilidae 
of Lacordaire and some others is now merged in Anthicidae. 
Thomson and Champion, on the other hand, separate some very 
minute Insects to form the family Xylophilidae, on account of 
certain differences in the form of the abdomen and tarsi. The 
Xylophilidae live in dead wood ; the Anthicidae, on the surface of 
the earth, after the manner of ground-beetles ; very little is, 
however, known as to their natural history. 

Fam. 73. Oedemeridae. Prothorax not forming sharp edges 
at the sides, head qcifjint/f a narrow neel: Pen /////' t/n/fr tarsal 
joint broad ; cla-n-s smooth. These Insects usually have a feeble 
integument, and bear a certain resemblance to Malacodermidae. 
Less than 500 species are known, but they are widely distri- 
buted, and occur in both temperate and tropical regions. The 
larvae live in old wood. Nac<'/'d>x melanura is common on our 



V 



HETEROMERA MORDELLIDAE 



267 




FIG. 138. Asdera cnernlea. A, Larva ; 
B, pupa (after Scbiudte) ; C, imago. 
Cambridge. 



coasts, where its larva lives in timber cast up by the sea, or 
brought down by Hoods, and it 
is able to resist immersion by 
the tide. It is remarkable from 
the possession of five pairs of 
dorsal false feet on the anterior 
segments, and two pairs on 
the ventral aspect. In Asdera 
caerv.lea there are six dorsal and 
three ventral pairs of these re- 
markable pseudopods. "We have 
six species of Oedemeridae in 
Britain, including Asdera as 
well as Naccrilcs. 

Fam. 74. Mordellidae (incl. 
Rhipiphoridae). Head peculiarly formed, vertex lobcd or ridged 
behind, so that in extension it looses on the front edge of the pvo- 
notum ; capable of great inflection and then covering the pro sternum ; 
hind coxae with laminae forming a sharp edge behind, frequently 
very large. This family is a very distinct one, though it exhibits 
great variety. Lacordaire has pointed out that Rmpiphoridae 
cannot at present be satisfactorily distinguished from Mordellidae. 
Leconte and Horn separate the two by the fact that the sides 
of the prothorax form a sharp edge in Mordellidae, but not in 
Khipiphoridae. A better character would perhaps be found by a 
study of the head, but as this would clearly result in a radical 
change in the composition of the two families it is preferable to 
treat them at present as only sub-families : if placed on a similar 
basis to the preceding families, the group would however form, 
not two, but several families. Besides the unusual shape of the 
head (Fig. 130, D) the ventral region of the body is remarkably 
formed, being very convex, and in many Mordellides terminating 
in a strong spinous process (Fig. 139, C). The elytra are, in 
several Ehipiphorids, of the groups Myoditini and Rhipidiini, 
reduced to a very small size, and the wings are not folded. The 
Mordellidae are remarkable for their activity; in the perfect 
state they usually frequent flowers, and fly and run with extreme 
rapidity. Mordellides are amongst the most numerous and 
abundant of the European Coleoptera, and in Britain the 
Anas-oini swarm on the flowers of bushes and Umbelliferae. The 



268 



COLEOPTERA 



CHAP. 



life-histories appear to be singularly varied : hut unfortunately 
they are incompletely known. The larvae of some of the 
Mordellids have been found in the stems of plants, and 
derive their nutriment therefrom. This is said by Schwarz 
to be undoubtedly the case with Moi'flcUistcna floridensis. 
Coquillett has found the larvae of J/ i>nx1iill. in plant-stems 
under circumstances that render it highly probable that they 
were feeding on a Lepidopterous larva contained in the stems ; 
and Osborn found a similar larva that was pretty certainly a 
Mordellist$na, and fed voraciously on Dipterous larvae in the 
steins of a plant. The little that is known as to the meta- 




FIG. IW.Mordelli- 

stenn tJi'i'iilensis. 
America. (After 
Pdley.) A, Larva ; 
B. pupa ; C, imago ; 
D, outline of de- 
tached head of im- 
ago of M. fin in iln, 
to show the ueck. 



morphoses of Mordclla and Anuspis shows that they live in old 
wood, but does not make clear the nature of their food. 

Although it has been ascertained that the Ehipiphorides 
exhibit instances of remarkable metamorphosis, their life- 
histories are still very imperfectly known. Dr. Chapman has 
ascertained some particulars as to Metoecus paradoxns, which has 
long been known to prey in the larval state on the larvae of the 
common social wasps. 1 The eggs are apparently not deposited in 
the nests of the wasps, but in old wood. The young larva is a 
triungulin, similar to that of the Cantharidae, we shall sub- 
sequently describe. It is not known how it makes its way to 
the was] is' nests, but it is possible that when a wasp visits some 
old wood haunted by these larvae, some of them may attach 
themselves to it and be carried to the wasps' nests. When 

1 Ann. Nat. Jlisf. (4) vi. 1870, p. 314 ; and Ent. Mu<j. xxvii. 1891, p. 18. 



v HETEROMERA CANTHARIDAE 269 

access is gained to the cells the little Metoecus pierces the skin of 
one of the wasp-grubs, and entering in it feeds on the interior ; 
after it has increased in size it emerges, changes its skin, and 
assumes a different form and habits ; subsequently, as an external 
parasite, entirely devouring the wasp-larva, and then becoming a 
pupa, and finally a perfect Metoecus, in the cell of the wasp. The 
wasps, though they investigate the cells, do not apparently entertain 
any objection to the Metoecus, though there may be sometimes as 
many as twenty or thirty of the destroyers in a single nest. A 
few hours after the Metoecus has become a winged Insect and has 
escaped from the cells, it appears however, from the observations 
of Erne l on nests of wasps in captivity, that the wasps become 
hostile to the foreigners, and it is probable that in a state of 
nature these leave the nest as quickly as possible. Emenadia 
Jitil>dlata, a genus allied to Metoecus, has been discovered by 
Chobaut to have a similar life-history, except that it attacks a 
solitary wasp of the genus Odynerus? An old record to the 
effect that a second species of Emenadia, E. bimaculata, lives in 
the stalks of Eryngium campestre, on the pith, is now thought to 
be erroneous. Fabre has found the larvae and pupae of another 
Ehipiphorid in the cells of a bee, Halictus sexcinctus. 

The most remarkable of the Bhipiphorids, from the point of 
view of its habits, is certainly Symlius blattarum, which is now 
treated as the same as an Insect previously described by Thunberg 
from specimens found in amber and called Ripidius pectinicornis. 
This species is parasitic in cockroaches : the male and female are 
very different, the former being an active winged Insect, while 
the female is worm-like, differing but little from the larva, and 
never leaving the body of the cockroach. It is to be regretted 
that the life-history is not better known. The species has been 
found on board ship in vessels coining from India ; the male has 
been met with in several European countries, but the female is 
excessively rare. 

Fam. 75. Cantharidae or Meloidae (Blister-beetles, Oil -beetles'). 
Head with an abrupt neck ; elytra and sides of the abdomen witli - 
out any coadaptation ; each claw of the feet with a long ^y^"'"'/"/" 
r/o.sv/// Applied beneath it. This distinct family consists of 
Heteromera with soft integument, and is remarkable for the fact 
that many of its members contain a substance that when extracted 
1 Mitt. Schu-eiz. cut. Ges. iv. 1876, p. 556. - Ann. Soc. ent. France, Ix. 1891, p. 447. 



2/O COLEOPTERA CHAP. 

and applied to the human skin, possesses the power of raising 
blisters. The life-history is highly remarkable, the most complex 
forms of hyper-metamorphosis being exhibited. The species now 
known amount to about 1500; there can be no difficulty in 
recognising a member of the family by the above characters, 
except that in a very few cases each claw bears a projecting 
tooth, instead of an elongate appendage parallel with itself. The 
penultimate tarsal joint is scarcely ever broader than the pi'e- 
ceding ; the colour and style of markings are extremely varied. 
There are two very distinct sub - families, Cantharides and 
Meloides ; the former are winged Insects, and are frequently 
found on flowers or foliage. The Meloides are wingless, and 
consequently terrestrial ; they have a very short metasternum, 
so that the middle coxae touch the hind ; and they also have 
very peculiar wing-cases, one of the two overlapping the other 
at the base ; in a few Meloids the wing - cases are merely 
rudiments. 

The post-embryonic development of these Insects is amongst 
the most remarkable of modern entomological discoveries. The 
first steps were made by Newport in 185 1, 1 and the subject 
has since been greatly advanced by Fabre, Eiley, and others. 
.Is an example of these peculiar histories, we may cite Eiley's 
account- of Epicauta vittata (Fig. 140), a blister-beetle living 
at the expense of North American locusts of the genus Calop- 
/' /ins. The locust lays its eggs underground, in masses sur- 
rounded by an irregular capsule, and the Epicauta deposits its 
eggs in spots frequented by the locust, but not in special 
proximity to the eggs thereof. In a few days the eggs of 
the blister-beetle hatch, giving rise to little larvae uf the kind 
called triungulin (Fig. 140, A), because each leg is terminated by 
three tarsal spines or claws. In warm, sunny weather these 
triungulins become very active ; they run about on the surface 
of the ground exploring all its cracks, penetrating various spots 
and burrowing, till an egg-pod of the locust is met with : into 
Ih is the triungulin at once eats its way, and commences to devour 
an egg. Should two or more Iriungulins enter the same egg-pod, 
battles occur till only one is left. After a few days passed in 

"On the Natural History, Anatomy, and Development of the Oil-Beetle, 
Meloe, ' Tr. LI mi. ,S'oe. xx. 1851, p. 297 ; ami xxi. 18.13, p. 167. 
Rep. U.S. ent. Commission, i. L878, p. 297. 



HETEROMERA CANTIIARIDAE 



271 



devouring a couple of eggs, the triungulin sheds its skin and 
appears as a different larva (Fig. 140, B), with soft skin, short 
legs, small eyes, and different form and proportions ; a second 
moult takes place after about a week, Lut is not accompanied by 
any very great change of form, thong] i the larva is now curved, 
less active, and in form like a larva of Scarabaeidae ; when 
another moult occurs the fourth instar appears as a still more 
helpless form of larva (Fig. 140, D), which increases rapidly 
in size, and when full grown leaves the remains of the egg-pod 




FIG. 140. Hypermetamorphosis of Epicauta vittata. North America. (After Riley.) 
A, Young larva or triungulin ; B, Caraboid iustar or second larva ; C, coarctate 
larva, or instar between the Scarabaeoid and Scolytoid larva ; D, Scarabaeoid larva, 
from which the Scolytoid, or sixth, instar differs but little ; E, pupa ; F, imago. 

it has been living on, and forms a small cavity near by ; here it 
lies on one side motionless, but gradually contracting, till the 
skin separates and is pushed down to the end of the body, dis- 
closing a completely helpless creature that has been variously 
called a semi-pupa, pseudo-pupa, or coarctate larva (Fig. 140, C) ; 
in this state the winter is passed. In spring the skin of the 
coarctate larva bursts, and there crawls out of it a sixth instar 
which resembles the fourth (Fig. 140, D), except in the somewhat 
n-durrd si/c and greater whiteness. It is worthy of remark that 
the skin it has deserted retains its original form almost intact. 
In this sixth instar the larva is rather active and burrows about. 



272 COLEOPTERA CHAP. 

but does not take food, and in the course of a few days again moults 
and discloses the true pupa (Fig. 140, E). As usual in Coleoptera 
this instar lasts but a short time, and in five or six days the perfect 
beetle appears (Fig. 140, F). It is extremely difficult to frame any 
explanation of this complex development ; there are, it will be 
noticed, no less than five stages interposed between the first 
larval instar and the pupal instar, and the creature assumes in 
the penultimate one a quasi-pupal state, to again quit it for 
a return to a previous state. It is possible to look on the 
triungulin and the pupal instars as special adaptations to external 
conditions ; but it is not possible to account for the intermediate 
instars in this way, and we must look on them as necessitated by 
the physiological processes going on internally. Nothing, how- 
ever, is known as to these. It may be well to mention that, 
after describing and figuring (lac. cit.} this series of instars, Eiley 
changed his views as to their nomenclature. 1 The following 
summary of the metamorphosis, to which we have added the two 
nomenclatures of Eiley the original one, when different from the 
amended one, being given in square brackets may therefore be 
useful, viz. Egg ; 1, triungulin-larva moult ; 2, Caraboid larva 
[second larva, Caraboid stage] moult ; 3, Scarabaeoid larva 
[second larva, Scarabaeoid stage] moult; 4, Scarabaeoid laiva 
[second larva, ultimate stage] (large amount of food and much 
growth) moult; 5, coarctate larva [pseudo-pupa, or semi pupa] ; 
6, Scolytoid larva [third larva] (active, but little or no food taken) 

moult ; 7, pupa moult ; 8, perfect Insect. 

M. Fabre has succeeded in elucidating the history of Sitaris 
Immeralis, a Cantharid that lives at the expense of bees of the 
genus Anthophora? The eggs of the Sitaris are deposited in 
the earth in close proximity to the entrances to the bees' nests, 
about August. They are very numerous, a single female pro- 
ducing, it is believed, upwards of 2000 eggs. In about a month 

- towards the end of September they hatch, producing a 
tiny triungulin of black colour ; the larvae do not, however, 
move away, but, without taking any food, hibernate in a heap, 
remaining in this state till the following April or May, when 
they become active. Although they are close to the abodes of 
the bees they do not enter them, but seek to attach themselves 

1 Amer. Nat. xvii. 1883, p. 790. 
For illustration of this metamorphosis, see Vol. Y. p. 159 of this work. 



CANTHARIDAE 2/3 



to any haiiy object that may come near them, and thus a certain 
number of them get on to the bodies of the AntTio-phora and are 
carried to its nest. They attach themselves with equal readiness 
to any other hairy Insect, and it is probable that very large 
numbers perish in consequence of attaching themselves to the 
wrong Insects. The bee in question is a species that nests in 
the ground and forms cells, in each of which it places honey and 
lays an egg, finally closing the receptacle. It is worthy of 
remark that in the case of the Anthophora observed by M. 
Fabre, the male appears about a month before the female, and it 
is probable that the vast majority of the predatory larvae attach 
themselves to the male, but afterwards seize a favourable 
opportunity, transfer themselves to the female, and so get 
carried to the cells of the bee. When she deposits an egg on 
the honey, the triungulin glides from the body of the bee on to 
the egg, and remains perched thereon as on a raft, floating on 
the honey, and is then shut in by the bee closing the cell. This 
remarkable act of slipping on to the egg cannot be actually 
witnessed, but the experiments and observations of the French 
naturalist leave little room for doubt as to the matter really 
happening in the way described. The egg of the bee forms the 
first nutriment of the tiny triungulin, which spends about eight 
days in consuming its contents ; never quitting it, because contact 
with the surrounding honey is death to the little creature, which 
is entirely unfitted for living thereon. After this the triungulin 
undergoes a moult and appears as a very different creature, being 
now a sort of vesicle with the spiracles placed near the upper part ;. 
so that it is admirably fitted for floating on the honey (Vol. V. 
Fig. 86, 10). In about forty days, that is, towards the middle 
of July, the honey is consumed, and the vesicular larva after a 
few days of repose changes to a pseudo-pupa (11 of the fig. 
cited) within the larval skin. After remaining in this state for 
about a month, some of the specimens go through the subsequent 
changes, and appear as perfect Insects in August or September. 
The majority delay this subsequent metamorphosis till the follow- 
ing spring, wintering as pseudo-pupae and continuing the series 
of changes in June of the following year ; at that time the pseudo- 
pupa returns to a larval form (12 of the fig. cited), differing com- 
paratively little from the second instar. The skin, though detached, 
is again not shed, so that this ultimate larva is enclosed in two 
vor, vi T 



2/4 COLEOPTERA CHAP. 

dead skins ; in this curious envelope it turns round, and in a couple 
of days, having thus reversed its position, becomes lethargic and 
changes to the true pupa, and in about a month subsequent to 
this appears as a perfect Insect, at about the same time of the 
year as it would have done had only one year, instead of two, 
been occupied by its metamorphosis. M. Fabre employs the 
term, third larva, for the instar designated by Riley Scolytoid 
larva, but this is clearly an inconvenient mode of naming the 
instar. Sitaris humeralis is now very rare in Britain, but it 
seems formerly to have been more common, and it is not 
improbable that its triungulin may have been the " Pediculus 
meliUae," that was believed by Kirby to be a sort of bee-louse. 
Some species of the genus Mdoe are still common in Britain, and 
the Insects maybe seen with heavy distended abdomen grazing on 
herbage in the spring. The females are enormously prolific, a single 
one producing, it is believed, about 10,000 eggs. Meloe is also 
dependent on Antlwpliora, and its life-history seems 011 the whole 
to be similar to that of Sitaris ; the eggs are, however, not 
necessarily deposited in the neighbourhood of the bees' nests, 
and the triungulins distribute themselves on all sorts of un- 
suitable Insects, so that it is possible that not more than one in 
a thousand succeeds in getting access to the Antliopliora nest. It 
would be supposed that it would be a much better course for these 
bee-frequenting triungulins to act like those of E^ncauta, and hunt 
for the prey they are to live on ; but it must be remembered that 
they cannot live on honey ; the one tiny egg is their object, and 
this apparently can only be reached by the method indicated by 
Fabre. The history of these Insects certainly forms a most 
remarkably instructive chapter in the department of animal 
instinct, and it is a matter for surprise that it should not yet 
have attracted the attention of comparative psychologists. The 
series of actions, to be performed once and once only in a 
lifetime by an uniiistructed, inexperienced atom, is such that we 
should a 2 ]r 'iori have denounced it as an impossible means of 
existence, were it not shown that it is constantly successful. It 
is no wonder that the female Jfeloe produces 5000 times more 
eggs than are necessary to continue the species without diminu- 
tion in the number of its individuals, for the first and most 
important act in the complex series of this life - history is 
accomplished by an extremely indiscriminating instinct; the 



HETEROMERA 275 



newly hatched 3Moe has to get on to the body of the female of 
one species of bee ; but it has no discrimination whatever of the 
kind of object it requires, and as a matter of fact, passes with 
surprising rapidity on to any hairy object that touches it ; hence 
an enormous majority of the young are wasted by getting on to 
all sorts of other Insects ; these larvae have been found in 
numbers on hairy Coleoptera as well as on flies and bees of wrong 
kinds ; the writer has ascertained by experiment that a camel's- 
hair brush is as eagerly seized, and passed on to, by the young 
Mdoe as a living Insect is. 

The histories of several other Cantharids have been more or 
less completely discovered. Fabre has found the larva of 
Germ-milt/, schaefferi attacking the stores of provisions laid up by 
a fossorial wasp of the genus Tacliytes, and consisting of 
Orthoptera of the family Mantidae. The student who wishes 
for further information may refer to M. Beauregard's work on 
this family. 1 

Some half-dozen species of the genus Ceplialoon found in 
Siberia, Japan, and North America, have, by some authorities, 
been separated as the family Cephaloidae. Nothing is known 
as to the metamorphosis of these rare beetles ; and at present it 
is not necessary to distinguish them from Cantharidse. 

Fam. 76. Trictenotomidae. Large Heteromera, with powerful 
free projecting mandibles ; the antennae long, but with the termi mil 
three joints short, with angular projections on one side. This 
family includes only two genera and seven or eight species. 
They are very remarkable Insects ; Autocrates aenea being three 
inches long. The family is of considerable interest, as it seems to 
have no affinity with any other Coleoptera. The appearance of 
the species somewhat reminds one of Lucanidae, or Prionides ; 
but Trictenotomidae have even less relation to those beetles th;in 
they have to the members of the Heteromerous series. The 
Trictenotomidae appear to be found only in the primitive forests 
of the Indian and Indo-Malayan regions. Nothing is known as 
to their life-histories. 

1 Les Inscctcs Vesicants, Paris 1890, 554 pp. Parts of this work were pre- 
viously published in J. de I'Anat. Phys., xxi. xxii. xxiii. 1886 and 1SS7. 



2/6 COLEOPTERA 



CHAP. 



Series V. Phytophaga. 

Tarsi apparently four-jointed, the three basal joints usually densely 
set with cushion -like pubescence beneath; the third joint 
different in form, being divided into two lobes, or grooved 
on its upper surface so as to allow of tit e fourth joint being 
inserted near its base instead of at its extremity. Head not 
forming a definite prolonged beak ; its labrum visible, the 
palpi rarely (and even then not con^letcly} occluded in the 
mouth. 

This great series of beetles includes something like 35,000 
species. It approaches, like all the other series, the Polymorpha, 
especially the family Erotylidae placed therein, Imt in the great 
majority of cases there is no difficulty in recognising its 
members. The tarsi have never the Heteromerous formula, the 
head is not constructed like that of Ehyuchophora, nor the 
mouth and feet like those of Adephaga ; the antennae are 
different from those of the Lamellicorns. The tarsi are really 
five-jointed, for careful inspection shows that the long claw-joint 
has at its extreme base a small nodule, which is undoubtedly 
the fourth joint (Fig. 142, B). In speaking of the joints it is, 
however, customary not to refer to this small and functionally 
useless joint at all, and to call the claw-joint the fourth ; when 
the little joint is referred to it may be called the true fourth 
joint, 

Nearly the whole of the enormous number of species of this 
series are directly dependent on the vegetable kingdom for 
their nutriment ; they are therefore well styled Phytophaga. 
This term is, however, restricted by some systematists to the 
family we have called Chrysomelidae. Although there is 
enormous variety in this series, three families only can be at all 
naturally distinguished, and this with difficulty. Of these the 
Bruchidae are seed -feeders, the Chrysomelidae, as a rule, leaf- 
feeders, the Cerambycidae wood and stem-feeders. The number of 
exceptions to this rule is but small, though certain Cerambycidae 
and certain Chrysomelidae live on roots. 

Fam. 77. Bruchidae. Prosternum extremely short ; in front 
perpendicular; behind the coxae, forming merely a transverse 
lamina with pointed extremity. Hind femora more or less 



PHYTOPHAGA BRUCHIDAE 



2/7 



thickened. This comparatively small family includes about 
70 species of small, unattractive beetles. The larvae live in 
seeds ; hence some of the species are liable to be transported by 
means of commerce ; some of them do considerable injury ; peas 
and beans being specially subject to their attacks. They are 
able to complete their growth with a very small amount of 
nutriment, some of them consuming only a portion a little larger 
than themselves of a bean or pea. The larvae are fat maggots 
without legs, but Riley has discovered that the young larvae of 
Bruclius pisi and B. fabae have, when first hatched, three pairs 
of legs which are subsequently lost. They also have peculiar 



FIG. \\.Bn>clivs 
pisi or pea-weevil. 
A, Young larva ; 
B.protlioracie spin- 
ous process ; C, 
post-embr yonic 
leg, greatly magni- 
fied. ; D, pea-pod, 
with tracks of 
entry ; E, portion 
of pod, with egg, 
and the ' subse- 
quently formed 
track, magnified ; 
F, imago. (After 
Riley.) 



spinous processes on the pronotum. Both of these characteristics 
may be correlative with the transient differences in the activities 
of the larva, for the little creature is not at first located in the 
pea, but mines a gallery in the pod, in which it moves about, 
subsequently entering the pea and losing its legs. There is a 
good deal of difference in these respects between the two species 
B. pisi and B. fabae examined by Riley, and as but little is 
known of the life-histories of other Bruchidae it is probable that 
still greater variety prevails. Heeger has found that Bruclius 
lentis sometimes requires two seeds to enable it to complete its 
growth ; it is, notwithstanding its legless state when half-grown, 
able to migrate by dropping to the earth, and dragging itself 
along by its mandibles till it conies to another pod into which it 
bites its way. 

The family has, until recently, been placed in the Rhyncho- 




2/0 



COLEOPTERA 



CHAP. 




phorous series, with which it has, however, no direct connection. 
On the other hand, it is so closely connected with Chrysomelidae 
that it is not possible to indicate good characters to distinguish 
the two at present. The Australian genus Carpophagus, and the 
large South American species of Garydborus appear to be quite 
indistinguishable as families, though Lacordaire and Chapuis 
placed one in Bruchidae, the other in Chrysomelidae. The 
definition we have given applies, therefore, to the majority of the 
family, but not to the aberrant forms just mentioned. The 
European genus Urodon appears to belong to Anthribidae, not to 
Bruchidae. The family Bruchidae is called Mylabridae by some. 
Fam. 78. Chrysomelidae. Antennae moderately long ; eyes 
moderately I// rt/r, usu filly not at all surrounding the insertion of tlir 
antennae; upper surface usually bare, frequently brightly coloured 
a ml shining. This enormous family comprises about 18,000 
species of beetles, in which the form and details of structure 
are very varied. No satisfactory character for distinguishing 
5 Chrysomelidae from Cerambycidae has 

^ e ^ ^ een di sco vere d, although the two 
families are certainly distinct and 
natural. Most of the Chrysomelidae 
live on foliage ; few of them are more 
than half an inch long, whereas the 
Cerambycidae are wood - feeders and 
usually of more elongate form and larger 
size. The potato beetle, or Colorado beetle, 
that occasioned so much destruction in 
North America some thirty years ago,and 
the introduction of which into Europe 
was anticipated with much dread, is a 
good example of the Chrysomelidae. The 
turnip flea, a tiny hopping beetle, is 

FIG. [4-2. Doryphoru decent- J 

i in fit ta, the potato beetle, among the smallest forms of the family. 

rSndTi. \ In S a d is mem ber of another very exten- 
joint ; !, true fourth joint ; sive subdivision of Chrysomelidae, viz. 

[>, so-called fourth joint. TT it- i TU -ri i 

Halticides. The term Phytophaga is 

by many naturalists limited to Chrysomelidae, the Cerambycidae 
being excluded. The classification of the family is but little 
advanced, but the enormous number of species of Chrysomelidae 
are placed in four divisions, viz. : 



v PHYTOPHAGA CHRYSOMELIDAE . 279 

Prothorax much narrower at the base than the elytra, and usually with- 
out side-margins (raised edges). Sub-fain. 1. EUPODA ; with three divisions, 
Sagrides, Donaciides, Criocerides. 

The basal ventral plates of the abdominal segments are somewhat shorter 
in tlie middle than at the sides, the fourth one being often invisible in the 
middle, while the fifth is very large. Sub-fam. 2. CAMPTOSOJIES ; with six 
divisions, Megascelides, Megalopides, Clythrides,.Cryptocephalides, Chlamydes, 
Sphaerocarides. 

In the other two groups there is no great disparity between the fourth 
and fifth ventral plates. 

Prothorax not greatly narrower at the base than the elytra, and usually 
with distinct edges at the outsides. Sub-fam. 3. CYCLICA ; with four divi- 
sions, Lainprosomides, Eumolpides, Chrysomelides, Galerucides. 

Front of the head bent downwards or inflexed, so that the mouth is on 
the lower aspect. Antennae inserted close together on the most anterior 
I'-ul of the head, so that they are more forward than the mouth. Sub-fam. 
4. CRYPTOSTOMES ; with two divisions Hispides, Cassidides. 

In the other three divisions the mouth is placed as usual, but the insertion 
<pf the antennae varies a good deal. 

The larvae of about 100 species of the family are known: 
they are arranged in accordance with their habits, by Chapuis, 1 
in six groups, viz. : 

1. Elongate larvae, living under water, and there undergoing their meta- 
morphosis. (Donaciides.) 

2. Larvae mining in leaves, and undergoing their metamorphosis in the 
leaf. (Hispides and some Halticide-. 

3. Short convex larvae, frequently with leathery and pigmented integuments, 

living exposed on plants. (Most of the Cyclica.) 

4. Larvae of short form ; covering the body with excrementitious matter. 

(Some Criocerides.) 
o. Peculiar larvae of short form, spiny, and protecting their bodies by 

excrementitious matter attached by a special apparatus, the excrement 

itself being modified so as to be suitable for retention. (Cassidides.) 
6. Elongate, pallid, -larvae with curved abdomen ; living in shell-like case.-, 

and undergoing metamorphosis therein. (Most of the Camptosomes, 

the habits of which are known.) 

Though our knowledge of these larvae extends to only about 
100 out of 18,000 species, the above category by no means 
includes all the kinds of larvae ; Captain Xambeu having 
recently discovered that, the larva of Chrysochus pretiosus HVQS in 
the earth feeding on roots after the manner of a Rhizotroyn* 
larva, which it resembles. The larva uf Xnym splcndida lives 



Genera dcs CoUopteres (Suites 7>'////'n/(). x. Paris. 1874, p. 15. 



280 COLEOPTERA 



CHAP. 



inside the stems of Dioscorea batatas, in swellings ; the group 
Sagrides, to which it belongs, is a very anomalous one. 

i. EUPODA. The beetles of the genus Donacia are of special 
interest. They form, with the genus Haemonia, a peculiar group, 
w r ell represented in Europe, and also in our own country. They are 
all connected with aquatic plants, the species of Haemonia living 
entirely under water, while the Donacia, live in the imago-state 
an aerial life ; though many of them enter the water with great 
readiness, and, it is said, are able to take wing from the surface. 
The larvae live 011 the roots of aquatic plants, and derive not only 
nutriment but air therefrom ; they pass several months as pupae 
(or as resting larvae waiting for pupation), under water in cocoons 
which they construct, and which, incredible as it may seem, are 
filled with air, not water. Exact details as to the construction 
of these cocoons are wanting. It was formerly absurdly supposed 
that the larva swelled itself out to the size of the cocoon it was 
about to make, and so served as a mould, subsequently con- 
tracting. The observations of Schmidt-Schwedt l make it, however, 
more probable that the plant itself furnishes the air which, under 
pressure of the water (so he supposes), fills the cocoon ; the larva 
wounds the root, piercing to an air-vessel and then constructs 
the cocoon on this spot, leaving to the last moment an orifice, 
according to Schmidt, as an exit for the water. The larva uses 
a similar artifice for obtaining air; it has no gills, but is pro- 
vided near the extremity of the body with two sharp chitinous 
processes which it drives into the root of the plant till it 
penetrates an air-vessel. Schmidt thinks the processes serve as 
conduits to conduct the air to the tracheae, but Dewitz thinks 
the air enters the larva in a more normal manner, by means of a 
stigma placed at the base of the piercing process. A similar 
larva exists in Haemonia ; which genus is additionally interesting 
from the fact that the imago lives entirely submerged. It is 
not known how it breathes. This genus is the only member of 
the Chrysomelidae that does not possess the structure of the 
feet that is characteristic of the Phytophaga. The late Professor 
Babington about sixty years ago found H. curtisi at Cley on 
the Norfolk coast on submerged Potamoycton pectinatus, but it has 
not been met with there for a great many years. 

The larvae of Criocerides are of two kinds, in one of which the 

1 JJ.Tliti. ent. Zdt. 1887, i>. o25, and 1889, p. 299. 



PHYTOPHAGA CHRYSOMELIDAE 



281 




body is peculiarly shaped in conformity with the curious habit 
of using the excrement as a covering. The larva is less elon- 
gate than usual, and has the anus placed on the upper surface, 
and formed so that the excrement when voided is pushed forward 
on to the Insect ; here it is retained by means of a slimy matter, 
and a thick coat entirely covering the creature, is ultimately 
formed. The larva of Lcma melanopa is not uncommon about 
Cambridge, where it feeds on the leaves of growing corn. It is 
a remarkable fact that even in one genus the species have some 
of them this habit, but others not. The species of Crioceris 
living on lilies C. merdiyera, 
e.g. are noted for possessing it ; 
while C. asparagi does not pro- 
tect itself in this way, but emits 
fluid from its mouth when dis- 
turbed. This larva is a serious 
nuisance in some localities to the 
cultivators of asparagus. The 
eggs are deposited on the stems 
of the plant as shown in our 
figure sometimes in great num- 
bers. 

The perfect Insects of many 
of the Criocerides possess a stridu- 
latiug organ. Two contiguous 
areas at the base of the last 
dorsal segment, where they can 
be rubbed by the tips of the 
elytra, are slightly elevated and 
bear very close and tine straight 
lines. 

ii. The CAMPTOSOMES, as we have already noticed, are distin- 
guished by a peculiar structure of the abdomen. This character 
appears to be connected with a very remarkable habit, viz. the 
formation of a case to envelop the egg. The tip of the abdomen 
is somewhat curved downwards, and, in the female, bears a 
hollow near the extremity ; when an egg is extruded the female 
holds it in this hollow by means of the hind legs, and 
envelops it with a covering said to be excrementitious. When 
the larva hatches, it remains within this case, and subse- 




FIG. 143. Crioceris asparagi. A, Eggs 
in position on stem of asparagus ; B, 
one egg much enlarged ; C, young 
larva. 



Cambridge. 



282 COLEOPTERA 



CHAP. 



quently enlarges it by additions from its own body. The 
beautiful Insects of the genus Cryptocephalus, which is fairly 
well represented in Britain, belong to this division. The 
exotic group Megalopodes is incorrectly placed in Camptosomes ; 
the side pieces of the prothorax meet in it behind the middle 
coxae, as they do in Khynchophora. The species of Megalopodes 
stridulate by means of an area on the base of the meso-scutellum 
rubbed by a ridge inside the pronotum, as in the Cerambycida?. 

iii. The division CYCLICA includes the great majority of Chryo- 
melidae ; we have not less than 170 species in Britain. The 
larvae live, like those of Lepidoptera, at the expense of foliage, 
and the species frequently multiply to such an extent as to be 
injurious. Some of them are destroyed in great numbers by 
Hymenopterous parasites, the Braconid genus Perilitus being one 
of the best known of these ; in some cases the parasite deposits 
its eggs in either the larva or perfect Insect of the beetle, and 
the metamorphoses of the parasites in the latter case are some- 
times, if not usually, completed, the larvae emerging from the 
living beetles for pupation. 

iv. The CRYPTOSTOMES, though comparatively few in number 
of species, include some very remarkable beetles. There are two 
groups, Hispides and Cassidides. The former are almost peculiar 
to the tropics and are not represented by any species in the 
British fauna. The head in this group is not concealed ; but in 
the Cassidides the margins of the upper surface are more or less 
expanded, so that the head is usually completely hidden by the 
expansion of the pronotum. Both the groups are characterised 
by the antennae being inserted very near together, and by the 
short claw-joint of the feet. Hi spa is one of the most extensive 
of the numerous genera of Hispides, and is remarkable from the 
imago being covered on the surface with long, sharp spines. But 
little is known as to the metamorphosis, beyond the fact already 
alluded to, that the larvae of several species mine the interior of 
leaves. The larva of Hispa testacea, according to Ferris, 1 makes 
use of the leaves of Cistus salvifolius in Southern Europe ; it is 
broad and flat, and possessed of six short legs. The eggs are not 
deposited by the parents inside the leaves, but are probably 
attached to various parts of the plant. After hatching, the young 
larva enters a leaf, and feeds on the parenchyma without rupturing 

1 Ann. Soc. Lie'/jc, x. 1855, p. 260. 



PHYTOPIiAGA CHRVSOMELIDAE 



283 



the epidermis ; but when it has consumed about three-fourths of 
the soft interior of the leaf it ruptures the epidermis of the upper 
surface, and seeks another leaf; this found, it places itself on the 
midrib, tears the upper epidermis, and lodges itself in the leaf. 
In the case of this second leaf it attacks the parenchyma in the 
neighbourhood of the petiole, and so forms an irregular tube 
which has an open mouth, the point of entry. In this tube it 
undergoes its metamorphosis. Each larva, it is said, alw;iy- 
makes use of two leaves, and of two opposed leaves. A know- 
ledge of the habits of some of the larger of the exotic Hispicles 
would be of much interest. 




FIG. 144. Pupa of 
C ass id id beetle 
(? AspidomorjiJi" 
s p.). A, With 
appendage ex- 
tended ; B, with 
the appendage re- 
posing on the back. 
New Britain. 



The Cassidides, in addition to the curious marginal expansion 
of their upper surface, have the power of withdrawing the head 
into the thorax, and hence they are often called shield or tortoise- 
beetles. They exhibit considerable variety in form and colour, 
and some of them display a peculiar metallic reflection of great 
delicacy and beauty ; this disappears entirely after death, but it 
may be restored by thoroughly moistening the dead Insect. The 
colour, therefore, probably depends on the presence of water in 
the integument. The larvae of Cassidides are notorious on 
account of their habit of covering their bodies with dried 
excrement, for which purpose they are provided with a forked 



284 COLEOPTERA CHAP. 

process at the posterior extremity; this serves to place the pro- 
tecting matter in a proper position and to retain it there. The 
excrement assumes in various species forms so peculiar that they 
cannot be considered merely incidental. In several species this 
covering-matter is like lichen. This is the case with Dolichotoma 
jxil iinn-tnii, the larva of which has, in place of the usual fork, a 
more complex appendage on the hack for the purpose of prepar- 
ing and retaining its peculiar costume. The pupae, too, some- 
times retain the larval skin. An extremely remarkable pupa of 
a Cassidid possibly of the genus Aspidomorplia was recently 
found by Dr. Arthur Willey in New Britain (Fig. 144). The 
back of the pupa is covered with a complex appendage, so that 
the creature has no resemblance to an Insect ; this appendage is 
perhaps capable of being moved, or even extended (Fig. 144, A), 
during life. Whether it may be formed by the retention of 
portions of the moulted gkins of the larva we cannot say with 
certainty. 

The most remarkable of the Cassidid coverings yet discovered 
are those formed by certain small beetles of the tropical Ameri- 
can genus Porpliyraspis. P. trtstts is apparently a common 
Insect at Bahia, where it lives on a cocoa-palm. The larva is 
short and broad, and completely covers itself with a very dense 
coat of fibres, each many times the length of the body, and 
elaborately curved so as to form a round nest under which the 
larva lives. On examination it is found that these long threads 
arc all attached to the anal extremity of the Insect, and there 
seems no alternative to believing that each thread is formed by 
small pieces of fibre that have passed through the alimentary canal, 
and are subsequently stuck together, end to end. The process of 

forming these long fibres, each one from 
scores of pieces of excrement, and giving 
them the appropriate curve, is truly remark- 
able. The fibres nearest to the body of the 
larva are abruptly curled so as to fit exactly, 
FIG. 1-45. Nest of intes- and make an even surface ; but the outside 

finally-made filaments fi j fc d fc j somewhat busllV 

under winch the larva ' 

of Poi-j>Jii/ri(yj>i.-i trintis fashion. The construction is much like that 
of a tiny bird's nest. Senor Lacerda informed 

the writer that the larva makes a nest as soon as it is hatched. 
Another 2 > <>fpJn/fitsj>is--P. palmarum- -lasts been recorded as 




PHYTOPHAGA CERAMBYCIDAE 



285 



forming similar nests on a species of Thrinax in St. Domingo. 
Candeze says l that when it lias completed its growth the larva 
ejects on to the leaf a quantity of semi-liquid matter, and this, on 
drying, sticks the nest to the leaf, so that the metamorphosis is 
effected under shelter. 

Fam. 79. Cerambycidae (Longicorns). Form usually oblong, 
not much, curved in outline at the sides; surface very frequent! '// ren- 
dered dull by a very minute hairiness, which often forms a ////*/// ; 
antennae usually long, and their insertion much embraced by ///*' 
eyes. This great family of beetles includes some 12,000 or 
13,000 known species. The elegance and variety of their forms 
and the charm of their colours have caused them to attract much 
attention, so that it is probable that a larger proportion of the 
existing species have been obtained than is the case in any other 
of the great families of Coleoptera. Still 
it is not likely that one-half of the living- 
forms are known. It is not possible at 
present to point out any one character of 
importance to distinguish Cerambycidae 
from Chrysomelidae, though the members 
of the two families have, as a rule, but 
little resemblance in external appearance. 
Most of them live on, or in, wood, though 
many are nourished in the stems of her- 
baceous plants. The larvae live a life of con- 
cealment, and are soft, w^hitish grubs with 

powerful mandibles, and usually with a comparatively small head, 
which is not much exserted from the thorax. Most of them are 
without legs, but a good many have three pairs of small legs, and 
there are numerous cases in which the surface of the body is 
furnished above or below with swellings believed to act as 
pseudopods (Fig. 84), and help the larvae to move about in their 
galleries ; but this is probably not the sole function of these 
organs, as their surface is varied in character, and often not of a 
kind that appears specially adapted to assist in locomotion. 
There is a slight general resemblance between the larvae of Ceram- 

O O 

bycidae and those of Buprestidae, and when the thorax of a 
Lougicorn larva is unusually broad, e.g. Asiynomus, this similarity 
is very pronounced. 

1 Mem. Soc. LUge, xvi. 1861, p. 387. 




1H! 



286 COLEOPTERA 



CHAT. 



The modes of life of Cerambycid larvae exhibit considerable 
variety, and much perfection of instinct is displayed by the larvae, 
as well as by the mother beetles. The larvae of Saperda popidnea 
are common in certain woods in the South of England in the 
stems of aspen ; they consume only a small quantity of the 
interior of the stem, and are probably nourished by an afflux of 
sap to the spot where they are situated. Elapliidion villosum is 
called the oak-primer in Xorth America. The parent beetle lays 
an eiro' near the axilla of a leaf-stalk or small stem, and the 

oo 

young larva enters this and feeds on the tender material ; as it 
grows it enters a larger limb, and makes an incision within this 
in such a manner that the wood falls to the ground with the 
larva within it, the dead wood serving subsequently as pa-bulum 
and as a shelter, within which the metamorphosis is completed. 
The species of the American genus Oncideres are called girdlers, 
because the parent beetle, after laying an egg in a small branch, 
girdles this round with a deep incision, so that the portion 
containing the larva sooner or later falls to the ground. The 

O O 

growth of a Longicorn larva frequently takes more than a year, 
and under certain circumstances it may be enormously prolonged. 
Monohammus confusus has been known to issue from wooden 
furniture in a dwelling-house when the furniture was fifteen 
years old. Individuals of another Longicorn have issued from 
the wood of a table, twenty and even twenty-eight years after the 
felling of the tree from which the furniture was made. Sereno 
Watson has related a case from which it appears probable that 
the life of a Longicorn beetle extended over at least forty-five 
years. 1 It is generally assumed that the prolongation of life in 
these cases is due to the beetle resting quiescent for long after it 
has completed the metamorphosis. Recent knowledge, however, 
renders it more probable that it is the larval life that is pro- 
longed ; the larva continuing to feed, but gaining little or no 
nutriment from the dry wood in these unnatural conditions. Mr. 
C. 0. TVaterhouse had for some years a Longicorn larva under 
observation, feeding in this way in the wood of a boot-tree ; ~ the 
burrows in the wood contained a great deal of minute dust 
indicating that the larva passed much matter through the ali- 
mentary canal, probably with little result in the way of nutriment. 

1 Packard, 5th Rep. U.S. Ent. Comm. 1S90, p. 689. 
- Xot a growing tree, but the instrument used for stretching boots. 



HYTOPHAGA CERAMBYCIDAE 287 

There are numerous Longicorns that bear a great resemblance 
in form and colour to Insects to which they are not related. 
Haensch J has noticed that species of the genus Odontocera 
resemble various Hymenoptera, one species being called 0. 
braconoides; he also observed that these Hymenoptera - like 
Longicorus, instead of withdrawing their underwings under the 
elytra as beetles generally dn. vibrate them rapidly like Hyn;en- 
"ptera. A large number of Loiigicnrn- ,-tridulate loudly by rubbing 
a ridge inside the pronotum on a highly specialised, striate >urf;i<-e 
at the base of the scutellum, and therefore covered up when the 
Insect is contracted in repose. A few produce noise by rubbing 
the hind femora against the edges of the elytra, somewhat after 
the fashion, of grasshoppers. In this case there appears to be 
comparatively little speciality of structure, the femora bearing, 
however, more or less distinct small granules. The species of the 
Hawaiian genus Plagithmysus produce sound in both these 
manners, the thoracic stridulating organ being beautifully de- 

o o o */ 

velupcd, while in some species the margin of the elytra and 
b:;se of the femora are also well adapted for the purpose of sound- 
production, and in a few species of the genus there are al.- 
highly-developed stridulating surfaces on the hind and middle 
coxae. This is the only case in which a beetle is known to 
possess more than one set of sound-organs in the imago state. 
Three divisions of this family are distinguished, viz. 

1. Front coxae large and transverse ; prothorax with distinct side mar- 
gins. Sub-fain. 1. PRIONIDES. 

2. Front coxae not greatly extended transversely, thorax not margined ; 
last joint of maxillary palpus not pointed, usually broader (more or less) 
than the preceding joint. Sub-fain. 2. CERAMBYCIDK-. 

3. Front coxae usually round and deeply embedded ; last joint of maxil- 
lary palpus pointed ; front tibiae with a more or less distinct, slanting groove 
"ii the inner side. Sub-fam. 3. LAMIIDES. 

The Prionides are on the average considerably larger in size 
than the members of the other divisions, and they include some of 
the largest of Insects. The Amazonian Titanus giganteus and the 
Fijian Mn<'r<itn,na heros are amongst the most gigantic. Some 
uf the Prionides have a great development of the mandibles in 
the male sex analogous to that we have already noticed in 
Lucanidae. The larvae of the large Prionides appear in various 
parts of the world to have been a favourite food with native 
1 Berlin, ent. Zeitsclo: xli. 1S96, SB. p. 22. 



288 COLEOPTERA 



CHAP. 



tribes, and Lumholz states that they are really good eating. 
In consequence of the destruction of forests that has progressed 
so largely of late years these gigantic Prionides have become 
much rarer. 

Several aberrant forms are included in Prionides. The genus 
Parandra has five-jointed tarsi; the third joint being much smaller 
than usual, so that the fourth joint is not concealed by it. 
The Brazilian Hypocephalus armatus was for long a subject of 
dispute as to its natural position, and was placed by different 
authorities in widely -separated families of Coleoptera. The 
structure of this aberrant Longicorn seems to be only explicable 
on the hypothesis of warfare amongst the males. 1 Nothing is, 
however, known as to the habits and history of the Insect, and 
only one or two specimens of the female have yet been obtained. 

The family Spondylidae has been proposed for some of 
these aberrant Longicorns, but as it includes but very few, and 
highly discrepant, species, it is neither natural nor of much use 
for systematic purposes. 

The Lamiides are the most highly specialised division of the 
Longicorns, and includes the larger number of the species. The 
front of the head is usually placed at right angles to the vertex, 
and in some cases (groups Hippopsini, Spalacopsini) it is strongly 
intiexed, so that the mouth is placed on the under side of the 
head. The extension of the eyes round the antennae is accom- 
panied by very curious shapes of those organs, and not 
infrequently each eye is divided into two more or less widely- 
separated parts, so that the Insect has, on the external surface, 
four eyes. 

Series VI. Rhynchophora. 

Head more or less prolonged in front to form a snout or leal', called 
rostrum. Tarsi four-jointed, usually nt least tlie f/iirtl 
joint Iroad and densely pubescent leneatJi. 

This enormous series includes about 25,000 species, and as 
may well be imagined shows a great variety of structure amongst 
its forms. The vast majority may, however, be readily recognised 
by the two characters mentioned above. There are some cases in 
which the beak is indistinct, and others in which the tarsi are 

1 Sharp, Ann. Soc. ent. Belyique, xxviii. 1384, CR. p. cvii. 



RHYNCHOPHORA 



289 



five-jointed (Dryophthorus), and even slender (Platypides). In 

these cases a close examination shows that the gular region on 

the middle of the back of the under surface of the head cannot 

he detected, and that the back of the prosternum is very strongly 

consolidated by the side-pieces of the thorax meeting together 

and being very firmly joined behind the coxae. The beak is in 

the great majority perfectly distinct, though it varies so extremely 

in form that it can only be briefly described by saying that it is 

a prolongation of the head in front of the eyes, or that the 

antennae are inserted on its sides near to, or far from, the tip. 

It has been ascertained in many cases that the rostrum is used 

by the female to assist in placing the eggs in suitable places, a 

hole being bored with it ; in some 

cases it is also used to push the 

egg far into the hole in which it 

has previously been placed by 

the ovipositor ; but there are 

many forms in which it is fairly 

certain that it is not so used. 

What purpose it serves in tbe 

male is totally unknown. In 

many members of the series, the 

rostrum differs in form in the 

two sexes, and in most, if not in 

all, these cases it is clear that the 

distinctions tend in the direction 

of making the beak of the female 

more efficient for the mechanical 

purpose we have mentioned. 

It was proposed by Leconte 
and Horn to separate this series 
from all the other Coleoptera as 
a primary division, and they 
looked on it as of lower or more 
imperfect structure. Packard has 
very properly protested against FIG. Itf.Evgnoristws mu-uachus ?. 

, -L , . Madagascar. A. The imago ; B, front 

this interpretation ; and there of pr0 notum, head, and rostrum, 
seems to be no reason whatever 

for considering the Khynchophora as " lower " than other beetles ; 
indeed we should be inclined to place such forms as Calandrides 
VOL. VI u 




290 COLEOPTERA CHAP. 

amongst the most perfect of Insects ; their external structure (as 
shown by Eugnoristus monachus, Fig. 147) being truly admirable. 
Only four families of Ehynchophora can lie at present 
accepted as satisfactory ; one of these Curculionidae includes 
an enormous majority of the whole series. Though it is probable 
that it will ultimately be divided into several families, the 
attempts to that end that have already been made are not 
satisfactory. 

Fam. 80. Anthribidae. Palpi usually not covered, Imt dis- 
tinct and flexible. Antennae often long, not elbowed, the first joint 
nt r< r;i lonij. Third joint of tarsus small, usually much concealed 
by being embraced by the second joint. Pi/gidii/ni exposed; pro- 
/if/i/idu/tn. deeply grooved in the midJfr. This family includes 800 
or more species, which are mostly tropical ; it is very sparsely 
represented in the faunas of Europe and North America. It is 
quite distinct from Curculionidae with which it was formerly 
associated. It contains many graceful Insects having a certain 
resemblance with Longicorns on account of the large development 

of the antennae. The habits and meta- 
morphoses are but little known. It 
seems probable that many species find 
their nutriment in old wood or boleti. 
The larvae of some genera (Cratoparis 
and Araeocerus) have legs, but in others 
the legs are wanting, and the larvae are 
said to completely resemble those of 
Curculionidae. In the larva of our 
tiny British species, Choragus sheppardi, 
the legs are replaced by three pairs 
of thoracic, sac-like pseudopods. This 
Insect makes burrows in dead branches 
FIG. i48.--P/r^/,v,/,,,|.< lati- of i iawt h m. The larvae of the genus 

/'.N//-/.V, Anthribidae. Britain. _ 

A, the perfect insect ; B, Bi'achytarsus have been ascertained to 

tarsus and tip o prey Qn Cocddae _ 

Fam. 81. Curculionidae ( Wecrils). The beak of rery variable 




and t/iir/,-/irss ; the palpi- small, nearly always concealed 
within Hie mouth, short, and rigid. Labr/n/i absent. Antennae 
of the majority r/bu/red, i.e. with the basal joint longer, and so 
formed that when 'it is faff rally extruded the other joints can be 
placed in a forward direction. This enormous family includes 



v RHYNCHOPHORA CURCULIONIDAE 291 

about 20,000 known species, and yet a large portion of the 
species yearly brought from the tropics still prove to be new. 
The rostrum or beak exhibits excessive variety in form, and is in 
many cases different in the sexes ; in this case it is usually longer 
and thinner in the female. As the rostrum is one of the chief 
characters by which a member of the family may be recognised, 
it is necessary to inform the student that in certain forms (the 
Australian Amycterid.es, e.g.') the organ in question may be so 
short and thick that it is almost absent. In these cases the 
Insect may be identified as a Curculionid by the gular area being 
absent on the under side of the head, and by the concealment of 
the palpi. The tarsi are usually of the same nature as those of 
Phytophaga, already described, but the true fourth joint is less 
visible. In the Brachycerides this joint is not present, and the 
third joint is not lobed. The palpi are flexible and more or 
less exserted in a very few species (Rhynchitides) ; in Rhinoma- 
cerides there is also present a minute labrum. The front coxae 
are deeply embedded, and in many forms the prosternum is 
peculiar in structure ; the side-pieces (epimera) meeting at the 
back of the prosternum in the middle line. This, however, is 
not universal in the family, and it occurs in some other beetles 
(e.g., Megalopodides of the Phytophaga). The larvae are without 
legs. They are vegetarian, the eggs being deposited by the 
mother-beetle in the midst of the food. These larvae may be 
distinguished from those of Longicorns by the general form, 
which is sub-cylindric or rather convex, not flattened, and 
more particularly by the free, exserted head, the mouth being 
directed downwards ; the attitude is generally a curve, and the 
anterior part of the body is a little the thicker. Xo part of 
plants is exempt from the attacks of the larvae of Curculionidae ; 
buds, twigs, leaves, flowers, fruits, bark, pith, roots and galls 
may each be the special food of some Curculionid. Certain 
species of the sub-families Rhynchitides and Attelabides prepare 
leaves in an elaborate manner to serve as food and dwelling for 
their young. If young birches, or birch bushes from 5 to 10 
feet in height, be looked at in the summer, one may often notice 
that some of the leaves are rolled so as to form, each one, a little 
funnel. This is the work of Ehynchites (or Deporaus) betulae, a 
little Curculionid beetle (Fig. 149). An inspection of one of 
these funnels will show that it is very skilfully constructed. The 



292 



COLEOPTERA 



CHAP. 



whole of a leaf is not used in the formation of a funnel, cuts being 
made across the leaf in suitable directions. The beetle stand- 
ing on a leaf, as shown in the figure, proceeds to cut with its 
mandibles an. incision shaped like an erect S, commencing at a 
certain part of the circumference, and ending at the midrib of the 
leaf; the beetle then goes to the other side of the midrib, and 
continues its incision so as to form another S-like curve con- 
siderably different from the first ; being prostrate and less abrupt. 
Thus the blade of the leaf is divided into two halves by certain 
curved incisions, the midrib remaining intact. The little funuel- 




FIG. 149. The leaf-rolling of Rh yncli itf* l^ti/hic. Britain. A, Female beetle, magnified : 
B, the beetle forming the first incision on a leaf ; C. the completed roll. (B and C 
after Debey. ) 

twister no\v commences to roll up the leaf to form the funnel ; 
and this part of the work is greatly facilitated by the shape of 
the incisions. Going back to the spot where it commenced work, 
by the aid of its legs it rolls one side of the leaf round an ideal 
axis, somewhat on the same plan as that adopted by a grocer in form- 
ing a paper-funnel for sugar. The incisions are found to be just 
of the right shape to make the overlaps in the rolling, and to re- 
tain them rolled-up with the least tendency to spring back. After 
some other operations destined to facilitate subsequent parts of its 
task, the beetle enters the rolled-up part of the leaf and brings it 
more perfectly together; it again comes out and, pursuing a 
different system, holds on with the legs of one side of the body 



v RHYNCHOPHORA CURCULIONIDAE 293 

to the roll, and with the other legs drags to it the portion of the 
leaf on the other side of the midrib so as to wrap this part (i.e. 
the result of its second incision) round the part of the funnel 
already constructed. This being done the Insect again enters 
the funnel, bites three or four small cavities on the inside of the 
leafy wall and deposits an egg in each. Afterwards it emerges 
and fits the overlaps together in a more perfect manner so as to 
somewhat contract the funnel and make it firmer ; then proceeding 
to the tip, this is operated on by another series of engineering 
processes and made to close the orifice ; this part of the opera- 
tion being analogous to the closing by the grocer of his paper- 
funnel after the sugar has been put in. The operation of the 
beetle is, however, much more complex, for it actually makes a 
sort of second small funnel of the tip of the leaf, bends this in, and 
retains it by tucking in some little projections. The work, which 
has probably lasted about an hour, being now completed, the creature 
takes a longer or shorter rest before commencing another funnel. 
We have given only a sketch of the chief points of the work, 
omitting reference to smaller artifices of the craft master ; but 
we may remark that the curved incisions made by the beetle 
have been examined by mathematicians and duly extolled as 
being conducted on highly satisfactory mathematical principles. 
It is impossible at present for us to form any conception as to the 
beetle's conceptions in carrying out this complex set of operations. 
Our perplexity is increased if we recollect its life-history, for we 
then see that neither precept or example can have initiated its 
proceedings, and that imitation is out of the question. The eggs 
hatch in their dark place, giving rise to an eyeless maggot, 
which ultimately leaves the funnel for the earth. The parts of 
this maggot subsequently undergo complete change to produce 
the motionless pupa of entirely different form, from which 
emerges the perfect Insect. Hence the beetle cannot be con- 
sidered to have ever seen a funnel, and certainly has never 
witnessed the construction of one, though, when disclosed, it 
almost immediately sets to work to make funnels on the complex 
and perfect system we have so imperfectly described. More 
general considerations only add to the perplexity we must feel 
when reflecting on this subject. Why does the Insect construct 
the funnel at all ? As a matter of protection it. appears to be 
of little use, for the larvae are known to suffer from the attacks 



294 COLEOPTERA CHAP. 

of parasites as other Insects do. We have not the least reason 
for supposing that this mode of life for a larva is, so far as 
utility is concerned, better than a more simple and usual one. 
Indeed, extraordinary as this may appear, it is well known that 
other species of the same genus adopt a simple mode of life, 
laying their eggs in young fruits or buds. We think it possible, 
however, that a knowledge of the mode of feeding of this larva 
may show that a more perfect nutrition is obtained from a well- 
constructed cylinder, and if so this would to a slight extent 
satisfy our longing for explanation, though throwing no light 
whatever on the physiology or psychology of the artificer, and 
leaving us hopelessly perplexed as to why a beetle in ages long 
gone by should or could adopt a mode of life that by long pro- 
cesses of evolution should, after enormous difficulties have been 
overcome, attain the perfection we admire. 1 

Fam. 82. Scolytidae. Rostrum extremely short, broad ; tibiae 
frequently denticulate externally; antennae, short, with a Iroad 
cluli. This family is not at all sharply distinguished from certain 
groups of Curculionidae (from Cossonid.es e.g.}, but as the species 
have somewhat different habits, and in the majority of cases 
can be readily distinguished, it is an advantage to separate 
the two families. About 1400 species are at present known. 
Most of them are wood- and bark-feeders; some bore into hard 
wood ; a few mine in twigs or small branches of trees, but the 
majority live in the inner layers of the bark ; and this also 
serves as the nidus of the larvae. A small number of species 
have been found to inhabit the stems of herbaceous plants, or to 
live in dry fruits. Owing to their retiring habits they are rarely 
seen except by those who seek .them in their abodes, when they 
may often be found in great profusion. The mother-beetle bores 
into the suitable layer of the bark, forming a sort of tunnel and 
depositing eggs therein. The young larvae start each one a 
tunnel of its own, diverging from the parent tunnel ; hence each 
batch of larvae produces a system of tunnels, starting from the 
parents' burrow, and in many species these burrows are charac- 

1 For a Tnore extensive account of Rhynchites bctuJae and others refer to 
Wasmann. Dcr Trickier wickler, Minister, 1884, and Debey, Beitrdgc zur Lebcus- 
und Entwickelungs-geschichte . . . <l?r AUcll>uJ?n, Bonn, 184(5. The first in- 
cludes an extensive philosophical discussion ; the second is a valuable collection 
of observation-. 



v KHYNCHOPHORA SCOLYTIDAE BRENTHIDAE 295 

teristic in form and direction, so that the work of particular 
Scolytids can be recognised by the initiated. 

The Platypides bore into the wood of trees and stumps ; they 
are chiefly exotic, and little is known about them. They are 
the most aberrant of all Rhynchophora, the head being remarkably 
short, flat in front, with the mouth placed on the under surface 
of the head, there being no trace of a rostrum : the tarsi are 
elongate and slender, the third joint not being at all lobed, while 
the true fourth joint is visible. Hence they have not the 
appearance of Rhynchophora. Some authorities treat the 
Platypides as a distinct family. 

Some of the members of the group Tomicides also bore into 
the wood. Recent observations have shown that there is an 
important feature in the economy of certain of these wood- 
borers, inasmuch as they live gregariously in the burrow, and 
feed on peculiar fungi that develop there, and are called ambrosia. 
According to Hubbard, 1 some species cultivate these fungi, 
making elaborate preparations to start their growth. The fungi, 
however, sometimes increase to such an extent as to seal up the 
burrows, and kill the Insects by suffocation. 

Scolytidae sometimes multiply to an enormous extent, attack- 
ing and destroying the trees in wooded regions. Much dis- 
cussion has taken place as to whether or not they are really 
injurious. It is contended by one set of partisans that they 
attack only timber that is in an unhealthy, dying, or dead con- 
dition. It may be admitted that this is usually the case ; yet 
when they occur in enormous numbers they may attack timber 
that is in a sort of neutral state of health, and so diminish its 
vigour, and finally cause its destruction. Hence it is of great 
importance that they should be watched by competent foresters. 

The larvae of Scolytidae are said to completely resemble those 
of Curculionidae : except in the group Platypides, where the body 
is straight and almost cylindrical, and terminates in an oblique 
truncation bearing a short hard spine. 2 

Fam. 83. Brenthidae. Form elongate; rostrum straight, 
ili/'cctly continuing the long axis of the body, often so thick as to 
form an elongate head ; antennae not elbou-ed. The Brenthidae 
form a family of about 800 species, remarkable for the excessive 

1 Bull. U.S. Dcp. Agric. ent. New series, No. 7, 1897. 
2 Penis, Ann. Sci. Nat. (2) xiv. 1840. p. 89, pi. iii. 



296 



COLEOPTERA 



length and sleuderness of some of its forms, and for the 



s 




extreme difference in the sexes that frequently exists. It 

well represented in the tropics 
only, and very little is known 
as to the natural history and 
development. These beetles are 
stated to lie wood-feeders, and 
no doubt this is correct in the 
case of the majority of the 
species ; but Mr. Lewis observed 
in Japan that Zemioses celtis and 
Cyphagogus segnipes are pre- 
d;iceous, and enter the burrows 

of wood-boring Insects to search 
fiG. 150. JSupsalis mmuta. Aorth 

America. A, Larva ; B. pupa ; c, for larvae as prey : they are 

female imago; D, head of male. (After veiy much modified ill Structure 

to permit this ; and as the other 

members of the group Taphroderides are similar in structure, 
it is probable that they are all predaceous. Nothing what- 
ever is known as to the larval history of these carnivorous 
forms. Indeed an uncertainty, almost complete, prevails as to 
the early stages of this family. Eiley has given a sketch of a 
larva which he had no doubt was that of .#///> W/.s minuta, the 
North American representative of the family ; if he is correct 
the larva differs from those of Curculionidae by its elongate form, 
and by the possession- of thoracic legs : these, though small, are 
three-jointed. Descriptions, supposed to be those of Brenthid 
larvae, were formerly published by Harris and Motschoulsky ; 
but it is now clear that both were mistaken. 

In the higher forms of Brenthidae the rostrum of the female 
is perfectly cylindrical and polished, and the mandibles are 
minute, hard, pointed processes placed at its tip. This organ is 
admirably adapted to its purpose ; it being used for boring a hole 
in wood or bark, in which ;in egg is subsequently deposited. The 
males in these cases are extremely different, so that considerable 
curiosity is felt as to why this should be so. In some cases their 
head is thick, and there may be no rostrum, while large powerful 
mandibles are present. 

In other cases the rostrum is slender, but of enormous 
length, so that it may surpass in this respect the rest of 



v BRENTHIDAE AGLYCYDERIDAE 297 

the body, although this itself is so drawn out as to be quite 
exceptional in the Insect world : l the antennae are inserted 
near the tip of the rostrum instead of near its base, as they 
are in the female. The size of the males is in these cases usually 
much larger than that of the female.' 2 The males of some species 
tight ; they do not, however, wound their opponent, but merely 
frighten him away. In Eupscdis it appears that the rostrum 
of the female is apt to become fixed in the wood during her 
boring operations ; and the male then extricates her by pressing 
his heavy prosternum against the tip of her abdomen ; the stout 
forelegs of the female serve as a fulcrum and her long body as a 
lever, so that the effort of the male, exerted at one extremity of 
the body of the female, produces the required result at the other 
end of her body. The New Zealand Brenthid, Lasiorhynchus 
barlini/'iiis, exhibits sexual disparity in an extreme degree: the 
length of the male is usually nearly twice that of the female, and 
his rostrum is enormous. It is at present impossible to assign 
any reason for this ; observations made at the request of the 
writer by Mr. Helms some years ago, elicited the information 
that the female is indefatigable in her boring efforts, and that 
the huge male stands near by as a witness, apparently of the 
most apathetic kind. 



Coleoptera of uncertain position. 

There are three small groups that it is impossible at present 
to place in any of the great series of beetles. 

Fam. 84. Aglycyderidae. - - Tarsi three-jointed, the second 
joint lobed ; head not prolonged, to form a beak. The two most 
important features of Rhynchophora are absent in these Insects, 
while the other structural characters are very imperfectly known, 
many parts of the external skeleton being so completely fused 
that the details of structure are difficult of appreciation. AVest- 
wood considered the tarsi to be really four-jointed, but it is not 

1 In the males of the genus Cedeocera the tips of the elytra are drawn out into 
processes almost as long as the elytra themselves, and rivalling the forceps of 
earwigs. 

' The stature of the individuals of the same species is, in some of these 
Brenthidae, subject to extreme variation, especially in the males, some individuals 
of which in the case of Brcntltus anchorage* are five times as long as others. 



298 



COLEOPTERA 



CHAP. 



at all clear that the minute knot he considered the third joint is 

more than the articulation 
of the elongate terminal 
joint. The family consists 
only of two or three species 
of Aglycyderes, one of which 
occurs in the Canary 
Islands, and one or two in 
New Zealand and Xew 
Caledonia. The former is 
believed to live in the stems 
of Euphorbia canariensis ; 
Canary islands, a Xew Zealand species has 

_ ITT 1 * 




151. A'jtiic,/<i,<,Yx 



A, Imago ; B, tarsus according to Westwood ; 



C, according to nature ; 
labium. 



been found in connection 



D, maxilla ; E, 

with the tree-fern Cyathea 
dealbata. 

Fam. 85. Proterhinidae. Tarsi three-jointed, the second joint 
lol>ed ; head of the male scarcely prolonged, hut that of the female 
fur ming a definite rostrum.; maxillae and ligula entirely covered 
l>y the mention-. As in the preced- 
ing family the sutures on the 
under side of the head and pro- 
sternum cannot be detected. The 
minute palpi are entirely enclosed 
in the buccal cavity. There is a 
very minute true third joint of 
the tarsus, at the base of the ter- 
minal joint, concealed between the 
lobes of the second joint. The 
family consists of the genus Pro- 
terhinus ; it is confined to 
Hawaiian Islands, where these In- 
sects live on dead wood in the 
native forests. The genus is numerous in species and individuals. 

Strepsiptera (or Rhipiptera, Stylopidae).--Jfae small or 
minute; prothorax extremely small ; mesothorax moderate, the elytra 
reduced to small, free slips; metathorax and wings very large; 
nervuration of the latter radiating. n-Hliunt cross nervules. female 
a mere sac, with one extremity smaller and forming a sort of neck 
or head. These curious Insects are parasitic in the interior of 





the Fio. 152. 1'i-iifi'rliiiinx /eciiiifei. Ha- 
waiian Islands. A, Male ; B, female ; 
C, front foot, more magnified. 



STREPSIPTERA 



299 



other Insects, of the Orders Hymenoptera and Hemiptera. Their 
structure and their life-histories entitle them to be ranked as 
the most abnormal of all Insects, and entomologists are not 
agreed as to whether they are aberrant Coleoptera or a distinct 
Order. The newly-hatched larva is a minute triungulin (Fig. 
154), somewhat like that of Meloe; it fixes itself to the skin 
of the larva of a Hymenopterous Insect, penetrates into the 
interior, and there undergoes its metamorphoses, the male emerg- 
ing to enjoy a brief period of an abnormally active, indeed agitated, 
existence, while the female never moves. It is important to 
note that these Strepsiptera do not, like most other internal 
parasites, produce the death of their hosts ; these complete their 
metamorphosis, and the development of the parasite goes on 
simultaneously with that of the host, so that the imago of the 



FIG. 153. Sexes of Strep- 
siptera. A, Male of 
Stylops dalii (after Cur- 
tis) ; B, female of Xenos 
rossii (after von Siebold). 



B 





A 



Strepsipteron is found only in the imago of the host. 1 After 
the young Stylops has entered its host it feeds for a week or so 
on the fat-body (apparently by a process of suction), then 
moults and assumes the condition of a footless maggot, in which 
state it remains till growth is completed. At the latter part of 
this period the history diverges according to sex ; the female 
undergoes only a slight metamorphic development of certain 
parts, accompanied apparently by actual degradation of other 
parts ; while the male goes on to pupation, as is normal in Insects. 
(We may remark that the great features of the development of 
the sexes are parallel with those of Coccidae in Hemiptera.) 
AVhen the Hymenopterous larva changes to a pupa, the larva 
of the Strepsipteron pushes one extremity of its body between two 
of the abdominal rings of its host, so that this extremity becomes 
external, and in this position it completes its metamorphosis, the 

1 This remark applies to the Strepsiptera parasitic on Hymenoptera : nothing 
whatever is known as to the life-histories of the species that attack Hemiptera. 



;oo 



STREPSIPTERA 



CHAP. 



male emerging very soon after the host has become an active 
winged Insect, while the female undergoes no further change of 
position, but becomes a sac, in the interior of which young 
develop in enormous numbers, finally emerging from the mother- 
sac in the form of the little triungulins we have already 
mentioned. This is all that can be given at present as a general 
account ; many points of the natural history are still obscure, 
others have been merely guessed ; while some appear to differ 

greatly in the different forms. A few brief 
remarks as to these points must suffice. 

Bees carrying, or that have carried, Strep- 
siptera, are said to be stylopised (it being a 
species of the genus Xtylops that chiefly infests 
bees) ; the term is also used with a wider 
application, all Insects that carry a Strepsip- 
terous parasite being termed stylopised, though 
it may be a Strepsipteron of a genus A T ery 
different from ^////n/is that attacks them. The 
development of one or more Strepsiptera in 
an Insect usually causes some deformity in 
the abdomen of its host, and effects consider- 
able changes in the condition of its internal 
organs, and also in some of the external char- 
acters. Great difference of opinion prevails 
as to what these changes are ; it is clear, how- 
FIG. 154. Young larva ever, that they vary much according to the 

of Stulops on a Lee's- -11 T <? 

hair. Greatly magni- species, and also according to the extent of 

lied. (After Newport.) the stylopisatioii. Usually only one Stylops 

is developed in a bee ; but two, three, and 

even four have been observed: 1 and in the case of the wasp, 

Polistcs, Hubbard has observed that a single individual may 

bear eight or ten individuals of its Strepsipteron (Xenos, 

n. sp. ?). 

There is no exact information as to how the young triungulins 
find their way to the bee -larvae they live in. Here again the 
discrepancy of opinion that prevails is probably due to great 

1 Although not an invariable, it seems that it is a general rule that the Stylops 

produced from the body of one individual are all of one sex ; it has even been 

bated that female bees produce moiv especially female Stylops. and male bees 

male kstylupK. If any correlation as to this latter point exist, it is far from general. 




STYLOPS 



301 



difference really existing as to the method. When a Stylops 
carried by an Insect (a Hymenopteron, be it noted, for we have 
no information whatever as to Hemiptera) produces young, 
they cover the body of the host as if it were powdered, being 
excessively minute and their numbers very great ; many hundreds, 
if not thousands, of young being produced by a single Stylops. 
The species of the wasp genus Polistes are specially subject to 
the attacks of Stylops ; they are social Insects, and a stylopised 
specimen being sickly does not as a rule leave the nest ; in this 
case the Stylops larva may therefore have but little difficulty in 
finding its way to a Hymenopterous larva, for even though it 
may have to live for months before it has the chance of attaching 
itself to a nest -building female, yet it is clearly in the right 
neighbourhood. The bee genus Andrena has, however, quite 
different habits ; normally a single female makes her nest under- 
ground ; but in the case of a stylopised female it is certain that 
no nest is built, and no larvae produced by a stylopised example, 
so that the young triungulins must leave the body of the bee in 
order to come near their prey. They can be active, and have 
great powers of leaping, so that it is perhaps in this way possible 
for them to attach themselves to a healthy female bee. 

We have still only very imperfect knowledge as to the struc- 
ture and development of Strepsip- 
tera. Indeed but little informa- 
tion has been obtained since 
1S43. 1 Before that time the 
mature female was supposed to be 
a larva, and the triungulins found 
in it to be parasites. Although 
the erroneous character of these FlQ> 15 5._p or ti n of early stages of 
views has been made clear, the 
problems that have been sug- 
gested present great difficulties. 
Apparently the change from the 
triungulin condition (Fig. 154) to 
the parasitic larvae (Fig. 1 5 5, A, B) 
is extremely great and abrupt, and it appears also that during 




Xenos rossii. (After YOU Siebold.) 
A, Small male larva ; B, small female 
larva ; C, full-grown male larva ; D, 
full-grown female larva ; E, the so- 
called " cephalothorax " and adjacent 
segment of adult female. (The newly- 
hatched larva is very much like that 
of Styloi>s shown in Fig. 154.) 



1 Yon Siebold, Arch. Naturgcs. ix. 1843, pp. 137-161. Xassonoff's recent paper 
is in Russian, but so far as we can gather (cf. Zool. Centralbl. i. 1894, p. 766), it does 
not add greatly to the data furnished by von Siebold. 



3O2 STREPSIPTERA CHAP. 

the larval growth considerable sexual differentiation occurs (Fig. 
155, C, D) ; details are, however, wanting, and there exists but 
little information as to the later stages. Hence it is scarcely 
a matter for surprise that authorities differ as to which is the 
head and which the anal extremity of the adult female. Yon 
Siebold apparently entertained no doubt as to the part of the 
female that is extruded being the anterior extremity ; indeed lie 
called it a cephalothorax. Supposing this view to be correct, we are 
met by the extraordinary facts that the female extrudes the head 
for copulatory purposes, that the genital orifice is placed thereon, 
and that the young escape by it. Meinert l contends that the 
so-called cephalothorax of the adult is the anal extremity, and 
that fertilisation and the escape of the young are effected by the 
natural passages, the anterior parts of the body being affected by 
a complete degeneration. Nassonoff, in controversion of Meinert, 
has recently pointed out that the " cephalothorax " of the young- 
is shown by the nervous system to be the anterior extremity. It' 
still remains, however, to be shewn that the " cephalothorax " of 
the adult female corresponds with that of the young, and we shall 
not be surprised if Meinert prove to be correct. The internal 
anatomy and the processes of oogenesis appear to be of a very unusual 
character, but their details are far from clear. Brandt has given 
some particulars as to the nervous system ; though he does not 
say whether taken from the male or female, we may presume it to 
be from the former ; there is a supra -oesophageal ganglion, and 
near it a large mass which consists of two parts, the anterior repre- 
senting the sub-oesophageal and the first thoracic ganglia, while 
the posterior represents two of the thoracic and most of the 
abdominal ganglia of other Insects ; at the posterior extremity, 
connected with the other ganglia by a very long and slender 
commissure, there is another abdominal ganglion. 2 

It is a matter of great difficulty to procure material for the 
prosecution of this study ; the fact that the instars to be observed 
exist only in the interior of a few Hymenopterous larvae, which 
in the case of the bee, Andrenu, are concealed under ground ; and 
in the case of the wasps, Pulistes, placed in cells in a nest of 
wasps, adds greatly to the difficulty. It is therefore of interest 
to know that Strepsiptera occur in Insects with incomplete 

1 Ent. Meddcl. v. 1890. j>. 14S, and <>c. Dunske Selsk. 1896,, p. 67. 
-' i- Soc. cut. lluss. xiv. 1879, p. 14. 



STYLO PS 303 




metamorphosis. They have been observed in several species of 
Homoptera ; and the writer has a large Pentatomicl bug of 
the genus Collided, which bears a female 
Strepslpteron apparently of large size. This 
bug 1 is abundant and widely distributed in 
Eastern Asia, and it may prove compara- 
tively easy to keep stylopised examples 
under observation. Both v. Siebold and 
Nassonofif think parthenogenesis occurs in 
Strepsiptera, but there appear to be no facts FlG> i56.-Abdpmen of a 

wasp (Polistes heb- 

to warrant this supposition. Von Siebold rams) with a strep- 
speaks of the phenomena of Strepsipterous gg^ ^^ 
reproduction as paedogenesis, or pseudo- dorsal plates of the 

j i -,-, wasp's abdomen being 

paedogenesis, but we must agree with removed. , Project 
Meinert that they cannot be so classed. tion of P ai- t of the 

mi i f c i- f parasite ; b, line in- 

Ihe males of Strepsiptera live for only a dicating the position 
very short time, and are very difficult of of the remove <l Dorsal 

lllfltf* 

observation. According to Hubbard the 
males of Xenos dash about so rapidly that the eye cannot see 
them, and they create great agitation amongst the wasps in the 
colonies of which they are bred. Apparently they are produced in 
great numbers, and their life consists of only fifteen or twenty 
minutes of fiery energy. The males of Stylops are not exposed 
to such dangers as those of Xenos, and apparently live somewhat 
longer a day or two, and even three days are on record. The 
individuals of Andrena parasitised by Stylops are apparently 
greatly affected in their economy and appear earlier in the season 
than other individuals ; this perhaps may be a reason, coupled 
with their short lives, for their being comparatively rarely met 
with by entomologists. 

It is not possible at present to form a valid opinion as to 
whether Stylopidae are a division of Coleoptera or a separate 
Order. Von Siebold considered them a distinct Order, and 
Nassonoff, who has recently discussed the question, is also of that 
opinion. 

1 Named by Mr. Distant Callidea baro ; according to the Brussels catalogue of 
Hexniptera, Chrysocoris yrandis var. baro. 



CHAPTEE VI 

LEPIDOPTERA OR BUTTERFLIES AND MOTHS 

Order VI. Lepidoptera. 

Wings four ; body and wings covered with scales usually varie- 
gate in colour, anil <>n the body frequent!!/ lllore r less like 
lair: nervures moderate in number, at the prripJiery of 
one winy not exceeding fifteen, but little irregular ; eross- 
nervules not more than four, there being usually only one or 
two closed cells on each wing, occasionally none. Imago 
with mouth incapable of biting, 'usually forming a long 
coiled proboscis capable of protrusion. Metamorphosis great 
and abrupt ; the wings developed inside the bod// ; the larva 
with large or moderate head and strong mandibles. Pupa 
with the apt>i'inl,iges usuU tj nd pressed and cemented to the 
body so that it presents mnre or less even, horny exterior, 
occasionally varied by projections that are not the appendages 
and that may make the form very irregular: in many 
of the smaller forms the appendages are only imperfectly 
cemented to the body. 

LEPIDOPTERA, or butterflies and moths, are so far as ornament is 
concerned the highest of the Insect world. In respect of 
intelligence the Order is inferior to the Hymenoptera, in the 
mechanical adaptation of the parts of the body it is inferior to 
Coleoptera, and in perfection of metamorphosis it is second to 
Diptera. The mouth of Lepidoptera is quite peculiar; the pro- 
boscis the part of the apparatus for the prehension of food- 
is anatomically very different from the proboscis of the other 
Insects that suck, and finds its nearest analogue in the extreme 
elongation of the maxillae of certain Coleoptera, e.g. Nemognatlut. 



VI 



LEPIDOPTERA 



305 



The female has no gonapophyses, though in certain excep- 
tional forms of Tineidae, there are modifications of structure 
connected with the terminal segments, that have as yet been 
only imperfectly investigated. As a rule, the egg is simply 
deposited on some living vegetable and fastened thereto. 
Lepidoptera are the most exclusively vegetarian of all the Orders 
of Insects ; a certain number of their larvae prey on Insects 
that are themselves filled with vegetable juices (Coccidae, 




FIG. 157. Metamorphosis of a Lepidopteron (Rhegmatophila alpinct, Notodontidae). 
(After Poujade, Ann. Soc. cnt. France, 1891.) Europe. A, Egg ; B, young larva, 
about to moult ; C, adult larva ; D, head and first body-segment of adult larva, 
magnified ; E, pupa, x ; F, male moth in repose ; G, female moth iu repose. 

Aphidae) and a very small number (Tinea, etc.) eat animal 
matter. In general the nutriment appears to be drawn ex- 
clusively from the fluids of the vegetables, the solid matter 
passing from the alimentary canal in large quantity in the form 
of little pellets usually dry, and called frass. Hence the 
quantity of food ingested is large, and when the individuals 
unduly increase in number, forest trees over large areas are 
sometimes completely defoliated by the caterpillars. 

Lepidoptera pass a larger portion of their lives in the pupal 
stage than most other Insects do ; frequently during nine months 
of the year the Lepidopteron may be a pupa. In other Orders of 

VOL. VI X 



306 LEPIDOPTERA 



CHAP. 



Insects it would appear that the tendency of the higher forms is 
to shorten the pupal period, and when much time has to he 
passed between the end of the feeding up of the larva and the 
appearance of the imago, to pass this time as much as possible 
in the form of a resting-larva, and as little as may be in the 
form of a pupa ; in Lepidoptera the reverse is the case ; the 
resting-larva period being usually reduced to a day or two. 
Hence we can understand the importance of a hard skin to the 
pupa. There are, however, numerous Lepidopterous pupae where 
the skin does not attain the condition of hardness that is 
secured for the higher forms by the chitinous exudation we 
have mentioned ; and there are also cases where there is a pro- 
longed resting-larva period: for instance Galleria mellonella 
spins a cocoon in the autumn and remains in it as a resting 
larva all the winter, becoming a pupa only in the spring. In 
many of these cases the resting-larva is protected by a cocoon. 
It is probable that the chief advantage of the perfect chitinous 
exudation of the Lepidopterous pupa is to prevent the tiny, 
complex organisation from the effects of undue transpiration. 
Bataillon has suggested that the relation of the fluid contents of 
the pupa to air and moisture are of great importance in the 
physiology of metamorphosis. 

The duration of life is very different in various forms 
of Lepidoptera. It is known that certain species (Ephestia, 
Jcuehniella, e.g.*) may go through at least five generations a year. 
On the other hand, certain species that feed on wood or roots 
may take three years to complete their life-history ; and it is 
probable that some of the forms of Hepialidae are even longer 
lived than this. 

Lepidoptera have always been a favourite Order with ento- 
mologists, but no good list of the species has ever been made, 
and it would be a difficult matter to say how many species are 
at present known, but it can scarcely be less than 50,000. In 
Britain we have about 2000 species. 

The close affinity of the Order with Trichoptera has long 
been recognised : Reaumur considered the latter to be practically 
Lepidoptera with aquatic habits, and Speyer pointed out the 
existence of very numerous points of similarity between the 
two. Brauer emphasised the existence of mandibles in the 
nymph of Trichoptera as an important distinction : the pupa 



VI 



EXTERNAL STRUCTURE 



307 



of Micropteryx (Fig. 211) has however been recently shown to 
be similar to that of Trichoptera, so that unless it should be 
decided to transfer Micropteryx to Trichoptera, and then define 
Lepidoptera and Trichoptera as distinguished by the condition of 
the pupa, it would appear to be very difficult to retain the two 
groups as distinct. 

Structure of Imago. The head of a Lepidopteroii is in large 
part made up of the compound eyes ; in addition to these it 
frequently bears at the top a pair of small, simple eyes so much 
concealed by the scales as to cause us to wonder if seeing 
be carried on by them. The larger part of the front of the 
head is formed by the clypeus, which is separated by a well- 




FIG. 158. External structure of a female butterfly, Anosia pleswppus. (After Scudcler.) 
, Base of antenna ; b, pronotum ; b", scutum of mesothorax ; c, clypeus ; ex, coxa ; 
d, scutellum ; d 1 , scutellum of metathorax ; e, post-scutellum ( = base of phragma) ; 
em, epimerou ; ep, episternum ; /, scutum of metathorax ; m, basal part of pro- 
boscis ( = maxilla); o, eye; p, labial palp; /, mesosteruum ; s, prothoracic 
spiracle ; t, tegula ; tr, trocliauter ; 1-9, dorsal plates of abdomen. 

marked line from the epicranium, the antennae being inserted 
on the latter near its point of junction with the former. There 
is sometimes (Saturnia, Castnici) on each side of the clypeus a 
deep pocket projecting into the head-cavity. The other parts of 
the head are but small. The occipital foramen is very large.' 

The antennae are always conspicuous, and are very various 
in form ; they are composed of numerous segments, and in the 
males of many species attain a very complex structure, especially 
in Bornbyces and Fsychidae ; they doubtless function in such 
cases as sense-organs for the discovery of the female. 

The largest and most important of the mouth-parts are the 
maxillae and the labial palpi, the other parts being so small as 
to render their detection difficult. The labruni is a very short, 

1 Kellogg, Kansas Quarterly, ii. 1893, p. 51, plate II. 



308 



LEPIDOPTERA 



CHAP. 



comparatively broad piece, visible on the front edge of the 
clypeus ; its lateral part usually forms a prominence which has 
often been mistaken for a mandible ; Kellogg has applied the 
term " pilifer " to this part. In the middle of the labrum a small 
angular or tongue-like projection is seen just over the middle of 
the base of the proboscis ; this little piece is considered by 
several authorities to be an epipharynx. 

MANDIBLES. Savigny, Westwood, and others considered the 
parts of the labrum recently designated pilifers by Kellogg to 
be the rudimentary mandibles, but Walter has shown that this 




FIG. 159. Mouth of Lepidoptera. Tiger-moth, Arctia cajn. A, Seen from front : B, 
from front and below. , Clypeus ; b, labrum ; c, epipharynx ; d, mandibular 
area ; d', prominence beneath mandibular area ; e, one side of haustellum or pro- 
boscis ; f, maxillary palp ; g, labial palp. 

is not the case. 1 The mandibles are usually indistinguish- 
able, though they, or some prominence possibly connected with 
them, 2 may frequently be detected in the neighbourhood of 
the pilifers ; they are, according to Walter, largest and most 
perfectly developed in Erioceplmla, a genus that was not dis- 
tinguished by him from Micropteryx and was therefore termed 
" niedere Micropterygiuen," i.e. lower Micropteryges. The 
opinion entertained by Walter that Micropteryx proper (his 
" hohere Micropteryginen ") also possesses rudimentary mandibles 
is considered by Dr. Chapman, no doubt with reason, to be 
erroneous. 3 The mandibles, however, in the vast majority of 
Lepidoptera can scarcely be said to exist at all in the imago ; 
there being only an obtuse projection without trace of 

1 Jena. Zcitschr. Naturw. xviii. 1885, p. 751. 

; The writer is not quite convinced that the supposed mandibles of these Macro- 
lepidoptera are really entitled to be considered as such. 
3 Tr. cnt. Soc. London, 1893, p. 263. 



vi MOUTH-PARTS 309 

articulation on each side of the labrum ; and even this pro- 
jection is usually absent. Meinert recognised these projections as 
mandibles in Smerinthus popttli, and Kellogg in Protoparce Caro- 
lina, another large Sphinx moth. They appear to be unusually 
well developed in that group. In Castnia they are even more 
definite than they are in Sphingidae. 

The MAXILLAE are chiefly devoted to the formation of the 
proboscis. Their basal portions are anatomically very indefinite, 
though they exist very intimately connected with the labium. 
Each usually bears a small tubercle or a segmented process, the 
representative of the maxillary palpus. The proboscis itself con- 
sists of the terminal, or outer, parts of the two maxillae, which 
parts are closely and beautifully coadapted to form the spirally 
coiled organ, that is sometimes, though incorrectly, called the tongue. 
The exact morphology of the Lepidopterous proboscis has not 
been established. The condition existing in the curious family 
Prodoxidae (see p. 432), where a proboscis coexists with another 
structure called a maxillary tentacle, suggests a correspondence 
between the latter and the galea of a typical maxilla ; and 
between the proboscis and the lacinia or inner lobe of a 
maxilla : but J. B. Smith is of opinion that the tentacle in 
question is a prolongation of the stipes. The condition of the 
parts in this anomalous family (Prodoxidae) has not, however, 
been thoroughly investigated, and Packard takes a different 
view of the proboscis ; he considers that " it is the two galeae 
which become elongated, united and highly specialised to form 
the so-called tongue or glossa of all Lepidoptera above the 
Eriocephalidae." The proboscis in some cases becomes very 
remarkable, and in certain Sphingidae is said to attain, when 
unrolled, a length of ten inches. In some cases the maxillary 
lobes do not form a proboscis, but exist as delicate structures, 
pendulous from the mouth, without coadaptation (Zeuzera aesculi, 
the Wood -leopard moth). In other forms they are absent 
altogether (Cossus, e.g.}, and in Jfcpialus we have failed to detect 
any evidence of the existence of the maxillae. On the other 
hand, in Micropteryx the maxillae are much more like those of a 
mandibulate Insect ; and various other Microlepidoptera approach 
more or less a similar condition. In the genus last mentioned 

1 Amcr. Natural, xxix. 1895, p. 637. It should be recollected that many 
Lepidoptera do not possess any proboscis. 



310 LEPIDOPTERA CHAP. 

the maxillary palpi are largely developed, flexible and slender. 
According to Walter various forms of palpus intermediate between 
that of Micropteryx and the condition of rudimentary tubercle 
may be found amongst the Microlepidoptera, 1 

LABIUM. The labial palpi are usually largely developed, 
though but little flexible ; they form conspicuous processes 
densely covered with scales or hairs, and curve forwards or up- 
wards, rarely downwards, from the under side of the head, some- 
what in the fashion of tusks. The other parts of the labium 
are frequently represented merely by a membranous structure, 
united with the maxillae and obstructing the cavity of the 
pharynx. Where the proboscis is absent it is difficult to find 
any orifice leading to the alimentary canal, such opening as 
may exist being concealed by the overhanging clypeus and 
labium. In some forms, Saturnia, e.g., there appears to be no 
buccal orifice whatever. In Hepialus the labium is in a very 
unusual condition ; it projects externally in the position usually 
occupied by the labial palpi, these organs being themselves 
extremely short. It is very difficult to form an opinion as to 
the structure of the labium and other mouth -parts when the 
maxillae are not developed, as in these cases the parts are of 
a delicate membranous nature, and shrivel after death. This 
is the explanation of the fact that in descriptive works we find 
vague terms in use such as " mouth aborted " or " tongue absent." 

The mouth of the Lepidopterous imago is a paradoxical 
structure ; it differs very greatly from that of the larva, the 
changes during metamorphosis being extreme. We should thus 
be led to infer that it is of great importance to the creatures ; 
but, on the other hand, the various structures that make up the 
mouth, as we have remarked, are frequently absent or reduced to 
insignificant proportions ; and even in forms where the apparatus 
is highly developed the individuals seem to be able to accomplish 
oviposition without taking food, or after taking only very minute 
quantities. It is therefore difficult to understand why so great 
a change should occur during the metamorphosis of the Insects 
of this Order. It has been ascertained that in some forms where 
the mouth is atrophied the stomach is in a correlative condition ; 
but we are not aware that any investigations have been made 
as to whether this correspondence is general or exceptional. 

1 Jena. Zdtschr. Natunv. xviii. 1885, p. 168. 



vi EXTERNAL STRUCTURE 3 1 I 

The exact mode in which the proboscis acts is in several 
respects still obscure, the views of Burmeister and Newport being 
in some points erroneous. Towards the tip of the proboscis 
there are some minute but complex structures considered by 
Fritz Miiller to be sense-organs, and by Breitenbach to be 
mechanical instruments for irritating or lacerating the delicate 
tissues of blossoms. It is probable that Miiller's view will prove 
to be correct. Nevertheless the proboscis has considerable 
power of penetration ; there being a moth, " Ophideres fullonica" 
that causes considerable damage to crops of oranges by inserting 
its trunk through the peel so as to suck the juices. 1 The canal 
formed by each maxilla opens into a cavity inside the front part 
of the head. This cavity, according to Burgess, 2 is a sort of sac 
connected with five muscles, and by the aid of this apparatus the 
act of suction is performed : the diverticulum of the alimentary 
canal, usually called a sucking-stomach, not really possessing the 
function formerly attributed to it. 

The PROTHORAX is very small, being reduced to a collar, be- 
tween the head and the alitrunk, of just sufficient size to bear the 
front pair of legs. Its most remarkable feature is a pair of pro- 
cesses, frequently existing on the upper surface, called " patagia." 
These in many cases (especially in Noctuidae) are lobes capable 
of considerable movement, being attached only by a narrow- base. 
In Hepialus, on the contrary, they are not free, but are merely 
indicated by curved marks on the dorsum. The patagia are 
styled by many writers " tegulae." They are of some interest in 
connection with the question of wing -like appendages on the 
prothorax of Palaeozoic insects, and they have been considered 
by some writers 3 to be the equivalents of true wings. The 
MESOTHORAX is very large, especially its upper face, the notum, 
which is more or less convex, and in the higher forms attains a 
great extension from before backwards. The notum consists in 
greater part of a large anterior piece, the meso-scuturn, and a 

1 Amer. Natiiral. xiv. 1880, p. 313. 

- For an account of the structures at the tip of the proboscis of this moth, and 
of the beautiful manner in which the lobes of the maxillae are dovetailed together, 
see Francis Darwin, Quart. J. Micr. Sd. xv. 1875, p. 385. For details as to 
numerous proboscides, and as to the difficulties that exist in comprehending the 
exact mode of action of the organ, refer to Breitenbach's papers, especially Jena. 
Zcitschr. Xaturw. xv. 1882, p. 151. 

3 See Cholodkovsky, Zool. Anz. ix. p. 615 ; Haase, t.c. p. 711 ;. also Riley, 
P. cut. Soc. Washington, ii. 1892, p. 310. 



3 I 2 LEPIDOPTERA CHAP. 

smaller part, the meso-scutellum behind. In front of the scutum 
there is a piece termed prae-scutum by Burgess. It is usually 
small and concealed by the front part of the scutum ; but 
in Hepialus it is large and horizontal in position. It is of 
importance as being the chief point of articulation with the pro- 
thorax. The scutellum is more or less irregularly rhomboidal in 
form ; its hinder margin usually looks as if it were a lobe or fold 
placed in front of the base of the abdomen or metathorax, accord- 
ing to whether the latter is concealed or visible. In some of the 
higher forms this meso-scutellar lobe is prominent, and there 
may be seen under its projection a piece that has been called 
the post-scutellum, and is really the base of the great meso- 
phragma, a chitinous piece that descends far down into the 
interior of the body. In addition to the front pair of wings the 
inesothorax bears on its upper surface another pair of appendages, 
the tegulae : in the higher forms they are of large size ; they are 
fastened on the front of the inesothorax, and extend backwards 
over the joint of the wing with the body, being densely covered 
with scales so that they are but little conspicuous. These 
appendages are frequently erroneously called patagia, but have 
also been called scapulae, pterygodes, paraptera, and shoulder- 
tufts, or shoulder-lappets. The lower surface of the inesothorax 
is much concealed by the large and prominent coxae, but the 
sternum and the two pleural pieces on each side, episternum and 
epimeron, are easily detected. The area for attachment of the 
anterior wing on each side is considerable, and appears to be of 
rather complex structure ; its anatomy has been, however, but 
little studied. 

The METATHORAX is small in comparison with the preceding 
segment, to which it is intimately co-adapted, though the two 
are really connected only by delicate membrane, and can conse- 
quently be separated with ease by dissection. The metanotum 
consists of (1) the scutum, which usually appears externally as 
an anterior piece on each side ; (2) the scutellum, forming a 
median piece placed behind the scutum, which it tends to 
separate into two parts by its own extension forwards. In order 
to understand the structure of the metathorax it is desirable to 
dissect it off from the larger anterior segment, and it will then 
be found that its appearance when undissected is deceptive, 
owing to its being greatly arched, or folded in the antero- 



vi EXTERNAL STRUCTURE 3 I 3 

posterior direction. A broad, but short phragma descends from 
the hind margin of the metascutellum into the interior of the 
body. It should be noted that though the metanotum is forced, 
as it were, backwards by the great extension of the mesonotum in 
the middle line of the body, yet at the sides the metanotum creeps 
forward so as to keep the points of attachment of the hind wings 
near to those of the front wings. In many forms of Hesperiidae, 
Sphingidae, Noctuidae, etc, the true structure of the metanotum 
is further concealed by the back of the mesoscutelluni reposing 
on, and covering it. 

Difference of opinion exists as to the thoracic SPIRACLES ; there 
is one conspicuous enough in the membrane behind the pronotum, 
and it is thought by some writers that no other exists. West- 
wood and Scudder, however, speak of a mesothoracic spiracle, and 
Dr. Chapman considers that one exists. Minot describes l a 
structure behind the anterior wing, and thinks it may be an 
imperfect spiracle, and we have found a similar stigma in 
Saturnia pavonia. At the back of the thorax there is on each 
side in some Lepidoptera (Noctuidae, Arctia, etc.), a curious large 
cavity formed by a projection backwards from the sides of the 
metasternum, and a corresponding development of the pleura of 
the first abdominal segment. Minot and others have suggested 
that this may be an organ of hearing. 

The ABDOMEN differs according to the sex. In the female 
seven segments are conspicuous dorsally, but only six ventrally, 
because the first segment is entirely membranous beneath, and 
is concealed between the second abdominal ventral plate and 
the posterior coxae. Besides these segments there are at the 
hind end two others smaller, more or less completely with- 
drawn into the body, and in certain cases forming an ovipositor. 
These nine segments are usually considered to constitute the 
abdomen; but according to Peytoureaiv a tenth dorsal plate is 
represented on either side of the anal orifice, though there is no 
trace of a corresponding ventral plate. In the male the segments, 
externally conspicuous, are one more than in the female. According 
to the authority quoted, 3 this sex has also truly ten abdominal 
segments, the ninth segment being withdrawn to a greater or 

1 Fourth Rep. U.S. Entom. Commission, 1885, p. 49. 

2 C.R. Ac. Set. Paris, cxviii. 1894, p. 360 ; and his Thesis, Bordeaux, 1895. 
3 C.R. Ac. Set. Paris, cxviii. 1894, p. 542. 



LEPIDOPTERA 



CHAP. 



less extent to the inside of the body, and modified to form part of 
a copulatory apparatus ; its dorsal portion bears a process called the 
" uncus " ; the anal orifice opens on the inner face of this process, 
and below it there is another process developed to a greater or 
less extent called the " scaphium." The ventral portion of the 
ninth segment bears a lobe, the " saccus " (Peytoureau, I.e.'). On 
each side of the ninth abdominal segment there is a process called 
the " valve," the internal wall of which bears some hook-like or 
other processes called " harpes " ; it is continued as a membrane 
surrounding the "oedeagus," or penis, and bearing more or 
less distinct prominences connects with the scaphium. In many 
forms the parts alluded to, other than the valves, are concealed 

by the latter, which 
come together when 
closed, and may lie 
covered externally with 
scales like the rest of 
the abdomen. Peytou- 
reau considers that the 
uncus is really the dorsal 
plate of a tenth segment, 
and that the scaphium 
is the tenth ventral 
Thus, according 
view, the ninth 




FIG. 160. Acherontia (ifr<i/><~is. The termination of <J 
body, one side removed. 7A', Ninth dorsal plate ; 
IX', ninth ventral ; s, lobe, saccus, of ninth 
ventral plate ; A', tenth dorsal plate, or uncus ; 
sc, scaphium, or tenth ventral plate ; , position of 
anus ; b, chitinised band of scaphium ; V, valve or Segment IS extensive and 
clasper : c, hooks or harpes, of clasper ; p, penis comp l ex ], e ing very 
(or oedeagus). (Alter Peytoureau.) - 1 - 

highly modified in all 

its parts : while the tenth segment is greatly reduced. The 
structure of the male organs is simpler in Lepidoptera, and less 
varied than it is in the other great Orders of Insects. There 
are seven pairs of abdominal spiracles on the upper parts of the 
membranous pleurae. 

LEGS. The legs are long, slender, covered with scales, and 
chiefly remarkable from the fact that the tibiae sometimes bear 
articulated spurs on their middle as well as at the tip. The 
front tibia usually possesses on its inner aspect a peculiar 
mobile pad ; this seems to be in some cases a combing organ ; 
it also often acts as a cover to peculiar scales. The tarsi 
are five-jointed, with two small claws and a small apparatus, 



vi WINGS 3 I 5 

the functional importance of which is unknown, between the 
claws. 

Wings. The wings are the most remarkable feature of this 
Order ; it is to them that butterflies owe their beauty, the sur- 
faces of the wings being frequently adorned with colours and 
patterns of the most charming and effective nature. These 
effects are due to minute scales that are implanted in the wing- 
membrane in an overlapping manner, somewhat similar to the 
arrangement of slates on the roof of a house. The scales are very 
readily displaced, and have the appearance of a silky dust. "We 
shall describe their structure and allude to their development 
subsequently. The wings are usually of large size in com- 
parison with the Insect's body : in the genus Morplio, the 
most gorgeous of the butterflies, they are enormous, though the 
body is small ; so that when deprived of these floats the Insect is 
insignificant. The great expanse of wing is not correlative with 
great powers of flight, though it is perhaps indicative of flying with 
little exertion ; for the small-winged Lepidoptera, Sphingidae, 
etc., have much greater powers of aerial evolution than the large- 
winged forms. The area of the wing is increased somewhat by 
the fact that the scales on the outer margin, and on a part or on 
the whole of the inner margin, project beyond the edges of the 
membrane that bears them : these projecting marginal scales are 
called fringes. In many of the very small moths the actual size 
of the wing-membranes is much reduced, but in such cases the 
fringes may be very long, so as to form the larger part of the 
surface, especially of that of the hind wings. Frequently the 
hind wings are of remarkable shape, being prolonged into pro- 
cesses or tails, some of which are almost as remarkable as those 
of Nemoptera in the Order Neuroptera. 

The wings are very rarely absent in Lepidoptera ; this occurs 
only in the female sex, no male Lepidopterous imago destitute of 
wings having been discovered. Although but little is known of 
the physiology of flight of Lepidoptera, yet it is clearly important 
that the two wings of the same side should be perfectly coadapted 
or correlated. This is effected largely by the front wing over- 
lapping the hind one to a considerable extent, and by the two 
contiguous surfaces being pressed, as it were, together. This is 
the system found in butterflies and in some of the large moths, 
such as Lasiocampidae and Saturniidae ; in these cases the hind 



316 LEPIDOPTERA 



CHAP. 



wing always has a large shoulder, or area, anterior to its point 
of insertion. In most moths this shoulder is absent, but in its 
place there are one or more stiff bristles projecting forwards and 
outwards, and passing under a little membranous flap, or a tuft 
of thick scales on the under face of the front wing ; the bristle is 
called the " frenulum," the structure that retains it a " retinaculum." 
In Cost iiia (Fig. 162) and in some Sphingidae there is the un- 
usual condition of a highly-developed shoulder (s) coexisting with 
a perfect frenulum (/) and retinaculum (r). The frenulum and 
retinaculum usually differ in structure, and the retinaculum in 
position, in the two sexes of the same moth ; the male, which 
in moths has superior powers of flight, having the better retaining 
organs. Hampson says " the form of the frenulum is of great 
use in determining sex, as in the males of all the forms that 
possess it, it consists of hairs firmly soldered together so as to 
form a single bristle, whilst in nearly all females it consists of 
three or more bristles which are shorter than that of the male ; 
in one female Cossid I have found as many as nine. Also in the 
large majority of moths the retinaculum descends from the costal 
nervure in the male, while in the female it ascends from the 
median nervure." This sexual difference in a structure for the 
discharge of a function common to the two sexes is a very re- 

O i/ 

markable fact. There are a few very few moths in which the 
bases of the hind wings are not well coadapted with the front 
wings, and do not possess a frenulum, and these species possess 
a small more or less free lobe at the base of the front wing that 
droops towards the hind wing, and may thus help to keep up an 
imperfect connexion between the pair ; this lobe has been named 
a juguin by Professor Comstock. Occasionally there is a jugum 
on the hind as well as on the front wing. There is usually a 
very great difference between the front and the hind wings ; for 
whereas in the front wing the anterior portion is doubtless of 
great importance in the act of flight and is provided with 
numerous veins, in the hind wing, on the other hand, the corre- 
sponding part has not a similar function, being covered by the 
front wing ; hence the hind wing is provided with fewer ncrvures 
in the anterior region, the divisions of the subcostal being less 
numerous than they are in the front wing. In the moths 
possessing a jugum the two wings differ but little from one 

1 Fauna of British India, Moths, i. 1892, p. 6. 



vi YVING-NERVURES 3 1 7 

another, and it is probable that they function almost as four 
separate wings instead of as two pairs. 

WING-NEEVURES. The nervures or ribs of the wings are of 
great importance in Lepidoptera, as at present they furnish the 
chief characters for classification and for the discussions of 
phylogeny that are so numerous in entomological literature. 
On looking at wings that have been deprived of their scales it 
will be noticed (Fig. 161) that the ribs are much more numerous 
at the outer margins than they are near the points of attachment 
of the wings, and that there is usually but one cell (or area com- 
pletely enclosed by ribs). This latter point is one of the chief 
peculiarities of the Lepidopterous wing ; in Insect-wings generally 
the number of cells in proportion to the area of the wings and to 
the number of nervures is greater than it is in Lepidoptera, for 
in the latter there are few or no cross-nervures. Hence there is 
sometimes no closed cell at all on the wing (Fig. 161, II. B). 
The maximum number of closed cells is six ; this is found in 
some species of Micropteryx, while in Hepialus there may be 
three or four ; but the rule is that there is only one cell in the 
Lepidopterous wing. When the number of cells is increased 
this is not necessarily due to an increase in the cross-nervures; and 
in fact it is generally due to irregular forking or to the sinuous 
form of the longitudinal nervures themselves (see wing of Castnia, 
Fig. 162, A.). Some authorities consider that all transverse or 
cross-veins in Lepidoptera are merely portions of longitudinal 
veins having diverted courses. When a portion of a nervure 
beyond the basal or primary portion serves as a common piece 
to two forked parts external to it, it is called a stalk (Fig. 162, 
A, e). There are cases in which the furcation takes place in the 
opposite direction, so that a nervure is double at the base of the 
wing (Fig. 161, I, A, la, and B, 16). This important condition 
has not yet been adequately discussed. 

Turning to the mode of designation of the nervures, 1 we may 

1 It is impossible for us to treat of the difficulties that exist on this point, and 
we must refer the student to the pamphlet, "The Venation of the Wings of 
Insects," by Prof. Comstock, Ithaca, 1895, being a reprint, with an important 
pivfatory note, from the Elements of Insect Anatomy, by J. H. Comstock and V. L. 
Kellogg, also to Packard's discussion of the subject in Mem. Ac. Sci. Wasliiujtvu. 
vii. 1895, pp. 84-86. The method of Spuler, alluded to in these two memoirs, is 
based on development, and, when extended, will doubtless have very valuable 
results. See Spuler, Zcitschr. wiss. Zool. liii. 1892, p. 597. 



LEPIDOPTERA 



CHAP. 



commence by remarking that no system satisfactory from a 
practical as well as from a theoretical point of view has yet been 
devised. The diagrams given in figure 1G1 will enable us to 
explain the methods actually in vogue ; I. representing the system, 
dating from the time of Herrich - Schaeffer, chiefly used by 
British naturalists, and II. that adopted by Staudinger and 
Schatz in their recent great work on the Butterflies of the world. 
The three anterior nervures in both" front and hind wings 
correspond fairly well, and are called, looking at them where 
they commence at the base of the wing, " costal," " subcostal," and 




FIG. 161. Wing-nervuratiou of Lepidoptera. I, Diagram of moths' wings (after Hamp- 
son) ; II, of a butterfly's wings (Mi,rji/n> ini'in'Inux ,5, after Staudinger and Schatz). 
A, front, B, hind wing. I. c, costal ; sc, subcostal ; //(, median ; In. Ifi, lr, in- 
ternal nervures ; /, frenulum ; 2, 3, 4, branches of median nervure ; 5, lower 
radial; 6, upper radial; 7-11, divisions of the subcostal; 12, termination of 
costal ; c, cell ; d, discocellular nervure. II. C, costal ; SC, subcostal ; M, median ; 
SM and SN, submedian nervures ; 1A, inner-margin nervure ; UR, lower radial ; 
OR, upper radial ; SC 1 to SC 5 , divisions of subcostal ; M 1 to M 3 , divisions of median 
uervure ; C, cell ; DC, discocellulars. 

" median " nervures. The nervures near the inner margin of the 
wing (that is the lower part in our figures) differ much in the front 
and hind wings, consisting either of two or of three separate 
portions not joined even at the base. British entomologists call 
these "branches or divisions of the internal nervure": the 
Germans call the more anterior of them the " submedian," and the 
more internal the "inner-margin nervure"; they are also frequently 
called anal nervures. The cross-nervure that closes the cell is 
called discocellular; when apparently composed of two or three 



WING-NERVULES 



319 



parts joined so as to form angles, the parts are called, according 
to position, upper, lower, and middle discocellulars. One or more 
short spurs may exist on the front part of the basal portion of 
the hind wing ; these are called praecostal. The branches or 
terminal divisions of the nervures should be called nervules ; they 
are usually mentioned by the numbers shewn in the diagram 
(Fig. 161, I.). In addition to this, it is only necessary to re- 
member that number 2 is always assigned to the posterior division 
of the median nervure, the nervules below this being all called 1, 
and distinguished by the addition of 
a, I), c when requisite. This course 
is necessary, because if it were not 
adopted the corresponding nervules 
on the front and hind wings would 
bear different numbers. 

The use of this system of num- 
bers for the nervules is becoming 
general, and it answers fairly well 
for practical purposes. On the other 
hand, extreme discrepancy exists as 
to the nomenclature of the nervures 
and nervules, and there are almost as 
many systems as there are authorities. 

The normal number of nervules 
is, on the front wing, 11+1 or 2 




B 



inner marginal, and on the 



wng 



+ 2 or 3 inner marginal. 



hind 
In 



genus 



FIG. 162. Wing - nervuration of 
Castnia. Undersides of, A, front, 
B, hind wings, la, lb, Ic, Irf, 
Inner marginal nervnres ; 2, lower 
branch of median ; 8, subcostal 
of hind wing ; 1'2, subcostal of 
front wing; e, "stalk" of 8 and 
9 ; _/", frenuluni ; r, retinaculum ; 
s, shoulder ; ij, articulation of 



the aberrant moths of the 

Castnia the nervuration is unusually 

complex and irregular (Fig. 162), and an analogous condition 

occurs in our common Goat - moth (Cossus ligniperda). In 

Jfi-jiinliis and Micropteryx (the jugate moths of Comstock) the 

hind wings are less dissimilar in nervuration from the front 

wings than they are in other Lepidoptera. 1 

Internal Anatomy. ~ The alimentary canal extends as a long, 

The structure and development of scales and nervures is dealt with as part of 
the brief study of the development of the wing, on p. 329, etc. 

2 The internal anatomy of Lepidoptera has not been extensively studied. For 
information refer to Dufour, L'.Il. Ac. Paris, xxxiv. 1852, p. 718; Scudder, JJi/tf. 
Xi'/r England, i. 1SS9, p. 47 ; Minot and Bur^rss. Fourth Jtcj>. I'. S. Entmn. 
1885, p. 53. 



320 



LEPIDOPTERA 



CHAP. 



slender oesophagus through the length of the thorax, dilating when 
it reaches the abdomen to form a tubular stomach ; before this 
it is somewhat enlarged to form an indistinct crop, and gives off 
a large diverticulum usually called a sucking stomach. According 
to Burgess, this structure does not possess the function ascribed 
to it by this name, and he terms it a food-reservoir. The Mai- 
pi ghian tubes are six in number, three on each side, and each set 
of three unite to form a common tube opening into the posterior 
extremity of the stomach ; behind them the alimentary canal 
continues in the form of a slender, tortuous intestine, expanding 




FIG. 163. Internal anatomy of Lepidoptera. Section of the body of a female butterfly, 
Anosia j>/r.ri/>i>its. (After Scudder. ) The portion to the left of the vertical line 
more magnified. I. II. Ill, thoracic segments ; 1-9, abdominal segments ; , 
antenna ; , anus ; ac, aortal chamber ; ag l , etc., abdominal ganglia ; agl 1 , agl~, 
accessory glands ; an, aorta ; br. brain ; c, colon ; cp, copnlatory pouch ; ex 1 , ex-, 
cjc 3 , coxae ; _/>. food-reservoir ; g 1 , suboesophageal ganglion ; h, dorsal vessel ; i, 
intestine ; I in, area filled by wing muscles ; Lor, ovary, or egg-tubes of left side ; 
mi-, Malpighian tube (the two others of the right side cut away, except small por- 
tions) ; ni.c, maxilla ; u, oviduct ; uo, its orifice ; oe, oesophagus ; ov.c, end of left 
ovary ; />, labial palp ; j/h, pharynx ; r.ov, terminal jiarts of right ovarian tubes, 
turned to one side, after the tubes have been cut away ; &</, salivary duct ; sgl, 
salivary gland ; .*//, .spermatheca ; st, stomach ; tg, thoracic ganglia ; r, copulatory 
orifice. 

at the extremity of the body to form a rectum. The dorsal or 
circulatory vessel commences near the posterior extremity of the 
body, but in the front part of the abdomen is deflexed to pass 
under the great phragma into the thorax, where it rises abruptly 
to the dorsal wall, but is again abruptly deflexed, forming a loop, 
and is then prolonged above the oesophagus into the head : at 
the summit of the thoracic loop there may be a dilatation called 
the aortal chamber. The supra- and infra-oesophageal ganglia 
are consolidated into a mass pierced by the oesophagus : there is 
a minute frontal ganglion ; the ventral chain consists of three 
much approximated thoracic ganglia and four abdominal ganglia 
separated from the thoracic by a long interval. 



vi INTERNAL STRUCTURE EGG 321 

The male sexual organs consist of the two testes placed in a 
common capsule, from which proceed a pair of contiguous vasa 
deferentia (dilated soon after their origin to form the vesiculae 
seminales) ; into each vas there opens a long, tubular gland ; the 
two vasa subsequently unite to form a long, coiled, ejaculatory 
duct. It is in the structure of the female sexual organs that the 
most remarkable of the anatomical characters of Lepidoptera is 
found, there being two external sexual orifices. The imago has, 
in the great majority of cases, four egg-tubes in each ovary ; the 
pair of oviducts proceeding from them unite to form a single un- 
paired (azygos) oviduct which terminates by an orifice quite at 
the posterior extremity of the body. There is a sac, the bursa 
copulatrix or copulatory pouch, which is prolonged in a tubular 
manner, to open externally on the eighth ventral plate : a tube, 
the seminal duct, connects the bursa with the oviduct, and on this 
tube there may be a dilatation the spermatheca. Besides these 
structures two sets of accessory glands open into the oviduct, an 
unpaired gland, and a pair of glands. The development of these 
structures has been described by Hatchett Jackson, 1 and exhibits 
some very interesting features. The exact functions of the bursa 
copulatrix and of the other structures are by no means clear. 
According to Riley, 2 the spermatheca in Pronvla contains some 
curious radiate bodies, and Godman and Salviii describe some- 
thing of the same sort as existing in butterflies. Several varia- 
tions in the details of the structure of these remarkably complex 
passages have been described, and the various ducts are some- 
times rendered more complex by diverticula attached to them. 
Some noteworthy diversities in the main anatomical features 
exist. According to Cholodkovsky, there is but one sexual 
aperture the posterior one in Nematois metallicus ; while, 
according to Brandt, the number of egg-tubes in a few cases 
exceeds the normal four being in Sesia scoliaeformis fourteen. 
In Nematois metallicus there is individual variation, the number 
of tubes varying from twelve to twenty. 

The egg has been more extensively studied in Lepidoptera 
than in any other Order of Insects. It displays great variety : 
we meet with elongate forms (Fig. 164) and flat forms like 
buttons, while in Limacodes (Fig. 83, Vol. Y.) the egg is a 

1 Tr. Linn. Soc. London (2), v. 1890, p. 143. 
" P. cut. Soc. U'ashiwjton, ii. 1892, p. 305. 
VOL. VI Y 



322 



LEPIDOPTERA 



CHAP. 



transparent scale of somewhat inconstant outline. Some are 
coloured and mottled somewhat after the fashion of birds'-eggs ; 
this is the case with some eggs of Lasiocampidae and Liparidae ; in 
some the sculpture of the egg-shell is of the most elaborate char- 
acter (Figs. 77, 78, Yol. Y.). The egg-shell or chorion is, accord- 
ing to Korschelt 1 and others, a cuticular product of the epithelium 
of the egg-chambers of the ovaries. The number of eggs deposited 
by an individual differs greatly in different species, and has been 
ascertained to be variable within certain limits in the same 
species. Speyer thought about 250 to be the average number 
of eggs deposited by an individual. The number in the case of 
Aporia crataeyi is believed to be from 60 to 100, and in some 
Ifepialus to be several thousands. The mode of deposition also 

differs greatly ; where the eggs are very 
numerous they seem to be discharged almost 
at random in suitable spots ; but moths such 
as Clisiocampa ncustria fasten their eggs 
round the stems of the food-plant in a very 
perfect and artistic manner. Butterflies 
seem as a rule to prefer to oviposit by placing 
an egg here and there rather than risk many 
in one situation ; but to this there are many 
conspicuous exceptions especially in the cases 
where the larvae live gregariously, as in the 
Yanessae. Some moths cover the eggs with 
fur from their own body, which, in the case 
of certain of the Eggers (Lasiocampidae), 

FIG. 164. Egg of Orange- seems to have a special supply for the pur- 
tip butterfly, Euchlue 

cardamines, magni- pose. ilie period that intervenes between 
tied. </, The egg of deposition and hatching of the eggs varies 

natural size on a stalk. 

from a few days to many months. There 
seems to be, as a rule, comparatively little power of extending the 
period of latency beyond a single season ; though certain facts 
have been recorded that would lead us to believe that in 
Australia eggs may last over the proper time during a drought, 
and be hatched as soon as rain falls. 

Larva. The young condition or larva of the Lepidopterous 
Insect is commonly called a caterpillar. It is a somewhat 

it was sometimes called 

1 Actu. Ac. German, li. 18S7, ]>. 238. 




worm-like creature in old English 



vi LARVA 323 

palmer-worm and is composed of a head and thirteen divisions 
or segments of the body ; the first three of the latter are called 
thoracic, the other ten, abdominal segments ; in most caterpillars 
the terminal two or three abdominal segments are more or less 
run together, and the ninth may be very small, so that the true 
number is indistinct. The first three segments bear each, on 
either side, a short limb, ending in a curved spine ; the next 
two (or three or more) segments are destitute of legs, but on 
some of the following divisions another kind of leg of a more 
fleshy character appears, while the body is terminated by a pair 
of these thick legs of somewhat different form. The front legs 
are usually called the true legs, the others prolegs, but this latter 
designation is a most unfortunate one, the term " pro " being in 
entomology used to signify anterior ; it is therefore better to 
call the three anterior pairs thoracic legs, and the others abdominal 
feet, . distinguishing the hind pair of these latter as claspers. 
There is, too, an unfortunate discrepancy amongst entomologists in 
their manner of counting the body-segments, some count the head 
as the first segment, while others apply this term to the first 
thoracic segment. The latter is the more correct course, for, as 
the head is not a single segment it should not be called such in 
a terminology that affects to be morphologically exact, not simply 
descriptive. The thoracic legs are transversely jointed (Fig. 165, 
B), but this is not the case with the abdominal feet, which are 
usually armed beneath with a circle, or with rows, of little hooks. 
The thoracic legs are, independent of their form, of a different 
nature from the abdominal, for these latter disappear subsequently, 
while the former jjive rise to the les;s of the imago. The number 

o o o 

of thoracic legs is always six, except in a few cases where there 
are none at all ; the abdominal feet are much more variable, and 
exhibit so many distinctions that we cannot here attempt to 
deal with them. M. Goossens has given a concise and interest- 
ing account of this subject, 1 and Speyer - a summary of the variety 
in number and position. 

The anatomy of the larva is simple in comparison with that 
of the perfect Insect; its main features will be appreciated from 
Fig. 165, from which it will be seen that the stomach is 
enormous, and the silk-vessels are also very extensive. There 
are three sets of glands opening by canals on the head, viz. the 

1 Ann. Soc. ent. France, 1887, pp. 384-404, PI. 7. " Isis, 1845, p. 835. 



LEPIDOPTERA 



CHAP. 



salivary glands proper, which open into the cavity of the mouth, 
one close to the base of each mandible ; the silk-glands, which 
terminate by a common canal, continued externally as the spin- 
neret ; and the glands of Filippi situate in the head itself, and 
opening into the ducts of the silk-glands, near their union 
into a common duct. It should be recollected that Fig. 165 
does not indicate all the details of the anatomy ; the muscular 
system, for instance, being entirely omitted, though there are 
an enormous number of muscles ; these however are not very 
complex, they being mostly repetitions in the successive seg- 
ments. 1 The mouth -parts are very different from those of the 

4- d:j. z i 




FIG. 165. A, Section of male caterpillar of Anosia plcxippus muscular and traclieal 
systems and fat-body not shown : I, pro-, II, meso-, III, meta-thorax ; 1-10, 
abdominal segments ; b, supra -oesophageal ganglion ; c, rectum ; d.v, dorsal vessel ; 
if-ff, ganglia of ventral chain ; h, head ; /, intestine ; iti.r, Malpighian tube ; n.c, 
nerve-cord of ventral chain ; ue, oesophagus ; , spinneret ; s.o.g, infra-oesophageal 
ganglion ; nt, stomach ; s.i; silk-vessel ; t, testis. B, One of the jointed prothoracic 
legs. C, An abdominal foot with its hooks. (After Scudder and Burgess, mag- 
nification about -j.) 

perfect Insect, inasmuch as the maxillae and labial palpi, which 
are the most remarkable structures of the imago, are small, and 
are differently constructed in the caterpillar, while the mandibles, 
which are the largest organs of the caterpillar, disappear in the 
adult. The little organ by which the caterpillar exudes its silk 
is called a spinneret ; according to Packard it is a " homologue 
of the hypopharynx." It is a more or less prominent point on 
the middle of the labium (Fig. 1 66, g} and sometimes forms a 
conspicuous spine projecting downwards. The eyes are ex- 
tremely imperfect organs, consisting merely of six, in some cases 

1 For anatomy of caterpillars refer to Lyounet's famous work, Trait anatomique 

de la c/teitilh' qui range Ic lois de saule, La Have, 1762. 




vi LARVA 325 

fewer, transparent, somewhat prominent, little spaces placed on 

each side of the lower part of the head ; they are called " ocelli," 

by Landois " ocelli compositi." Under each of these external 

facets there are placed percipient 

structures, apparently very imperfect 

functionally, the caterpillar's sight 

being of the poorest character. 1 The 

spiracles of the caterpillar are nine 

on each side, placed one on the first 

thoracic segment and one on each of ^/. <-- 

/ */ 

the first eight abdominal segments ; . 

FIG. 166. Front view of head of 

there are no true stigmata on the a caterpillar, with the jaws 

second and third thoracic segments. Partially opened, a Labrum ; 

o, mandible ; c, antenna ; tf, 
though traces Of their rudiments Ol' ocelli ; e, maxilla ; /, lingua ; g, 

vestiges are sometimes visible. sphmeret ; 7 '' 1:ihial pal1 '- 

In the caterpillar there are no traces of the external sexual 
organs, so that the two sexes cannot be distinguished on super- 
ficial inspection ; it was however long ago demonstrated by 
Herold 2 that the ovaries and testes exist in the youngest cater- 
pillars, and undergo a certain amount of growth and development 
in the larval instars ; the most important feature of which is 
that the testes are originally separate but subsequently coalesce 
in the middle line of the body, and become enclosed in a common 
capsule. In a few forms especially of Liparidae (Lymaiitriidae 
of modern authors) the caterpillars are said to be of different 
colours in the two sexes. Most of what is known on this point 
has been referred to by Hatchett Jackson. 3 

The SILK -GLANDS of Lepidoptera are of great interest from the 
physiological point of view, as well as from the fact that they 
have furnished for many ages one of the most beautiful of the 
adornments made use of by our own species. The sericteria, or 
vessels that secrete silk, are of simple structure, and differ greatly 
in their size in the various forms of the Order; they sometimes 
become of great length ; in the Silk-worm each of the two 
vessels is nearly five times as long as the body, while in 
Bombyx yamamai and others, even this is exceeded. They 

1 See Plateau, Bull. Ac. Belgique, xv. 1888, p. 28 ; in reference to structure of 
ocelli, Blanc, Tete du Bombyx mori . . . 1891, pp. 163, etc. ; and Landois in 
Zcitsehr. wiss. Zool. xvi. 1866, p. 27-. 

- Entvrickelungsgeschicfite der Schmetterlinge, Cassel, 1815. 

3 Tr. Linn. Soc. London, Zool. 2nd Ser., v. 1890, pp. 147, 148. 



326 LEPIDOPTERA 



CHAP. 



grow with remarkable rapidity, being in the young silk-worm 
only 3 mm. long, in the adult 22 mm. The increase in weight is 
still more remarkable ; when the silk-worm is thirty-one days old, 
the sericteria weigh only 3 mgr., but when the age is fifty days 
their weight has increased to 541 mgr., being then | of the whole 
weight of the body. In the pupa they undergo a gradual atrophy, 
and in the moth they are, according to Helm, no longer to be 
found, though earlier authors were of a contrary opinion. 1 Ac- 
cording to Joseph," the silk-vessels begin to develop at an ex- 
tremely early age of the embryo, and are very different in their 
nature from the salivary glands, the former being derivatives of 
the external integument (ectoderm), while the salivary glands 
belong to the alimentary system. This view is to some extent con- 
firmed by the observations of Gilson as to the different manner 
in which these two sets of glands discharge their functions. 

The chief feature in the anatomy of the larva is the great 
size of the stomach. There is a very short oesophagus and crop ; 
the latter becomes enlarged, spreading out so as to form the 
stomach, a great sac occupying the larger part of the body-cavity 
(Fig. 165). On the hinder end of this sac the Malpighian tubes 
open ; they are similar in their disposition to those of the imago ; 
behind the stomach the canal expands into two successive, short 
dilatations, the first called an intestine, the second a rectum ; 
they are connected by very short isthmuses. The dorsal vessel 
is a simple, slender tube, extending from the eighth abdominal 
segment to the head. The main nervous system consists of 
supra- and infra -oesophageal ganglia, a small frontal ganglion, 
and a ventral chain of eleven ganglia, three thoracic and eight 
abdominal, the last of these latter being double. The sexual 
organs are quite rudimentary, and the passages connected with 
them very incompletely developed. 

Pupa. The pupa, which is one of the most remarkable of 
the instars of an Insect's life, attains its highest development in 
Lepidoptera. The Lepidopterous pupa is frequently called a " chry- 
salis," a term originally ;ippli'd t<> certain metallic butterfly pupae. 
The Lepidopterous pupa differs from that of other Insects in the 
fact that its outer skin forms a bard shell, all the appendages of 

1 For information as to the structure and function of the silk-vessels, refer to 
Ifrhn, Zci/sc/tr. -wiss. Zuol. xxvi. 1876, p. 434 ; and Gilson, La Cellule, vi. 1800, 
p. 116. - Jahresbcr. Schlcsisch. Gcs. Iviii. 1881, p. 116. 



VI 



PUPA 



327 



the body being glued together by an exudation so as to form a 
single continuous outer skin. This form of perfect pupa is called 
" pupa obtecta." The obtected pupa is exhibited in various stages 
of perfection in the Lepidoptera ; the maximum of perfection is 
attained by the pupae of such butterflies as are exposed without 
protection or concealment ; on the other hand, we find in various 
small moths conditions of the pupa that do not differ in any 
marked manner from the pupae of Insects of other Orders. 
Moreover, certain Coleoptera and Diptera exhibit obtected pupae 
of a more or less perfect kind. Hence the pupa obtecta is to be 
considered as a perfected condition that exists more frequently 
in the Lepidoptera than in other Orders. 

The pupa has no orifices to the alimentary canal or sexual 

FIG. 167. Section of 
female pupa of 
Anosia plexippus, 
3-4 days old. I, 
pro-, II, meso-, III, 
meta-thorax ; 1-9, 
abdominal s e g- 
ments ; a, anten- 
na(?) ; ac, aortal 
chamber ; ay 1 - <nf\ 
abdominal ganglia ; 
a g I, accessory 
glands ; ao, aorta ; 
br, brain ; c, colon ; 

cj), bursa copulatrix ; cr, cremaster ; f l , first femur ; fr, food-reservoir ; /;, dorsal vessel ; 

i, part of intestine ; MI; Malpighian tube ; mx, base of maxilla ; oe, oesophagus ; ov, ovary ; 

l)k, pharynx; sd, salivary duct; sgl, salivary gland ; st, stomach; t\ first tarsus; tg, 

compound thoracic ganglion ; ts 2 , is 3 , second and third tarsus.. (After Scudder.) 

organs, but the respiratory openings are pervious. It has no 
means of locomotion, but it can move a certain number of the 
posterior segments (the number variable according to kind). In 
some cases it is provided with spines, " adminicula," by means of 
which, aided by the wriggling movements of the abdominal seg- 
ments, considerable changes of position can be effected. The 
pupae of the genus Micropteryx apparently use the legs for 
locomotion, as do the pupae of 'Trichoptera. 

The study of the pupa of Lepidoptera is less advanced than 
that of the imago and larva, between which it is, in many points 
of structure, intermediate. 1 The interior of the pupa contains a 

1 The student will find important information as to the varieties of external 
form of pupae in Dr. T. A. Chapman's writings ; see especially Tr. cut. Soc. London, 
1893, 1894, and 1896. 




iff 



mx. 



328 



LEPIDOPTERA 



CHAP. 



quantity of cream-like matter, including the results of histolysis 
but this, as well as the condition of the internal organs, differs 
much according to whether the change from the caterpillar to 
the moth is much or little advanced. 

Many pupae are protected by cocoons. These are masses of silk 
very various in form disposed by the caterpillar around itself 
during the last stage of its existence. Some of these cocoons are 
so perfect that the moth has considerable difficulty in escaping 
when the metamorphosis is complete. Various devices are used 
for the purpose of emergence ; the Puss-moth excretes a corrosive 
fluid, containing potassium hydroxide, and then protects itself 
from this by retaining on the head while passing through it a 
shield formed of a portion of the pupa-skin. 1 Lepidopterous pupae 
usually have the body terminated by a projection of very various 
and peculiar form called " cremaster." In certain cases these 
projections are used for the suspension of the pupa, and are then 
frequently provided with hooks (Fig. 177, C, D). In other cases 
the cremaster is frequently called the anal armature (Fig. 205, B). 

The development of the wings 
of Lepidoptera has recently been 
much studied. It has been known 
since the time of Lyonnet, that 
the rudiments of the wings exist 
inside the body of the caterpillar 
when it is nearly adult. Verson 
considers that he has detected the 
rudiments in the silk-worm larva 
even before hatching, and he 
attributes their origin to a 
modification of form of those 

FlG. 168. Wing -rudiments of Pieris hypodemial cells that OCCUpy 

brassicae. A, Rmlinieuts of a wing , -, ,-, 

before the first moult of the cater" the s P ots where the Spiracles 

pillar : ce, embryonic cells ; ch, ex- o f the Second and third thor- 

ternal cuticle ; h. hypodermis ; <>. . , 

opening of the imagination; tr, acic segments might be looked 

trachea. B, posterior wing-rudiment f or- rt^ ^{\[ j^g recollected that 
of full - grown caterpillar ; b, semi- 
circular pad; c, a bundle of the there are no spiracles on these 

rolled tracheae ; e, envelope ; i, pedi- twQ thoracic segments ill Lepi- 

cel ; tr, trachea. (After Goniu.) 

dopterous larvae). Gonm has 
examined the wing-rudiments in the caterpillar, a few days old, of 

1 Latter, Tr. cut. Soc. London, 1895, p. 399. 




vi DEVELOPMENT - WINGS - NERVURES 329 

Pier is brassicae, 1 and finds that the future wing is then indi- 
cated by a thickening and bagging inwards of the hypodermis, 
and by some embryonic cells and a trachea in close relation with 
this mass (Fig. 168, A). The structure grows so as to form a sac 
projecting to the interior of the body, connected with the body- 
wall by a pedicel, and penetrated by a trachea forming branches 
consisting of rolled and contorted small tracheae (Fig. '168, B). 
If the body -wall be dissected off the caterpillar immediately 
before pupation the wings appear in crumpled form, as shown 

in Fig. 169. This fact was known 

a a 

to the older entomologists, and gave sti 
rise to the idea that the butterfly 
could be detected in a caterpillar by 
merely stripping off the integument. 

The exact mode by which the 
wings become external at the time 
of appearance of the chrysalis is not 
ascertained ; but it would appear from p" p .-" 

Goilill's Observations that it is not FIG. 169. Anterior parts of a cater- 

by a process of evagination, but by P illar of p - brassieae, the body- 

. ' wall having beeii dissected oft, 

destruction Ot the hypodermiS lying immediately before pupation. 

outside the wing. However this may a \ a '> Anterior and posterior 

wings ; st 7, first spiracle ; p, p , 
be, it is Well kllOWll that, when the second and third legs. (After 




caterpillar's skin is finally shed and 
the chrysalis appears, the wings are free, external appendages, 
and soon become fastened down to the body by an exudation 
that hardens so as to form the shell of the chrysalis. 

Scales and nervures. Before tracing the further develop- 
ment it will be well to discuss the structure of the scales and 
nervures that form such important features in the Lepidopterous 
wing. 

If a section be made of the perfect wing of a Lepidopteron, 
it is found that the two layers or walls of the wing are firmly 
held together by material irregularly arranged, in a somewhat 
columnar manner. The thickness of the wing is much greater 
where the section cuts through a nervure (Fig. 1VO, A). The 
nervures apparently differ as to the structures found in them. 
Spuler observed in a nervure of Triphaena pronuba, a body having 
in section a considerable diameter, that he considered to be a 

1 Bull. Soc. Vaudoise, xxx. 1894, No. 115. 



330 



LEPIDOPTERA 



CHAP. 



trachea, and also a " wing-rib " and blood-cells. He remarks that 
even in nervures, perfectly formed as to their chitinous parts, 
either wing-rib or trachea or both may be absent. 1 Schaffer 2 
was unable to find any tracheae in the completed wings he 
examined, and he states that the 
matrix of the tracheae and even 
their inner linings disappear. The 
wing-ribs were, however, found by 
him to be present (Fig. 170, A 
and B). 

The scales that form so con- 
spicuous a feature in Lepidoptera 
exist in surprising profusion, and 




B 





FIG. 170. Structure of wing of imago. A, 
Transverse section of basal portion of wing 
[of Vanessa ?] containing a nervure : <; 
cuticle ; ft; wing-rib ; //, wall of nervure 
(" Grunilmembran ") ; //, hypodermis ; p, 
connecting columns : r, lumen of nervure ; B, 
section of a rib ; b, one of the chitinous pro- 
jections ; sir, central rod. (After Schafl'er.) 



Fiu. 171. Scales of male Lepidoptera. 
A, Scale from upper surface of 
Eccres /<niii//t/"x ; B, from upper 
surface of Pieris m/Hic : C, from 
inner side of fold of inner margin 
of hind wing of Laertias phihnor ; 
D, one of the cover-scales from the 
costal androconinm of Eudamiis 
jirofeus ; E, F, G, scales from andro- 
coniunt of Thorybes pylades. (After 
Scudder). 



are of the most varied forms. They may be briefly described 
as delicate, chitinous bags ; in the completed state these bags 
are flattened, so as to bring the sides quite, or very nearly, 
together. Their colour is due to contained pigments, or to stria- 
tion of the exposed surface of the scale ; the latter condition 
1 ZdtKclir. n-iss. Zwl . liii. 1892, p. 623. 2 Zool. Jahrl. Annt. iii. 1889, p. (346. 



VI 



SCALES 



331 




A. 

Socket holding the stalk in Galleria 
mellonella ; B, insertion of the scale 
of Polyommatus phloeas. b, Base of 
scale ; ;, holding-ring ; w, surface of 
wing. (After Spuler.) 



giving rise to metallic " interference-colours." The walls of the 
scales are themselves, in some cases, tinted with pigment. It is 
said that some of the scales contain air, and that the glistening 

O O 

whiteness of certain scales is due to this. The exposed surface of 

the scale usually differs from the 

surface that is pressed down on 

the wing in being delicately and 

regularly striated ; the colours 

of the upper and under surfaces 

of a scale may also be quite 

different. Scales are essentially 

of the nature of hairs, and all 

the transitions between hairs and F IG> 179. insertion of scales. 

true scales may be found on the 

wings of certain Lepidoptera that 

baar both hairs and scales, e.g. 

Ithomia. It has been calculated 

that there are a million and a half of scales on the wings of an 

individual of the genus Moiylio. The scales are arranged on 

the wing in an overlapping manner, somewhat like slates on the 

roof of a house. Each scale has a short stalk, and is maintained 

in position by the stalk fitting into a cavity in a projection of 

the wing-membrane (Fig. 172). 

Androconia. - -The males of numerous butterflies possess 
scales peculiar in kind and various in arrangement. They may 
be either irregularly scattered over the w r ing, or they may form 
very complex definite structures (Fig. 173). They were formerly 
called " plumules," but Scudder has replaced this name by the 
better one, " androconia." The function of the androconia is 
still obscure. An odour is believed to be connected with them. 
Thomas supposes l that these scales are hollow tubes in connec- 
tion with glands at their bases, and that matter secreted by the 
glands passes through the scales and becomes diffused. In 
nearly all Lepidoptera it is the male that seeks the female ; if 
therefore odorous scales were present in one sex only we should 
have supposed that this would have been the female rather than 
the male. As, however, the reverse is the case, the function of 
the androconia is supposed to be that of charming the female. 
Scudder considers that the covering part of the androconial 

1 Amer. Natural., xxvii. 1893, p. 1018. 



332 



LEPIDOPTERA 



CHAP. 



structures is sometimes ornamental. As a rule, however, the 
' brands " of male Lepidoptera detract from their beauty to 
our eyes. 




FIG. 173. A, section of part of wing showing the complex androconia of Thanaos tages, 
a Skipper butterfly. The turning over of the costal margin of the wing is in this 
case part of the arrangement. , Upper covering-scales attached to the costal 
portion of the under surface of the wing ; b, edge of costal margin of the wing ; 
c, costal nervure with its scales ; d, field of the wing next the costal nervure, bearing 
stunted scales ; e, the androconia proper, or male scales ; /, posterior covering 
scales ; g, lumen of the costal nervure : B, a portion of the costal area flattened 
out and seen from above ; lettering as before : C, section of audroconium on the 
second nervure of Argynnis paphia. (After Aurivillius.) 

Resuming our consideration of the development of the wings, 
we may remark that the history of the changes during the 
pupal state is still imperfect. By the changes of relative size 
of the thoracic segments the hind wing is brought to lie under 
the anterior one (i.e. between it and the body), so that in the 
newly formed pupa the arrangement is that shown in Fig. 1 74. 
The wings are two sacs filled with material surrounding peri- 
tracheal spaces in which run tracheae. The subsequent history 
of the tracheae is very obscure, and contrary opinions have been 
expressed as to their growth and disappearance. We have 
alluded to the fact that in some nervures tracheae are present, 
while in others they are absent; so that it is quite possible that 



VI 



WING-DEVELOPMENT 



ao, 



the histories of the formation of the nervures and of their 
relation to tracheae are different in various Lepidoptera. This 
conclusion is rendered more probable by the statement of Corn- 
stock and JSTeedharn, 1 that in some Insects the " peritracheal 
spaces " that mark out the position of 
the future nervures are destitute of 
tracheae. Gonin thinks the nervures 
are derived from the sheaths of the 
peritracheal spaces, and a review of all 
the facts suggests that the tracheae 
have only a secondary relation to the 
nervures, and that the view that a 
study of the pupal tracheae may be 
looked on as a study of the pre- 
liminary state of the nervures is not 
sufficiently exact. It is, however, 
probable that in Lepidoptera the 
pupal tracheae play an important 
though not a primary part in the 
formation of the nervures ; possibly 
this may be by setting up changes in 
the cells near them by means of the 
air they supply. Semper long ago 
discovered hypodermal cylinders tra- 
versed by a string (Fig. 170, B), 
placed near the tracheae in the 
pupa, 2 It appears probable that the "wing -ribs" found in 
the nervures (Fig. 170, Afr and B) are the final state of these 
cylinders, but the origin and import of the cylinders are still 
unknown. 

The formation of the scales of the wing commences very 
early apparently soon after the casting of the larval skin- 
though the completion of the scales and their pigmentation is 
delayed to a late period of the pupal life. The scales are formed 
by special cells of the hypodermis that are placed deeper in the 
interior of 'the wing than the other hypodermal cells. Each 
scale is formed by one cell, and protrudes through the over- 
lying hypodermis ; the membrane into which the scales are 
inserted is a subsequently developed structure, and the beautiful 

1 Aincr. Natural., xxxii. 1898. p. 256. - Zeitschr. u-iss. Zool. viii. 1857, p. 326. 




chrysalis of Pieris brassicae, 
showing the position and struc- 
ture of the wings, hanging 
from one side of the body. 
aa, Anterior wing ; up, pos- 
terior wing ; e, e, peritracheal 
spaces ; t, t, tracheae. (After 
Gouin.) 



LEPIDOPTERA 



CHAP. 



articulation of the scale with the wing takes place by a division 
of the stalk of the scale where it is encompassed by the mem- 
brane. Semper was not able to show that the scale-forming 
cells are certainly hypodermal cells, but this has since been 
demonstrated by Schaffer, who also shows that each of the cells 
contains an excretory vesicle. 

Very little is positively known as to the development of the 
colour in the wing-scales. It has been pointed out by Hopkins l 
that in some cases the colours are of the nature of unites ; that 
is, of excretory matter of the kind that usually passes from the 
body by direct channels, and in the case of Lepidoptera, by the 
Malpighian tubes. Miss IsTewbigin suggests that the organic 
pigments used in scale -coloration will be found to be of two 



"i;L B 




FIG. 175. Early condition of scales and nervures. (After Semper.) A, Section of 
portion of \viug of pupa of N//////M: /li/n/xtr/ ; n, basal membrane with trachea 
beneath it ; c, scale-forming cell ; </, early state of a scale ; e, e, more advanced 
stages ; /, hypodermal cells. B, part of a cellular cylinder that excretes the 
nervure [or more probably the rib or " Rippe " of Schaffer; cf. Fig. 170, B] ; b. 
epithelial [hypodermal] cells ; a, central string [supposed by Semper to be a nerve]. 

kinds, urates and melanins, the urates being derivatives from nitro- 
genous, the melanins from carbonaceous, matters. 2 Marchal, who 
has devoted a great deal of attention to the study of the Malpi- 
ghian tubes, informs us that the siibdermal pigments of cater- 
pillars are fretpiiently in large part deposits of urates, and he is 
of opinion that, the function of the Malpighian tubes being 
arrested at certain periods of the metamorphosis, elimination of 
the matter they separate when functionally active then takes 
place in a variety of other ways. 3 A similar condition as to 
the melanin-pigments and the respiratory functions appears also 

1 Pit II Trans. 186 B, 1896, No. 15. - Xatwal Science, viii. 1896, p. 94. 

s Bull. Soc. ent. France, 1896, p. 257. 



vi WING-PATTERN SENSITIVENESS 335 

probable. The scales when first formed are pallid, and the physi- 
ology of their pigmentation is not fully ascertained ; it is, how- 
ever, known that when the scales are pallid the hypodermis is 
either pigmented or in close contact with pigmentary matter, 
and that as the scales become coloured this pigmentation of the 
hypodermis diminishes ; so that it is clear that the colour of 
the scales is obtained from matter in the interior of the develop- 
ing wing, and probably by the agency of the hypodermis. 

The pattern on the wings of Lepidoptera is formed before 
the emergence from the pupa. In the Tortoiseshell butterfly, 
according to Schaffer, it commences to appear about the ninth 
day of the pupal life, and the pattern is completed about the 
eleventh or twelfth day. He also states that the process varies 
in its rapidity, and this, he thinks, may depend on the previous 
condition of the larva. According to Buckell the pupa of 
Ncmeoliius lucina is sufficiently transparent to allow the develop- 
ment of the colour of the imago to be watched. He says that 
the coloration occurred first in front; that its entire production 
occupied less than twenty-fonr hours, and only commenced about 
forty-eight hours before the imago emerged. 1 When the butter- 
fly leaves the pupal skin the wings are soft, crumpled sacs, of 
comparatively small size, but, as everyone knows, they rapidly 
expand and become rigid ; the physiology of this process is 
apparently still unknown. 

A great deal of evidence, both direct and indirect, has 
accumulated showing that the organisation of many Lepidoptera 
is excessively sensitive, so that slight changes of condition pro- 
duce remarkable results ; and it has also been shown that in the 
early part of the life this sensitiveness is especially great at the 
period of ecdysis. Numerous butterflies produce more than one 
generation a year, and sometimes the generations are so different 
that they have passed current with entomologists as distinct 
species. The phenomena of this character are styled " seasonal 
variations " or " seasonal dimorphism." It has, however, been 
shown that, by careful management, the eggs of a generation (say 
form .) may be made to produce form a, whereas in the usual 
course of nature they would produce form 1>. A very remarkable 
condition is exhibited by the North American Papilio ajax. There 
are three forms of the species, known as P. ajax, P. telamonides, 

1 Ent. Record, vi. 1895, p. 258. 



336 LEPIDOPTERA CHAP. 

and P. marcellus. It is uncertain how many generations there 
may be in one year of this species, as the length of the life- 
cycle varies greatly according to circumstances. But in West 
Virginia all the butterflies of this species that emerge from the 
chrysalis before the middle of April are the form marcellus ; 
those produced between the middle of April and the end of May 
are telamonides ; while those that appear after this are ajax. 
P. telamonides is not, however, the offspring of marcellus, for both 
forms emerge from pupae that have passed through the winter 
(and are the offspring of ajax'), those that emerge early being 
marcellus, those that appear later telamonides. 

In various parts of Asia and Africa the butterflies produced 
during the wet season differ more or less markedly from those of 
the same species produced during the dry season. These are 
called " wet " and " dry season " forms. Their aetiology has not 
been investigated, this discovery being comparatively recent. 

Turning to the early life we find that some larvae vary in 
colour, and that this variation is sometimes of a definite char- 
acter, the larva being one of two different colours green or 
brown. In other cases the variation of the species is less 
definitely dimorphic, a considerable range of variation being 
exhibited by the species. In tracing the life-histories of Lepi- 
dopterous larvae it is not rare to find species in which the larva 
abruptly changes its form and colour in the middle of its life, 
and so completely that no one would believe the identity of the 
individual in the two successive conditions had it not been 
shown by direct observation ; in these cases the change in 
appearance is usually associated with a change in habits, the 
larva being, perhaps, a miner in leaves in its first stages, and an 
external feeder subsequently. In the case of the larval variation 
we have alluded to above, it is understood that there is no 
marked change of habits. 1'oulton has shown * that it is not 
infrequent for some of these latter kinds of variable larvae to 
change colour during life, and he considers that light or conditions 
of illumination, that he speaks of as " phytoscopic," are the in- 
ducing causes. Great difference is, however, exhibited according 
to species, some variable species not being so amenable to these 
influences as others are. In dimorphic forms the change was 
observed to take place at a moult, the larva changing its skin 

1 Tnuts. cut. SOL: London, 1892, p. 293, etc. 



vt SENSITIVENESS MIMICRY 337 

and appearing of another colour. In some cases the result of 
the change was to bring the colour of the larva into harmony 
with its surroundings, but in others it was not so. During the 
final stage many larvae are susceptible, the result being made 
evident only when the pupa is disclosed. Variably coloured 
pupae of certain species of butterflies have long been know r n, and 
it has been shown that some of the varieties can be induced by 
changing the surroundings. The result of the changes is in 
certain cases correspondence between the colour of the individual 
and its surroundings. In the case of other species having pupae 
of variable colour, the colour of the pupa is without relation to, 
or harmony with, the surroundings. 

Experiments have been made on pupae by Merrifield and 
others, with the result of showing that by changes of tempera- 
ture applied at certain moments some of the colours or marks of 
the butterfly that will emerge can be altered. 

It is found that in certain localities the colour of various 
kinds of butterflies more or less agrees, while it differs from 
that of the same butterflies found in other localities. Thus 
Weir speaks of a duskiness common to various butterflies in 
Java, and calls it "phaeism" ; and Bates states that in the Amazon 
valley numerous species of butterflies vary in a similar manner, 
as regards colour, in a locality. This phenomenon is now called 
" homoeochromatism," and is supposed to be due to the effect of 
local conditions on a susceptible organisation, though there is no 
experimental evidence of this. 

Mimicry. There are many cases in Lepidoptera of species 
that depart more or less strongly in appearance from those forms 
to which they are considered to be allied, and at the same time 
resemble more or less closely species to which they are less allied. 
This phenomenon is called mimicry. 1 Usually the resembling 
forms are actually associated during life. Bates, w r ho observed 
this phenomenon in the Amazon valley, thought that it might 
be accounted for by the advantage resulting to the exceptionally 
coloured forms from the resemblance ; 2 it being assumed that 
these were unprotected, while the forms they resembled were 

1 The term mimicry is sometimes used in a wider sense ; but we think it better 
to limit it to its original meaning. The word is a most unfortunate one, being 
both inadequate and inaccurate. 

' Trans. Linn, Soc. xxiii. 1862, p. 507. 
VOL. VI 



338 LEPIDOPTERA CHAP. 

believed to be specially protected by nauseous odours or taste. 
It was, in fact, thought that the destroying enemies were 
deceived by the resemblance into supposing that the forms that 
were in reality edible were inedible. This subject has been 
greatly discussed, and in the course of the discussion numerous 
cases that could not be accounted for by Bates's hypothesis have 
been revealed. One of these is the fact that resemblances of the 
kind alluded to very frequently occur amongst inedible forms. 
This also has been thought to be accounted for by a supposed 
advantage to the Insects ; it being argued that a certain number 
of " protected " forms are destroyed by enemies the instincts of 
which are faulty, and which therefore always require to learn by 
individual experience that a certain sort of colour is associated 
with a nasty taste. The next step of the argument is that it 
will be an advantage to a protected butterfly to form part of 
a large association of forms having one coloration, because 
the ignorant enemies will more easily learn the association of a 
certain form of coloration with nastiness ; moreover such destruc- 
tion as does occur will be distributed over a larger number of 
species, so that each species of a large, similarly coloured, inedible 
association will have a less number of its individuals destroyed. 
It is scarcely a matter for surprise that many naturalists are 
very sceptical as to these explanations ; especially as the pheno- 
mena are supposed to have occurred in the past, so that they 
cannot be directly verified or disproved. It has not, however, 
been found, as a matter of fact, that even unprotected butterflies 
are much destroyed in the perfect state by birds. Moreover, in 
endeavouring to realise the steps of the process of development 
of the resemblance, we meet with the difficulty that the amount 
of resemblance to the model that is assumed to be efficient at 
one step of the development, and to bring safety, is at the next 
step supposed to be inefficient and to involve destruction. In 
other words, while analysis of the explanation shows that it 
postulates a peculiar and well-directed discriminative power, 
and a persistent selection on the part of the birds, observation 
leads to the belief that birds have been but little concerned in 
the matter. If we add to this that there is no sufficient evidence 
that the species now similar were ever dissimilar (as it is sup- 
posed they were by the advocates of the hypothesis), we think 
it is clear that the explanation from our point of view is of but 



vi MIMICRY CLASSIFICATION 339 

little importance. 1 The comparatively simple, hypothetical 
explanation, originally promulgated by Bates, is sometimes called 
Batesian mimicry ; while the " inedible association " hypothesis 
is termed Miillerian mimicry. 

There is one branch of the subject of mimicry that we think 
of great interest. This is the resemblance between Insects of 
different Orders ; or between Insects of the same Order, but be- 
longing to groups that are essentially different in form and 
appearance. It is not infrequent for beetles to resemble Hymen- 
optera, and it is still more frequent for Lepidoptera to resemble 
Hymenoptera, and that not only in colour and form, but also in 
movements and attitude. Druce says : " Many of the species of 
Zygaenidae are the most wonderful of all the moths ; in some 
cases they so closely resemble Hymenoptera that at first sight it 
is almost impossible to determine to which Order they belong." 
W. Miiller says: "The little Lepidoptera of the family Glaucopides, 
that are so like certain wasps as to completely deceive us, have 
when alive exactly the same manner of holding their wings, the 
same restless movements, the same irregular flight as a wasp." 
Seitz and others record a case in which a Brazilian Macroglossa 
exactly resembles a humming-bird, in company with which it 
flies ; and the same naturalist also tells us 4 of a Skipper butterfly 
that greatly resembles a grasshopper of the genus Tettix, and that 
moreover makes movements like the jumping of grasshoppers. 
In most of these cases the probabilities of either original 
similarity, arrested evolution, or the action of similar conditions 
are excluded : and the hypothesis of the influence, by some means 
or other, of one organism on another is strongly suggested. 

The classification of Lepidoptera was said by Latreille a 
century ago to be a reproach to entomologists. Since that time 
an enormous number of new species and genera have been 
described, but only recently has much advance been made in 

1 A summary of the chief aspects of the question is contained in Beddard's 
Animal Coloration, London, 1892. An account of the subject with numerous illus- 
trations has been given by Haase, " Untersuchungen iiber die Mimicry," Bill. 
ZonJ. iii. 1893, Heft viii. Those who wish to see the case as stated by an advocate 
may refer to Professor Poulton's work, The Colours of Animals (International 
Scientific Series), Ixviii. London, 1890. 

- P. Zool. Sue. London, 1883, p. 372. 

3 Kosmos, xix. 1886, p. 353. The Insects alluded to by both these naturalists 
are now, we believe, placed in the Family Syntomidae (see p. 388). 

4 Stctt. cut. Zdt. li. 1891, p. 264 ; and Ivi. 1895, p. 234. 



340 



LEPIDOPTERA 



CHAP. 





the way of improvement of classification. The progress made 
has been limited to a better comprehension and definition of the 
families. The nervuration of the wings is the character most 
in vogue for this purpose. As regards the larger groups, and 
Phylogeny, there is a general opinion prevalent to the effect that 
Micropterygidae, Eriocephalidae and Hepialidae are in a com- 
paratively primitive condition, but as to the relations of these 
families one with the other, or with other Lepidoptera, there is a 
wide difference of opinion. 

The primary divisions of the family most often met with in 
literature are : either Ehopalocera ( = butterflies^ and Hetero- 



FlG. 176. Clubs of butterflies' 
antennae. Terminal portions 
of antenna of, 1, Pier is bras- 

siaif ; '_', >'///.! infernalis ; 3, 
Hetstiu ill en (sub-family Dan- 
aicle.s) ; 4, Eudamus proteus, 
ami 5, Lii/HH-lini-i'x tinnnas 
(Hesperiidae). (Alter Bchatz 
and Sc mliler.) 



cera ( = moths.) ; or Macrolepidoptera and Microlepidoptera ; 
the Macrolepidoptera including the butterflies and large moths, 
the Microlepidoptera being limited to the families Tineidae 
(now itself in process of division into numerous families) and 
Tortricidae ; some entomologists including also Pvr;ilidac, Ptero- 
phoridae and Orneodidae in Microlepidoptera. The division of all 
Lepidoptera into two series is merely a temporary device necessi- 
tated by imperfect acquaintance with morphology. The division 
into Macro- and Micro- lepidoptera is entirely uuscu'iitilir. 

Series 1. Rhopalocera or Butterflies. Antennae knobbed at the tip or 
thickened a little before the tip, without pectinations, projecting 
processes, or conspicuous arrangements of cilia. Hind wings with- 
out a frenulum, but with the costal nervure strongly curved at the 
base (Fig. 161, II, B). 

Series II. 7/r/rrwrm or Cloths. Antennae various in form, only rarely 
knobbed at the tip, and in such cases a frenulum present. In the 
large majority a riviiulimi is present, and the costal nervure of the 
hind-wing is either but little a relied at the base (as in Fig. KM. 
I, B) or it, has a large area between it and the front margin; 
but in certain families the hind wing is formed much as in 
Rhopalocera. 



vi BUTTERFLIES 34! 

It may be inferred from these definitions that the distinc- 
tion between the two snb-Orders is neither sharply defined nor of 
great importance. The club of the antenna of the Rhopalocera 
exhibits considerable variety in form (Fig. 17G). 1 Butterflies 
are as a rule diurnal in their activity and moths nocturnal ; but 
in the tropics there are numerous Heterocera that are diurnal, 
and many of these resemble butterflies not only in colour but 
even in the shapes of their wings. 



Series I. Rhopalocera. Butterflies. 

Classification and Families of Butterflies. Although 
considerable unanimity exists as to the natural groups of butter- 
flies, there is much diversity of opinion as to what divisions are 
of equivalent value some treating as sub-families groups that 
others call families and as to the way the families should be 
combined. There is, however, a general agreement that the 
Hesperiidae are the most distinct of the families, and E. Renter 
considers them a distinct sub-Order with the name Grypocera. 2 

Four categories may be readily distinguished, as follows, 
viz. : 

1. The majority of butterflies ; having the first pair of legs more or less strik- 

ingly different from the other pairs ; frequently very much smaller 
and not used as legs ; when not very small, then differing according 
to sex of the same species, being smaller in the male than in 
the female ; the part most peculiar is the tarsus, which is modified 
in various manners, but in the males of this great series is always 
destitute of its natural form of a succession of simple joints five in 
number. There is no pad on the front tibia, 

Fam. NYMPHALIDAE, ERYCINIDAE, LYCAENIDAE. 
[The distinctions between these three families are found in the 
amount and kind of the abortion of the front legs ; for 
definition refer to the heading of each of the families.] 

2. The front legs are in general form like the other pairs ; their tibiae 

have no pads; the claws of all the feet are bifid, and there is an 
empodiuui in connection with them. Fam. PIERIDAE. 

1 For an account of the antenna of butterflies, see Jordan, J\~oi: Zool. v. 1898, 
pp. 37-1-415. 

2 Haase first proposed the name Netrocera (Deutsclie cut. Zeit. Lcp. iv. 1891, 
p. 1) for Hesperiidae, as a division distinct from all other butterflies ; Karsch 
replaced the name in the following year by Grypoceva, because Netrocera is the 
name of a genus. 



34 2 LEPIDOPTERA CHAP. 

3. The front legs are like the other pairs ; their tibiae however possess 

pads ; the claws are large, not bifid, and there is no empodium ; 
the metanotum is completely exposed at the base of the abdomen. 

Fam. PAPILIONIDAE. 

4. The front legs are like the other pairs ; their tibiae however possess 

pads ; the claws are small, toothed at the base, and there is an 
empodium ; the metanotum is concealed by the prolonged and 
overhanging mesonotum. Fam. HESPERIIDAE. 

The relations between the families Erycinidae, Lycaenidae, 
and Nymphalidae are very intimate. All these have the front 
legs more or less modified, and the distinctions between the 
families depend almost entirely on generalisations as to these 
modifications. These facts have led Scudder to associate the 
Lycaenidae and Erycinidae in one group, which he terms 
' Eurales." It is however difficult to go so far and no farther : 
for the relations between both divisions of Eurales and the 
Nymphalidae are considerable. "NVe shall subsequently find that 
the genus Libytliea is by many retained as a separate family, 
chiefly because it is difficult to decide whether it should be 
placed in Erycinidae or in Nymphalidae. Hence it is difficult to 
see in this enormous complex of seven or eight thousand species 
more than a single great Nymphalo-Lycaenid alliance. The 
forms really cognate in the three families are however so few, 
and the number of species in the whole is so very large, that it 
is a matter of great convenience in practice to keep the three 
families apart. It is sufficient for larger purposes to bear in 
mind their intimate connexions. 

The Papilionidae and Pieridae are treated by many as two 
sub-divisions of one group. But we have not been able to find 
any justification for this in the existence of forms with connect- 
ing characters. Indeed it would, from this point of view, 
appear that the Pieridae are more closely connected with the 
Lycaenidae and Erycinidae than they are with Papilionidae ; 
in one important character, the absence of the pad of the front 
tibia, the Nyinphalo-Lycaenids and the Pierids agree. It has 
also been frequently suggested that the Papilionidae (in the 
larger sense just mentioned) might be associated with the 
Hesperiidae. But no satisfactory links have been brought to 
light; and if one of the more lowly Hesj>eriids, such as Tlianaos, 
be compared with one of the lower Papilionidae, such as 
Parnassius, very little approximation can be perceived. 



vi BUTTERFLIES 343 

It appears, therefore, at present that Hesperiidae, Papi- 
lionidae, Pieridae, and the Nymphalo-Lycaenid complex are 
naturally distinct. But in the following review of the families 
and sub-families of butterflies, we shall, in accordance with the 
views of the majority of Lepidopterists, treat the Lycaenidae 
and Erycinidae as families distinct from both Nymphalidae and 
Pieridae. 1 

The number of described species of butterflies is probably 
about 13,000; but the list is at present far from complete; 
forms of the largest size and most striking appearance being still 
occasionally discovered. Forty years ago the number known 
was not more than one-third or one-fourth of what it is at 
present, and a crowd of novelties of the less conspicuous kinds is 
brought to light every year. Hence it is not too much to antici- 
pate that 30,000, or even 40,000 forms may be acquired if 
entomologists continue' to seek them with the enthusiasm and 
industry that have been manifested of late. On the other 
hand, the species of Rhopalocera seem to be peculiarly liable to 
dimorphic, to seasonal and to local variation ; so that it is 
possible that ultimately the number of true species that is, 
forms that do not breed together actually or by means of inter- 
mediates, morphological or chronological may have to be con- 
siderably reduced. 

In Britain we have a list of only sixty-eight native butter- 
flies, and some even of these are things of the past, \vhile others 
are only too certainly disappearing. New Zealand is still 
poorer, possessing only eighteen ; and this number will prob- 
ably be but little increased by future discoveries. South 
America is the richest part of the world, and Wallace informs us 
that 600 species of butterflies could, forty years ago, be found 
in the environs of the city of Para. 

Fam. 1. Nymphalidae. The front pair of legs much reduced 
in size in each sex, their tarsi in the male with but one joint, 

' The literature of butterflies has become extremely extensive. The following 
works contain information as to general questions : 1, Scudder's Butterflies of Neio 
England, a beautifully illustrated work completed in 1889, and replete with 
interesting discussions. 2, Staudinger, Schatz and Rober, Exotischc Tag/alter, 
in three folio volumes (Fiirth, 1884-1887), with illustrations of exotic butterflies 
and a detailed sketch of their characters. 3, Enzio Renter, "Tiber die Palpen 
der Rhopaloceren," in Acta Soc. Sci. Fenn. xxii. 1896, treating fully of classifica- 
tion and phylogeny. 



344 



LEriDOPTERA 



CHAP. 



though in the female there are usually five hut without any claws. 
Pupa xiixjii'iided l>y the tail so as to hang down freely. AVe 
include in this family several sub -families treated by some 
taxonomists as families ; in this respect we follow Bates, whose 
arrangement l still remains the basis of butterfly classification. 
With this extension the Nymphalidae is the most important of 
the families of butterflies, and includes upwards of 250 genera, 
and between 4000 and 5000 species. There are eight sub- 
families. 

It is in Nymphalidae that the act of pupation reaches its 
acme of complication and perfection ; the pupae hang suspended 
by the tail, and the cremaster, that is the process at the end 

of the body, bears highly- 
developed hooks (Fig. 177, 
C, D). The variety in 
form of the chrysalids is 
extraordinary; humps or 
processes often project 
from the body, making 
the Insect a fantastic 
object ; the strange ap- 
pearance is frequently in- 

FIG. 177. Pupa of the Purple Emperor butterfly, Creased by patches like 
Apatura in New Forest. A, Lateral, B, M ^ j j 

dorsal aspect ; C, enlarged view of cremaster 

with the suspensory hook ; D, one hook still Various parts of the body, 

more enlarged. -r, IT j -i ,-1 

It is believed that the 




I 




term chrysalid was first suggested by these golden pupae. 
The Purple Emperor, Apatura iris, differs strikingly in the pupa- 
as well as in the larva- stage from all our other Nymphalids ; it 
is of green colour, very broad along the sides, but narrow on 
the dorsal and ventral aspects (Pig. 177). The skin of this 
pupa is less hard than usual, and the pupa seems to be of a 
very delicate constitution. The Purple Emperor, like some of 
the Satyrides as well as some of its more immediate congeners, 
hibernates in our climate as a partially grown larva and passes 
consequently only a very brief period of its existence in the 
form of a pupa. 

Sub-Fam. 1. Danaides. Front wing with inner-margin 

1 Journal of Entomology, i. 1862, p. 218 : for early instars of South American 
Nymphalidae sue JM tiller, Zoo 1. Jahrl. Syst. i. 1886, p. 117. 



vi RHOPALOCERA DANAIDES 345 



(submedian) nervure, with a short fork at the base. Cell of hind 
i/'iug dosed. Front foot of the female ending in a corrugate 
knob. Caterpillars smooth, provided with a feio long fie shy pro- 
cesses. The claws are in a variable state, being sometimes simple, 
as in Papilionidae, sometimes with an empodium, apparently of 
an imperfect kind. The Danaides are usually large Insects with 
an imperfect style of ornament and colour ; they have a great 
deal of black or very dark scaling, and in some Euploea this 
is agreeably relieved by a violet or purple suffusion, and these 
are really fine Insects. Usually there are large pale spaces, of 
some neutral indefinite tint, on which black blotches are dis- 
tributed in a striking but inartistic manner. In many of the 
species the markings are almost spot for spot the same on the 
upper and under sides. About seven genera and 250 species are 
recognised. Danaides occur in all the warmer parts of the 
world, but are most numerous in the Eastern tropics. In Europe 
the family is represented only by an Asiatic and African species, 
Limnas chrysippus, that has extended its range to Greece. 
Besides this another species, Anosia erippus, Or. (unfortunately 
also called Anosia menippe, Hb., and Danais archippus or even I). 
plexippus) has in the last two or three decades extended its 
range to various islands and distant localities, concomitantly, it 
is believed, with an extension of the distribution of its food-plant, 
Asdepias. This Insect has several times been taken in this 
country, and may probably be a natural immigrant. It is a 
common butterfly in North America, where it is called the 
Monarch. 1 

Some, at least, of the Danaides are unpleasant to birds in 
odour or in taste, or both. Among them there occur, according 
to Moore 2 and others, numerous cases of resemblance between 
forms that are thus protected. It is possible that the odour 
and taste are of some value to the Insects ; 3 as, however, butter- 
flies of any kind appear to be but rarely attacked in the imago- 
state by birds, and as their chief enemies are parasitic Insects 
that attack the larval instar, it is impossible to consider this 
protection of such prime importance to the species as many 
theorists assume it to be. 

1 This is the subject of Scudder's Life of a Butterfly, 1893. 

2 P. Zoul. Soc. London, 1883, p. 205. 

3 Finn, J. Asiat. Soc. Bengal, Ixvi. 1896, p. 528 ; Ixvii. 1897, p. 213. 




346 LEPIDOPTERA CHAP. 

Sub-Fam. 2. Ithomiides. Differs from. Danaides ly the 
female front foot having a true, though somewhat abbreviate 
fi'/'sus. The caterpiUcrs have no long processes. There has been 
considerable difference of opinion as to this division of butter- 
flies. It is the family Neotropidae of Schatz, the Mechanitidae 
of Berg ; also the " Danaioid Heliconiidae " of several previous 
writers, except that Itunci and Lycorea do not belong here 
but to Danaides. Godman and Salvin treat it as a group 
of the Danaid sub -family. The Ithomiides are peculiar to 
tropical America, where some 20 or 30 genera and about 500 

species have been discovered. 
^/ ^-^^ There is considerable variety 
amongst them. Ithomia and 
Hy in i a if is are remarkable for the 
small area of their wings, which 
bear remarkably few scales, these 

ornaments being in many cases 
FIG. 1/S. Itfiomia pusio. Brazil. 

limited to narrow bands along 

the margins of the wings, and a mark extending along the 
discocellular nervule. Wallace says they prefer the shades of 
the forest and flit, almost invisible, among the dark foliage. 
Many of these species have the hind-wings differently veined 
in the two sexes on the anterior part, in connection with the 
existence in the male of peculiar fine hairs, placed near the 
costal and subcostal veins. Tithorea and other forms are, how- 
ever, heavily scaled insects of stronger build, their colours usually 
being black, tawny-red or brown, yellow, and white. In the 
sub-fam. Danaides, according to Fritz Miiller, the male has scent- 
tufts at the extremity of the abdomen, whereas in Ithomiides 
analogous structures exist on the upper side of the hind-wing. 
Ithomiides have various colour-resemblances with members of 
the Heliconiides and Pieridae ; Tithorea has colour analogues in 
Heliconius, and Ithomia in Dismorpliia (formerly called Leptalis). 
Crowds of individuals of certain species of Ithomia are occasion- 
ally met with, and mixed with them there are found a small 
number of examples of Dismr/>// ia coloured like thrmst-hrs. 
They arc placed by Haase in his category of secondary models. 
Belt states that some Ithomiides are distasteful to monkeys and 
spiders, but are destroyed by Fossorial Hymenoptera, which use 
the butterflies as food for their young ; and he also says that 



vi RHOPALOCERA 1THOMIIDES SATYRIDES 347 

they are very wary when the wasp is near, and rise off their 
perches into the air, as if aware that the wasp will not then 
endeavour to seize them. " Much information is given about 
the habits by Bates in the paper in which he first propounded 
the " theory of mimicry." The larvae are said to live on 
Solanaceae. 

The genus Hamadryas is placed by some writers in Danaides, 
by others in Ithomiides ; and Haase has proposed to make it the 
group " Palaeotropinae." The species are small, black and white 
Insects, somewhat like Pierids. They are apparently hardy 
Insects, and are abundant in certain parts of the Austro-Malay 



region. 



Sub-Fam. 3. Satyrides. Palpi strongly pressed tor/ether, set 
in front with long, stiff hairs. Front wings frequently with one 
or more of the nervures swollen or 'bladder-like at the base of the 
wing. Cells of both wings closed. Caterpillar thickest at the 
middle, the hind end of the body bijid. Pupa generally suspended 
by the cretnastcr, without girth: but sometimes terrestrial. This 
is a very extensive group, consisting of upwards of 1000 species. 
The Insects are usually of small size, of various shades of brown 
or greyish colours, with circular or ringed marks on the under 
sides of the wings. It is found all over the world, and is well 
represented in Europe; our Meadow-browns, Heaths, and Marbled- 
whites, as well as the great genus Erelria of the highlands and 
mountains belonging to it. Most of these Insects have but 
feeble powers of flight, and rise but little from the surface of the 
ground. The caterpillars live on various grasses. They are 
usually green or brown, destitute of armature, and a good deal 
like the caterpillars of Noctuid moths, but the hind end of the 
body is thinner and divided to form tw T o corners, while the head 
is more or less free, or outstanding. The pupae are of great 
interest, inasmuch as in a few cases they do not suspend them- 
selves in any way, but lie on the ground ; sometimes in a very 
feeble cocoon or cell. There are no cremasteral hooks. The 
pupae of the Grayling butterfly, Hipparchia semelc, has been 
found in loose soil a quarter of an inch below the surface. The 
chrysalis of the Scotch Argus, Erebia aethiops, was found by 
Mr. Buckler to be neither suspended nor attached, but placed 
in a perpendicular position, head upwards, amongst the grass. 

1 Trans. Linn. Soc. xxiii, 1862, p. 495. 



348 LEPIDOPTERA CHAP. 

In the majority of cases the pupa is, however, suspended 
as is usual in Nymphalidae. Nothing is known as to the 
nature of the peculiar inflation of the bases of the nervures of 
the front wings ; it is well shown in our common species of 
Coenonympha] this character is not, however, constant through- 
out the family. There is in South America a very remarkable 
group of Satyrides consisting of the genera Cithaerias and 
Haetcm, in which the wings are very delicate and transparent, 
bearing on the greater part of their area remote fine hairs instead 
of scales ; there are nevertheless some scaled patches about the 
margins, and one or more of the ringed marks characteristic of 
the Satyrides ; while in some species the distal portions of the 
hind wings are tinted with carmine, The species of the genus 
Pierella connect these transparent Satyrids with the more 
ordinary forms. According to Wallace the habits of these 
fairy-like forms are those characteristic of the family in general. 
The genus Elymnias has been separated by some authorities ;is 
a sub-family, or even as a family, Elymniidae, chiefly on the 
ground of a slight peculiarity in the termination of the branches 
of the veins at the outer angle of the front wings. The Elymnias 
are said to be of a mimetic nature, having a greater or less 
resemblance to butterflies of various other divisions ; there is 
also a considerable difference in appearance between their own 
sexes. The larva of E. undularis is known ; it is of the form 
usual in Satyrides, and lives on the palm Corypha. About 
50 species, ranging from India to Australia, with two in Africa, 
are known of this interesting group. 

Sub-Fam. 4. Morphides. There is no cell on the hind ?r/////, 
the discocellular ncrrulc Icing absent (Fig. 1 6 1, II. B). Caterpillars 
smooth or spiny, with the extremity of the u<><ly divided ; frequently 
gretjitrioiix. These Insects have become notorious from the extra- 
ordinary brilliancy of blue colour exhibited by the upper surface of 
the wings of the typical genus Morpho. The species ofMorpho are 
all Insects of large size, but with wings enormous in proportion 
to the body ; this latter part is carried in a sort of cradle formed 
by the inner parts of the margins of the hind wings. Although 
an arrangement of this kind is seen in numerous other butter- 
flies, yet there is perhaps none in which it is carried to quite 
such a pitch of perfection as it is in MorpJio, where, on the 
under surface no part of the body behind the posterior legs can 



vi RHOPALOCERA MORPHIDES BRASSOLIDES 349 



be seen. There are only about 100 species of Morphides, and 
50 of these are included in Morplio, which is peculiar to tropical 
and sub-tropical America ; the other half of the family is divided 
among ten or twelve genera, found in the Indo-Malay region ; 
there being none in Africa. The eastern Morphides, though 
fine Insects, are not to be compared, either in size or brilliancy, 
with their American allies. The species of Morplio are ap- 
parently found only in the great forests of South America, 
where they are far from rare ; some have a flapping and undulat- 
ing flight, straight onwards along the alleys of the forest, and 
near the ground ; others are never seen except steadily gliding 
with outstretched wings from 20 to 100 feet above the ground, 
where they move across sunny spaces between the crowns of the 
taller trees ; the low - flyers settle frequently on the ground 
to suck the juices from fallen fruit, but the members of the 
other section never descend to the ground. As regards the 
caterpillars, "\V. M tiller tells us l that the spines they are armed 
with break off, and enter the skin, if the creatures are carelessly 
handled. Four of the five species known to him are conspicu- 
ously coloured with black, red, yellow and white. The individuals 
are gregarious. The larvae of M. achilles sit in companies, often 
of more than 100 individuals, on trunks of trees, and so form a 
conspicuous patch. The caterpillars of M. epistrophis hang to- 
gether as red clumps on the twigs of their food-plants. Hence 
it appears that in this genus w r e have an exception to the rule 
that night-feeding caterpillars rest in a hidden manner during 
the day. 

Sub-Fain. 5. Brassolides. Large butterflies, with the cell of 
the hind wing 'closed, and usually with a. small adjoining predis- 
coidal cell. Larva not very s-piny ; thinner at the two ends, the tail 
l>ijid, the head perpendicular and margined with spines. This 
small sub-family includes less than 100 species arranged in about 
eight genera, all South American. They have the very unusual 
habit of resting during the day like moths, becoming active only 
late in the afternoon. They are truly noble Insects ; although 
not possessed of the brilliant colours of Morplio, they are 
adorned, especially on the under surface, with intricate lines 
and shades most harmoniously combined, while the upper surface 
is frequently suffused with blue or purple. This sub-family 

1 Kosmos, xix. 1886, p. 355. 



350 



LEPIDOPTERA 



CHAP. 






attains its highest perfection in the 
genus Caligo ; they are enormous 
Insects, and some of them not rare. 
The larva of C. eurylochu.s (Fig. 179) 
during early life is green, and sits 
on the leaf of a Masa, but after the 
third moult it becomes brown and 

hides itself among the dry leaves. It 
is common in the gardens of Eio de 

Janeiro, where its pupae are found on 

~ the walls, like those of our white 

^ butterflies here. 

Sub - Fam. 6. Acraeides. - - Sul- 

x median ncrrure of fore- wl mjs not forked 
at the lase ; the -median without spur. 

Q 

Cells closed. Palpi in section cylindric, 
sparingly set with Ixdrs. Larva armed 
-r with Iranched spines. A somewhat 
2 monotonous and uninteresting division : 
r _ ; the size is moderate or small, and the 
colours not artistic, but consisting of 
ill-arranged spots ; the under side of 
the hind wings very frequently cliver- 
"| sifted by numerous line-like marks, 
radiately arranged, and giving place at 
the base to a few spots. There are 
about 200 species known, of which the 
majority are African ; there are but 
few Oriental or South American species. 
Some authorities consider there is 
o only one Eastern genus, but others 
prefer to adopt seven or eight divisions. 
Aliicna is now placed in Lycaenidae, 
though until recently it was con- 
sidered to belong here. The females of 
some species possess an abdominal pouch 
somewhat similar to that of Parnassius. 
The members of this suit-family 
are considered to be of the protected 
kind. 



vi RHOPALOCERA HELICONIIDES 3 5 I 

Sub-Fam. 7. Heliconiides. S/ibmedian nervure of front wing 
not forked ; median with a short spar near the base. Cell of hind 
wing dosed by a perfect nervule. Palpi compressed, witli scales at 
the sides, in front covered with hairs. Male with an elongate 
n n jointed, female witli a four-jointed, front tarsus. Caterpillars set 
with lr ni'iu'd, spines. This family is peculiar to tropical America 
and consists of only two genera, Heliconius and Eueides, with 
about 150 species; but it is one of the most characteristic of the 
South American groups of Butterflies. It is very closely allied to 
the Nymphalides, especially to the genera Metamorpha and Colaenis, 
but is readily distinguished by the perfectly-formed nervules that 
close the wing-cells. The wings are longer and narrower than 
in Nymphalides, and the colour, though exhibiting much diver- 
sity, is on the whole similar to that of the heavily-scaled forms 
of Ithomiides of the genera Titliorea, Melinaea, Melanitis ; there 
being in several cases a great resemblance between species of the 
two groups. A frequent feature in one group of Heliconius is 
that the hind wing bears a patch of red prolonged outwards by 
angular radiating marks. The individuals of certain species If. 
melpomene and H. rhea are known to execute concerted dances, 
rising and falling in the air like gnats ; when some of them 
withdraw from the concert others fill their places. H. erato 
exhibits the very rare condition of trichroism, the hind wings 
being either red, blue, or green. Schatz states that the different 
forms have been reared from a single brood of larvae. The cater- 
pillars of Heliconiides live on Passiflorae, and are said to be very 
similar to our European ^4;v/?/ /mis-caterpillars. The chrysalids 
are very spinous. "We may here remark that considerable con- 
fusion exists in entomological literature in consequence of Itho- 
miides having been formerly included in this sub-family ; for 
remarks formerly made as to " Heliconiides," but that really 
referred only to Ithomiides, have been interpreted as referring 
to Heliconiides of the present system. 

The Heliconiides seem remarkably plastic as regards colour, 
and are therefore exponents of " homoeochromatism." Bates 
says, as regards them : " In tropical South America a numerous 
series of gaily-coloured butterflies and moths, of very differ- 
ent families, which occur in abundance in almost every locality 

/ i/ 

a naturalist may visit, are found all to change their hues and 
markings together, as if by the touch of an enchanter's wand, at 



352 LEPIDOPTERA CHAP. 

every few hundred miles, the distances being shorter near the 
eastern slopes of the Andes than nearer the Atlantic. So close 
is the accord of some half-dozen species (of widely different genera) 
in each change, that he had seen them in large collections classed 
and named respectively as one species." Many of them are 
believed to be permeated by nauseous fluids, or to possess glands 
producing ill-smelling secretions. 

Sub-Fam. 8. Nymphalides. Cells, of both front and hind 
/'/////, citjicr closed only l:nj imperfect transverse nervules or entirely 
open. Fi-onl tiii'xiis of the male unjointed and without sjiin.cs, 
of the female four- or fire-jointed. Caterpillar either spincd 
or smooth; in the latter case the head nn'c. or lexs stromjly 
horned or seined, mid the apex of the body bijid. This sub- 
family is specially characterised by the open cells of the 
wings ; the discocellulars, even when present, being frequently 
so imperfect as to escape all but the most careful observa- 
tion. The Nymphalides include upwards of 150 genera and 
2000 species. The divisions having smooth larvae are separated 
by Kirby ' 2 and others as a distinct sub-family (Apaturides). In 
Britain, as in most other parts of the world, Nymphalides is the 
predominant group of butterflies. We have eighteen species, am< >i ig 
which are included the Fritillaries, Admirals, Purple Emperor, and 
the various Vanessa Peacock, Camber well Beauty, Red Admiral, 
Tortoise-shells, and Painted Lady. All have spined caterpillars 
except the Emperor. In the temperate regions of the northern 
hemisphere Vanessa may be considered the dominant butter- 
flies, they being very numerous in individuals, though not in 
species, and being, many of them, in no wise discomfited by 
the neighbourhood of our own species. Several of them are 
capable of prolonging and interrupting their lives in the winged 
condition to suit our climate ; and this in a manner that can 
scarcely be called hibernation, for they frequently take up the 
position of repose when the weather is still warm, and on the 
other hand recommence their activity in the spring at a very 
early period. This phenomenon may frequently be noticed iu 
the Tortoise-shell butterfly ; it is as if the creature knew 1:1 uit 
however warm it may be in the autumn there will be no more 
growth of food for its young, and that in the spring vegetation 

1 1'. i'ii/. ,W. LfDulun, 1879, p. xxix. 
2 Allen's Naturalists' Library, Butterflies, i. 1896. 



vr BUTTERFLIES NYMPHALIDES 353 

is sure to be forthcoming and abundant before long, although 
there may be little or none at the time the creature resumes 
its activity. It is probable that the habit may be in some 
way connected with an imperfect activity of the sexual organs. 
It should, however, be recollected that many larvae of butterflies 
hibernate as young larvae after hatching, and, sometimes, with- 
out taking any food. Pyrameis cardui, the Painted Lady, is, 
taking all into consideration, entitled to be considered the most 
ubiquitous of the butterfly tribe. Its distribution is very wide, 
and is probably still extending. The creature is found in 
enormous numbers in some localities, especially in Northern and 
Eastern Africa ; and when its numbers increase greatly, migration 
takes place, and the Insect spreads even to localities where it 
cannot maintain itself permanently. In Britain it is probably 
during some years nearly or quite absent, but may suddenly 
appear in large numbers as an immigrant. The favourite food 
of the larva is thistles, but many other plants serve the Insect 
at times. 

I'linessa, or Pyrameis, 1 atalanta, the Red Admiral, is common 
in the Palaearctic and Nearctic regions, and extends its range to 
various outlying spots. The most remarkable of these is the 
remote Hawaiian Islands, where the Insect appears really to be 
mw at home, though it is associated with a larger and more 
powerful congener, P. tameamea. Another interesting Yauessid is 
Araschnia levana, which is peculiar to Europe, where it produces 
annually two generations so dissimilar to one another that they 
passed current as two species, V. levana and V. prorsa. Although 
intermediate forms are rare in nature they can be induced by 
certain treatments applied to the larvae under human control. 

The dead-leaf butterflies of the genus Kallima belong to 
Xymphalides. They are so shaped and coloured that when 
settled, with wings closed, on a twig, the appearance is exactly 
that of a dry leaf: the exposed surface is mottled with spots 
that look just like the patches of minute fungi, etc, that are so 
common on decaying vegetation. The colour and the spots on 
the under surface of this butterfly are very variable. According 
to Mr. Skertchly,' 2 we may presume that in the minute details of 

1 A most unfortunate diversity exists in the generic names applied to these 
1'unessa, as well as in those of many other Lepidoptera. 
-Ann. Xat. Hist. (6), iv. 1889, p. 212. 
VOL. VI 2 A 



354 LEPIDOPTERA CHAP. 

these resemblances we have a case of hypertely similar to that 
of the resemblance to Insects' minings exhibited by certain marks 
OH the tegmina of Pterochroza (mentioned in Vol. V. p. 322). 

In South America there is a somewhat peculiar genus of 
Nymphalides Agcronia that delights in settling on the trunks 
of trees rather than on flowers or leaves. It was long since noticed 
that the species of Ayeronia make a clicking noise ; in some cases 
when on the wing, in other cases by moving the wings when the 
Insect is settled. The object of the noise is quite uncertain ; it 
has been suggested that it is done in rivalry or courtship, or to 
frighten away enemies. Bigg- Wether found, however, that in 
South Brazil there is a lazy little bird to which this sound serves 
as a signal, inducing it to descend from its perch and eat the clicker. 
The mode in 'which the noise is produced is not quite clear. Sir 
George Hampson has pointed out l that the fore wing bears at 
the extreme base a small appendage bearing two hooks, and that 
two other processes on the thorax play on these when the wing 
moves. His suggestion that these hooks are the source of the 
sound seems highly probable. 

There is a great variety in the larvae of Nymphalides. In 
the Vanessa group the body is armed with spines, each one of 
which bears shorter thorns, the head being unadorned. The 
Fritillaries (Argynnis, Melitaea] also have caterpillars of this kind. 
In many other forms the head itself is armed with horns or spines 
of diverse, and frequently remarkable, character. In Apatwa and 
its allies the body is without armature, but the head is perpen- 
dicular, the vertex bifid and more or less prolonged. The 
caterpillar of our Purple Emperor, Apatura iris, is quite unlike 
any other British caterpillar ; in colour it is like a Sphingid 
larva -green with oblique lateral stripes of yellow and red but 
in form it is slug-like, pointed behind, and it has on the head 
two rather long tentacle-like horns. In the South American genus 
Prepona, the larva of which in general form resembles that of 
Apatura, there are no anal claspers, but the extremity of the body 
is prolonged, forming a sort of tail. 

Fam. 2. Erycinidae (Lemoniidae of some authors). --The 

female has six perfectly formed leys, tlmuyli fj/e front pair is smaller. 

The male has the coxae of the front leys for in ing a spine, and the 

tarsi unjointed, withoitt claws. This family consists of about 1000 

1 P. Zool. Sue. London, ]S92, ]>. 191. 



vi BUTTERFLIES ERYCINIDAE 355 

species, usually of rather small size, exhibiting a great variety of 
shape and coloration, some of them being remarkably similar to 
some of the gay, diurnal moths of South America. The palpi are 
usually small, but in Ourocnemis they are large and porreet. The 
family is specially characteristic of tropical America, but there is 
one small group of 30 or -40 species, Nemeobiides, in the Eastern 
Hemisphere. We have one species in Britain, Nemeohius lucina, 
the Duke of Burgundy Fritillary. Neither the larvae nor the 
pupae of Erycinidae present any well-marked characteristic 
feature, but exhibit considerable variety. According to Bar, 1 
some of the larvae are like those of moths ; the caterpillar of 
Meliboeus is said to be like that of a Liparis : the chrysalis has 
the short, rounded form of that of the Lycaenidae, and is sus- 
pended with the head down, and without a band round the 
body. The larvae of Eurygona .are gregarious. The pupae of some 
other forms adhere, heads downwards, to branches. Scudder 
considers that this family is not distinct from Lycaenidae, and 
that the Central American genus Eumaeus connects the two. 
Eeuter also treats Erycinidae as a division of Lycaenidae. 

Sub-Fam. 1. Erycinides. [Characters of 'the family .] Palpi 
not unusually large. We place all the Erycinidae in this sub- 
family except the following 

Sub-Fam. 2. Libytheides. Butter flics of average size, with 
the palpi large and porreet: the front legs of the male small, the 
tarsus reduced to one joint: the front leg of the female of the 
normal structure, and hut little reduced in size. This division 
consists of the single genus Liliythea, with only a score of species. 
They are Insects somewhat like Vanessa, in appearance, but tan- 
not fail to be recognised on account of the peculiar palpi. The 
genus is of very wide distribution, occurring in most parts of the 
warm and temperate continental regions, and it also occurs in 
Mauritius and the Antilles. 

The Libytheides have given rise to much difference of opinion 
amongst systematists, some of whom assign them as a sub- 
family to the Erycinidae, some to the Nymphalidae ; while others 
treat them as a family apart. The families Nymphalidae, Ery- 
cinidae and Lycaenidae are so intimately allied, that Scudder is 
probably correct in considering them to form really one huge 
family ; if this view were adopted there would be no difficulty 

1 Bull. 8oc. cnt. France, 1856, pp. c, ci. 



LEPIDOPTERA CHAP. 



iu locating Libythea therein. If they he kept apart, it is almost 
necessary to separate Libythca also; though possibly its claims 
to he placed in Erycinidae slightly preponderate. The recently 
described germs Ourocnemis to some extent connects Erycinides 
with Libythaeides. 1 

Fam. 3. Lycaenidae. The front legs but little smaller than 
the others : in t/tr male, however, the tarsus, though elongate, is only 
of one joint, and is terminated by a single claw. No pad on the 
front tibia. Claws not toothed. The Lycaenidae, or Blues, are, as 
a rule, of small size, but in the tropics there are many that reach 
the average size of butterflies, i.e. something about the stature of 
the Tortoise-shell butterfly. The family is one of the larger of 
the divisions of butterflies, considerably more than 2000 species 
being at present known, and this number is still rapidly increas- 
ing. Although blue on a part of the upper, surface is a very 
common feature in the group, it is by no means universal, for 
there are many " Coppers," as well as yellow and white Lycae- 
nidae. Many species have delicate, flimsy appendages tails 
to the hind wings, but in many others these are quite absent ; 
and there are even tailed and tailless forms of the same species. 
The members of the group Lipteninae (Liptena, Vanessnla, 
Miinarraea, etc.) resemble members of other sub-families of Xym- 
phalidae, and even of Pieridae. Lycaenidae are well represented 
wherever there are butterflies ; in Britain we have 1 8 species. 

The larvae of this family are very peculiar, being short, thicker 
in the middle, and destitute of the armature of spines so remark- 
able in many other caterpillars. It has of late years been fre- 
quently recorded that some of these larvae are attended by ants, 
which use their antennae to stroke the caterpillars and induce 
them to yield a fluid of which the ants are fond. Gueiiee had 
previously called attention L> to the existence of peculiar structures 
contained in small cavities on the posterior part of the cater- 
pillar of Lycacna, baetica. These structures can be evaginated, 
and, it is believed, secrete a fluid ; Edwards and M'Cook are of 
opinion that they are the source of the matter coveted by the 
ants. The larvae are without spines. 

The caterpillars of the Blues have some of them strange tastes ; 
more than one has been recorded as habitually feeding on Aphidae 

1 Baker, Tr. ent. Soc. London, 1887, p. 175, PI. ix. 

2 Ann. Soc. ent. France (4), vii. 18(i7, ]i. (!(.>,">, PI. xiii. 



vi BUTTERFLIES LYCAENIDAE PIERIDAE 357 



and scale-Insects. The pupae are, like the larvae, of short 
inflated form. By a remarkable coincidence, the pupae of two 
species bear a considerable resemblance to the heads of monkeys, 
or mummies. The Lycaenid pupa is usually extremely consoli- 
dated, destitute of movement, and is supported in addition to 
the attachment by the cremaster by a silk thread girdling the 
middle. There are exceptions to these rules, and according to 
Mr. liobson the pupa of Tajuria diaeus hangs free, suspended 
from a leaf, and can move the body at the spot where the 
abdominal segments meet the wing-cases in the dorsal line. 1 

Fam. 4. Pieridae. The six legs well developed, and similar 
in the sexes ; there is no pad on the front tibia. The claws of all 
the feet are li/id, or toothed, and there is an empodium. There 
are upwards of 1000 species of Pieridae already known. Al- 
though several taxonomists treat the Pieridae and Papilionidae 
as only subdivisions of one family, yet they appear to be quite 
distinct, and the relationships of the former to be rather with 
Lycaenidae. In Pieridae, white, yellow, and red are the pre- 
dominant colours, though there is much black also. It lias 
recently been ascertained that the yellow and red pigments, as 
well as the white, are uric acid or derivatives therefrom. 2 The 
physiology of this peculiarity has not yet been elucidated, so 
that we do not know whether it may be connected with some 
state of the Malpighiaii vessels during metamorphosis. 

Our Garden-White, Brimstone, Clouded-yellows and Orange-tip 
butterflies belong to this family ; as does also the South American 
genus formerly called Leptalis. This generic name, which is 
much mentioned in literature owing to the resemblance of the 
species of the genus to Heliconiides, has now disappeared ; Leptalis 
having been divided into various genera, while the name itself 
is now considered merely a synonym of Dismoi^h ia. 

The African Insect, Pseudopontia paradoxa, has nearly trans- 
p; i rent wings, no club to the antennae, a remarkably small cell on 
the wing, and an arrangement of the nervules not found in any 
other butterfly ; there being only ten nervules at the periphery of 
the front wing, and both upper and lower radial nervules uniting 
with the posterior branch of the subcostal. It has been treated 
as a moth by several entomologists. Aurivillius considers that it 

1 /. Bombay Soc. ix. 1895, pp. 338-341. 

- Hopkins, Phil. Trans. 186 B, 1895, p. 661. 



353 



LEPIDOPTERA 



CHAP. 



is certainly a butterfly ; but as the metamorphoses are unknown, 
we cannot yet form a final opinion as to this curious form. The 
extraordinary Peruvian Insect, Styx inf emails, is also placed in 
this family by Staudinger ; it is a small, pale Insect, almost white, 
and with imperfect scales ; a little recalling a Satyrid. It appears 
to be synthetic to Pieridae and Erycinidae. 

The caterpillars of Pieridae are perhaps the least remarkable 
or attractive of all butterfly-caterpillars ; their skins are as a 

rule bare, or covered only with fine, short 
down or hair ; their prevalent colour is 
green, more "or less speckled with black 
and yellow, and they are destitute of any 
prominent peculiarities of external struc- 
ture. Pupation is accomplished by the 
larva fixing itself to some solid body by the 
posterior extremity, with the head upwards 
(or the position may be horizontal), and then 
placing a girdle round the middle of the 
body. The pupa never hangs down freely 
as it does in Xymphalidae. It has been 
by experiment that if the 




B 



9 ? 



FIG. ISO. Pupation of the 

Orange - tip butterfly, ascertained 

XucMoe examines. A, ifdl d t} j b fc t j 

Ine completed pupa ; B, 

the larva, with its girdle, tion can nevertheless be accomplished by 

prepared for the change. ft consitlerable propol . t i on o f larvae. Some 

of the pupae are of very peculiar form, as is the case in the 

Orange-tip (Fig. 180, A) and Brimstone butterflies. The Orange- 

tip butterfly passes nine or 

ten months of each year as 

a pupa, which is variable 

in colour ; perhaps to some 

extent in conformity with 

its surroundings. The North 

American E. genutia has a 

similar life-history, but the 

larva leaves its Cruciferous 

food-plant, wanders to an FIG. 181. Newly-hatched larva of Euchloe car- 

A ' ^ If? j n P rofil< : ; * 01 f 

segment more magnified, showing the liquid- 
bearing setae ; C, one of the setae still more 
magnified, and without liquid. 




oak tree, and there turns to 

a pupa, resembling in colour 



the bark of the tree. 

It is riot unusual for caterpillars to change their habits and 



VI 



BUTTERFLIES PIERIDAE PAPILIONIDAE 



359 




appearance in a definite manner in the course of the larval life. 
The caterpillar of Eucliloe cardamines exhibits a larval meta- 
morphosis of a well-marked character. The young larva (Fig. 
181) is armed with peculiar setae, furcate at the tip, each of 
which bears a tiny 
ball of fluid. In this 
stage the caterpillar 
makes scarcely any 
movement. In the 
middle of the cater- 
pillar's life a new 
vestiture appears 
after an ecdysis ; 
numerous fine hairs 

are present, and the FIG. 182. Larva of Euchloe. cardamines in middle life. 
fluid -bearilio- spines A> thu larva iu f rotile ; B - one se g m e"t more magnified. 

nearly disappear, being reduced to a single series of spines of a 
comparatively small size on each side of the upper middle region 
of the body (Fig. 182). The colour is also a good deal 
changed, and concomitantly there is a much greater voracity 
and restlessness. 

Fam. 5. Papilionidae. All the legs well developed. Claws 
large, simple, without empodium. Front tibiae with a pad. The 
metanotum free, conspicuously exposed between mesonotum and 
abdomen. This series of butterflies includes some of the most 
magnificent of the members of the Insect world. It is considered 
by some authorities to be the highest family of butterflies ; and 
in one very important feature sexual differentiation it cer- 
tainly is entitled to the rank. There are about 700 recorded 
species, the larger portion of which are included in the genus 
Papilio. The great variety of form has led to this genus being 
divided ; the attempts have, however, been partial, with the 
exception of an arrangement made by Felder, who adopted 75 
sections, and a recent consideration of the subject by Haase, who 
arranges Felder 's sections into three sub-genera. Many of the 
sections have received names, and are treated by some authors as 
genera, so that an unfortunate diversity exists as to the names 
used for these much-admired Insects. The genus is distributed 
all over the world, but is perhaps nowhere more numerous in 
species than in South America. Wallace informs us that the great 



360 



LEPIDOPTERA 



CHAP. VI 



majority of the species of the Amazon valley frequent the shady 
groves of the virgin forest. In many cases the sexes are ex- 
tremely different in appearance and habits, and are but rarely 
found together in one spot. The genus Ornitlioptera is closely 




allied to Papilio, and contains some of the most remarkable of 
butterflies, the homes of the species being the islands of the Malay 
Archipelago, and outlying groups of islands, there being a smaller 
number of species in the neighbouring continents. The females 
are of great size, and are so excessively different from their 



362 l.KI'IDOPTERA CHAP. 

consorts of the other sex, as to arouse in the student a feeling of 
surprise, and a strong desire to fathom the mysteries involved. It 
would L)e difficult to surpass the effective coloration of the males 
in many of the species of Ornithoptera ; they are, too, very diverse 
in this respect ; 0. brookiana is of an intense black colour, with 
a band of angular green marks extending the whole length of 
its wings, while behind the head there is a broad collar of crimson 
colour. Perhaps the most remarkable of all is the 0. paradisea, 
recently discovered in New Guinea ; in this species the sexual 
disparity reaches its maximum. The female (Fig. 184) is a 
large, sombre creature of black, white and grey colours, but the 
male (Fig. 183) is brilliant with gold and green, and is made 
additionally remarkable by a long tail of unusual form on each 
hind wing. 

We may anticipate that these extraordinary cases of sexual 
total dissimilarity in appearance are accompanied by equally 
remarkable habits and physiological phenomena. In the case of 
0. li-imkitrtift. the female is extremely rare, so that the collector, 
Kiinstler, could only obtain fifteen females to a thousand males. 
According to Mr. Skertchly, instead of the crowd of males being- 
eager to compete for the females, the reverse is the case ; the 
female diligently W T OOS the male, who exhibits a reluctance to 
coupling. This observer apparently considered that the " emerald 
feathers " of the male are a guide or incitement to the female. 1 

In Africa Ornithoptera is to a certain extent represented 
by two extremely remarkable forms, Papilio zalmoxis and P. 
(Druri/a*) antirnachus. There are about a dozen other genera of 
Papilionidae ; most of them contain but few species. Parnassius, 
however, is rich in species inhabiting the mountains and elevated 
plateaus of the northern hemisphere in both the Old and Xew 
Worlds ; it is remarkable for the small amount of scales on the 
wings, and for the numerous variations of the species. The female 
possesses a peculiar pouch at the end of the body ; although only 
formed during the process of coupling, it has a special and 
characteristic form in most of the species. The curious Indian 
genus Leptt>rir<-tfH has parts of the front wings transparent, while 
the hind pair form long tails. This genus is of interest in that 

1 Ann. Nat. ///*/. (6), iv. 1889, p. 213. "We trust there will not bo many more 
Kiinstlcrs. as this beautiful butterfly must certainly become extinct, if the female 
1)0 really as rare as is supposed. 



vi BUTTERFLIES SKIPPERS 363 

it is said to connect Papilionidae to some extent with Hesperiidae. 
The larvae of this family are remarkable on account of a curious 
process on the thoracic segment called an " osmeterium." It is 
usually retracted, but at the will of the caterpillar can be everted 
in the form of a long furcate or Y-shaped process ; there is a 
gland in the osmeterium, and as a result a strong odour is 
emitted when the exstulpation occurs. 

The pupation of Papilionidae is similar to that of Pieridae, 
the pupa being placed with the head upwards, fixed by the tail, 
and girt round the middle. A very curious diversity of pupation 
occurs in the genus Thais, in which the pupa is attached by the 
tail as usual, and which is quite exceptional also by a thread 
placed at the top of the head. Scudder thinks there is also a 
girdle round the middle, but Dr. Chapman inclines to the view 
that the thread attaching the head is really the median girdle 
slipped upwards. The pupation of Parnassius is exceptional, 
inasmuch as, like Satyrides, it is terrestrial, in a slight construc- 
tion of silk. 

Fam. 6. Hesperiidae (Skippers). -Six perfect legs: metanotum 
not free, largely covered Toy the mesonotum. A pad on the front tibia. 
Claws short and thick ; empodium present. Although this family 
has been comparatively neglected by entomologists, upwards of 
2000 species and more than 200 genera are known, and it is not 
improbable that it may prove to be as extensive as JSTymphalidae. 
We have already said that Hesperiidae is generally admitted to 
be the most distinct of the butterfly groups. It has been thought 
by some taxonomists to be allied to Papilionidae, but this is a 
mistake. It is undoubtedly more nearly allied to Heterocera, 
and when the classification of Lepidoptera is more advanced, so 
that the various natural groups placed in that sub-Order are 
satisfactorily distinguished, it is probable that Hesperiidae will 
be altogether separated from Rhopalocera. We have already 
mentioned that E. Reuter considers the Hesperiidae to be phylo- 
genetically unconnected with Rhopalocera proper; but though 
quite ready to admit that he will probably prove correct in this, 
w T e think Lepidopterists will not be willing to recognise the 
family as a sub-Order equivalent in value to all Heterocera. 

The body is shorter and thicker than it is in most butterflies, 
and is pointed at the tip rather than knobbed or bent down- 
wards; the wings are less ample; the antennae are not truly 



364 LEPIDOPTERA CHAP. 

knobbed, but are thicker before the actual tip, which is itself 
pointed and more or less bent backwards, so that the antennae 
are somewhat hook-shaped. 

In habits as well as structure the family is markedly distinct 
from butterflies; the pupation is peculiar, and the name Skipper 
has been applied to the perfect Insects, because so many of them 
indulge in a brief, jerky flight, instead of the prolonged aerial 
courses characteristic of the higher butterflies. 

There is great difference among the members of the family, 
and some of them possess a very high development of the powers 
of locomotion, with a correspondingly perfect structure of the 
thoracic region, so that, after inspection of these parts, we can 
quite believe Wallace's statement that the larger and strong- 
bodied kinds are remarkable for the excessive rapidity of their 
flight, which, indeed, he was inclined to consider surpassed that 
of any other Insects. " The eye cannot follow them as they dart 
past ; and the air, forcibly divided, gives out a deep sound louder 
than that produced by the humming-bird itself. If power of 
wing and rapidity of flight could place them in that rank, they 
should be considered the most highly organised of butterflies." 
It was probably to the genera Pyrrliopyge, Ery tides, etc., that 
Mr. Wallace alluded in the above remarks. Although the Hes- 
periidae are not as a rule beautifully coloured, yet many of these 
higher forms are most tastefully ornamented ; parts of the wings, 
wing-fringes, and even the bodies being set with bright but agree- 
able colours. We mention these facts because it is a fashion to 
attribute a lowly organisation to the family, and to place it as 
ancestral to other butterflies. Some of them have crepuscular 
habits, but this is also the case with a variety of other Ehopalo- 
cera in the tropics. 

In their early stages the Skippers so far as at present known 
depart considerably from the majority of butterflies, inasmuch 
as they possess in both the larval and pupal instars habits of con- 
cealment and retirement. The caterpillars have the body nearly 
bare, thicker in the middle, the head free, and more or less 
notched above. They make much greater use of silk than other 
butterfly-larvae do, and draw together leaves to form caves for 
concealment, and even make webs and galleries. Thus the habits 
are almost those of the Tortricid moths. Pupation takes place 
under similar conditions ; and it is interesting to find that Chap- 



VI 



HESPERIIDAE SKIPPERS 



365 



man considers that the pupa in several points of structure re- 
sembles that of the small moths. Not only does the larva draw 
together leaves or stalks to make a shelter for the pupa, hut it 
frequently also forms a rudimentary cocoon. These arrangements 
are, however, very variable, and the accounts that have been 
given indicate that even the same species may exhibit some 
amount of variation in its pupation. Scudder considers that, in 
-the North American Skippers, the cremaster is attached to a single 
Y-like thread. In other cases there is a silk pad on the leaf for 
the cremaster to hook on. An interesting account given by Mr. 
Dudgeon of the pupation of a common Indian Skipper, Badamia 
exclamationis, shows that this Insect exercises considerable in- 



FIG. 185. Pupation of 
Badam ia exclamation is. 
(After Dudgeon. J. 
.Bombay Soc. x. 1895, 
p. 144). A, One side 
of the leaf-cradle, the 
other (nearest to the 
observer) being broken 
away ; B, transverse sec- 
tion of entire cradle. </, 
The pupa ; b, fastenings 
of perpendicular threads 
round pupa ; c, cross 
thread retaining the leaf 
m cradle form ; d, mar- 
gins of the leaf ; e, mid- 
rib of leaf. 



genuity in the structure of the puparium, and also that the 
arrangements it adopts facilitate one of the acts of pupation most 
difficult for such pupae as suspend themselves, viz. the hooking the 
cremasters on to the pad above them. Badamia uses a rolled-up 
leaf (Fig. 185); the edges of the leaf are fastened together by 
silk at d ; from this spot there descends a thread which, when 
it reaches the pupa, a, forks so as to form an inverted Y, and is 
fastened to the leaf on either side ; the two sides of the leaf are kept 
together by a cross thread, cc. Mr. Dudgeon was fortunate enough 
to observe the act of pupation, and saw that " although the anal 
prolegs of the larva were attached to a tuft or pad of silk in the 
usual way, and remained so until nearly the whole skin had been 
shuffled off, yet when the last segment had to be taken out, the 
pupa drew it entirely away from the skin and lifted it over the 




366 LEPIDOPTERA CHAP. 



riiipty skin, and by a series of contortions similar to those made 
by an Insect in depositing an egg, it soon re-attached its anal 
segment or cremaster to the wel), throwing away the cast-off skin 
by wriggling its body about." 

Series II. Heterocera. Moths. 

Although Rhopalocera if exclusion be made of the Hes- 
periidae is probably a natural group, yet this is not the case 
with Heterocera. The only definition that can be given of 
Heterocera is the practical one that all Lepidoptera that are not 
butterflies are Heterocera,. Numerous divisions of the Heterocera 
have been long current, but their limits have become more and 
more uncertain, so that at the present time no divisions of greater 
value than the family command a recognition at all general. This 
is not really a matter of reproach, for it arises from the desire to 
recognise only groups that are capable of satisfactory definition. 

Several attempts have recently been made to form a rough 
forecast of the future classification of moths. Professor Comstock, 
struck by some peculiarities presented by the Hepialidae, Mierop- 
terygidae (and Eriocephalidae), recently proposed to separate them 
from all other Lepidoptera as a sub-order Jugatae. Comstock's 
discrimination in making this separation met with general ap- 
proval. The character on which the group Jugatae is based is, 
however, comparatively trivial, and its possession is not suffi- 
cient, as pointed out by Packard, 1 to justify the close association 
of Hepialidae and Micropterygidae, which, in certain important 
respects, are the most dissimilar of moths. The characters 
possessed by the two families in common may be summarised by 
saying that the wings and wing-bearing segments remain in a 
low stage of development. In nearly all other characters the 
two families are widely different. Packard has therefore, while 
accepting Comstock's separation of the families in question, 
proposed a different combination. He considers that Eriocepha- 
lidae should l)e separated from all others as " Protolepidoptera " 
or " Lepidoptera Laciniata," while the whole of the other Lepi- 
doptera, comprised under the term "Lepidoptera Haustellata," 
are divided into Palaeolepidoptera (consisting only of Microp- 
terygidae) and jSTeolepidoptera, comprising all Lepidoptera (in- 

1 Mem. Ac. Jl / '(is.'iingtun, vii. 1895, p. 57. 



vi IIETEROCERA MOTHS 367 

elusive of Hepialidae) except the Eriocephalidae and Microptery- 
gidae. The question is rendered more difficult by the very close 
relations that exist between Micropterygidae and a sub-Order, 
Trichoptera, of jSTeuroptera. Dr. Chapman, by a sketch of the 
classification of pupae, 1 and Dyar, by one on larval stages, 2 have 
made contributions to the subject ; but the knowledge of early 
stages and metamorphosis is so very imperfect that the last two 
memoirs can be considered only as preliminary sketches ; as indeed 
seem to have been the wishes of the authors themselves. 

Simultaneously with the works above alluded to, Mr. Mey- 
rick has given 3 a new classification of the Order. We allude, 
in other pages, to various points in Mr. Meyrick's classifica- 
tion, which is made to appear more revolutionary than it really 
is, in consequence of the radical changes in nomenclature com- 
bined with it. 

As regards the various aggregates of families that are widely 
known in literature by the names Bombyces, Sphinges, JSToctuae, 
Geometres, Pyrales, we need only remark that they are still 
regarded as to some extent natural. Their various limits being 
the subject of discussion and at present undecided, the groups 
are made to appear more uncertain than is really the case. The 
group that has to suffer the greatest changes is the old Bom- 
byces. This series comprises the great majority of those moths 
that have diurnal habits. In it there were also included several 
groups of moths the larvae of which feed in trunks of trees or 
in the stems of plants, such as Cossidae, that will doubtless prove 
to have but little connection with the forms with which they were 
formerly associated. These groups with aberrant habits are those 
that give rise to the greatest difficulties of the taxonomist. 

The following key to the families of Heterocera is taken from 
Sir G. F. Hainpson's recent work, Fauna of British India J/oM.s. 4 
It includes nearly all the families at present recognised among 
the larger Lepidoptera ; certain families 5 not mentioned in this 
key are alluded to in our subsequent remarks on the families : 

1 Tr. cnt. Soc. London, 189-3, p. 97, with Suppl. up. dt. 1896, pp. 129 and 567. 

2 Amcr. Natural, xxix. 1S95, p. 1060. See also Ann. X. York Ac. viii. 1895, 
p. 194, and Ent. Record, 1897, pp. 136 and 196. 

3 Handbook of British Lepidoptera, 1895. 

4 London, 1892. Published under the authority of the Secretary of State for 
India in Council. 

' Those numbered 2, 8, 10, 17, 22. 27. 44, and 46 in our arrangement. 



368 LEPIDOPTERA CHAP. 



KEY TO THE FAMILIES OP MOTHS l 

N.B. This table is not simply dichotomic ; three contrasted categories are used 
in the case of the primary divisions, A, B, C, and the secondary divisions, 

I, II, III. 

A. Fore wing with nervule 5 coming from the middle of the discocellulars, 

or nearer 6 than 4 (Categories I, II, III = 1-18). 
I. Frenulum rudimentary . . Fam. 38. Epicopeiidae, see p. 418 

II. Frenulum absent (Categories 1-8). 

1. Proboscis present, legs with spurs (Cat. 2-5). 

2. Hind wing with nervule 8 remote from 7 (Cat. 3 and 4). 

3. Fore wing with nervule 6 and 7 stalked 

Fam. 39. Uraniidae, see p. 419. 

4. Fore wing with nervules 6 and 7 not stalked 

Fam. 5. Ceratocampidae, see p. 375. 

5. Hind wing with nervule 8 nearly touching 7 beyond end of cell 

Fam. 4. Brahmaeidae, see p. 374. 
6. Proboscis absent, legs without spurs (Cat. 7 and 8). 

7. Hind wing with one internal nervure 

Fam. 3. Saturniidae, see p. 372. 

8. Hind wing with two or three internal nervures 

Fam. 6. Bombycidae, see p. 375. 
III. Frenulum present (Cat. 9-18). 

9. Antennae fusiform [spindle-shaped] Fam. 9. Sphingidae, seep. 380. 
10. Antennae not fusiform (Cat. 11-18). 

11. Proboscis absent . . Fam. 7. Eupterotidae, see p. 376. 

12. Proboscis present (Cat. 13-18). 

13. Hind wing witli nervule 8 curved and almost touch- 
ing 7 after end of cell ; nervure la reaching anal angle 

Fam. 12. Cymatophoridae, see p. 386. 

14. Hind wing with nervule 8 remote from 7 after end of 

cell (Cat, 15-18). 

1 5. Tarsi as short as tibia, hairy ; stoutly built moths 

Fam. 11. Notodontidae,' 2 see p. 383. 

16. Tarsi long and naked ; slightly built moths (Cat. 

17 and 18) 

17. Fore wing with nervule 7 remote from 8, and 
generally stalked with 6 

Fam. 40. Epiplemidae, see p. 420. 

18. Fore wing with nervule 7 given olf from 8 ; hind 
wing with nervure la short or absent 

Fam. 36. Geometridae, see p. 411. 

1 For explanatory diagram of the wings, see Fig. 161, I. When the nervuration 
is obscured by the wing-scales, it may be rendered temporarily visible by the appli- 
cation, with a camel's-hair brush, of a little benzine. The wings may be per- 
manently denuded of their scales by being placed for a short time in Eau de Javelle 
(hypochlorite of potash). 

2 The genus Cyphanta (one species from India) has nervule 5 of the fore wing 
proceeding from the lower angle of the cell. 



vi HETEROCERA MOTHS 369 

B. Fore wing with nervule 5 coming from lower angle of cell or nearer 4 

than 6 [see figures 161 and 162, pp. 318, 319] (Categories 19-58). 
19. Hind wing with more than 8 nervules (Cat. 20, 21). 

20. Proboscis absent, no mandibles nor ligula ; size not very small 

Fam. 23. Hepialidae, see p. 396. 

21. Mandibles, long palpi and ligula present ; size very small 

Fam. 47. Micropterygidae, sec p. 435. 
22. Hind wing with not more than 8 nervules (Cat. 23-58). 

23. Hind wing with nervule 8 remote from 7 after origin of 

nervules 6 and 7 (Cat. 24-51). 
24. Frenulum absent (Cat. 25-29). 

25. Hind wing with one internal nervure ; nervule 8 with 
a precostal spur, Fain. 31. Pterothysanidae, sec p. 406. 

26. Hind wing with two internal nervures (Cat. 27 and 28). 

27. Hind wing with a bar between nervules 7 and 8 
near the base ; nervure la directed to middle of inner 
margin . Fam. 30. Endromidae, see p. 406. 

28. Hind wing with no bar between nervules 7 and 8 ; 
nervure la directed to anal angle 

Fam. 29. Lasiocampidae, see p. 405. 
29. Hind wing with three internal nervures 

Fam. 21. Arbelidae, see p. 396. 
30. Frenulum present (Cat, 31-51). 

31. Hind wing with nervule 8 aborted, 

Fam. 15. Syntomidae, see p. 388. 

32. Hind wing with nervule 8 present (Cat. 33-51). 

33. Antennae knobbed Fam. 1. Castniidae, see p. 371. 

34. Antennae filiform, or (rarely) dilated a little towards 

the tip (Cat. 35-51). 

35. Fore wing with nervure Ic present (Cat. 36-43). 
36. Hind wing with nervule 8 free from the 
base or connected with 7 by a bar (Cat. 
37-42). 

37. Proboscis present 

Fam. 16. Zygaenidae, see p. 390. 

38. Proboscis absent i^Cat. 39-42). 

39. Palpi rarely absent ; J Dinged ; 
larvae wood-borers 

Fam. 20. Cossidae, see p. 395. 

40. Palpi absent ; 9 apterous (Cat. 41, 

' 42). 

41.9 rarely with legs; ^ au( l 
larvae case-dwellers 
Fam. 19. Psychidae, see p. 392. 
42. 9 an d larvae free ' 
Fam. 18.Heterogynidae,seep.392. 

1 This is a mistake of Sir George Hampson's. It has long been known that the 
female of Hcteroyi/nis does not leave the cocoon (for references see p. 392) ; the 
larvae, however, do not live in cases, as those of Psychidae do. 

VOL. VI - P 



3/0 LEPIDOPTERA CHAP. 

43. Hind wing with, nervule 8 anastomosing 
shortly with 7 

Fam. 26. Limacodidae, see. p. 401. 
44. Fore wing with nervure Ic absent (Cat, 45-51). 

45. Hind wing with nervule 8 rising out of 7 

Fam. 34. Arctiidae, see p. 408. 

46. Hind wing with nervule 8 connected with 7 

by a bar, or touching it near middle of 
cell (Cat. 47, 48). 

47. Palpi with the third joint naked and 
reaching far above vertex of head ; 
proboscis present 

Fam. 33. Hypsidae, see p. 408. 

48. Palpi not reaching above vertex of 
head ; proboscis absent or very minute 

Fam. 32. Lymantriidae, see p. 406. 
49. Hind wing with nervule 8 anastomosing 
shortly with 7 near the base ; proboscis 
well developed (Cat. 50, 51). 

50. Antennae more or less thick towards tip 

Fam. 35. Agaristidae, see p. 410. 

51. Antennae filiform 

Fam. 37. Noctuidae, see p. 414. 

52. Hind wing with nervule 8 curved and nearly or quite 
touching nervule 7, or anastomosing with it after origin of 
nervules 6 and 7 (Cat, 53-58). 
53. Hind wing with nervure Ic absent (Cat. 54-57). 

54. Hind wing with nervule 8 with a precostal spur 

Fam. 24. Callidulidae, see ]). 400. 

")"). Hind wing with nervule 8 with no precostal spur 
(Cat. 56, 57). 

56. Hind wing with, nervure la absent or very short 

Fam. 25. Drepanidae, see p. 400. 

57. Hind wing with nervure la almost or quite 
reaching anal angle 

Fam. 28. Thyrididae, see p. 404. 
58. Hind wing with nervure Ic present 

Fam. 41. Pyralidae, see p. 420. 

C. Fore wing with 4 nervules arising from the cell at almost even dis- 
tances apart (Cat. 59-66). 
59. Wings not divided into plumes (Cat. 60-63). 

60. Hind wing with nervule 8 coincident with 7 

Fam. 13. Sesiidae, see p. 386. 
61. Hind wing with nervule 8 free (Cat. 62, 63). 

62. Fore wing with nervure Ib simple or with a very 
minute fork at base 

Fam. 14. Tinaegeriidae, see p. 387. 

63. Fore wing with nervure la forming a large lurk 

with Ib at base Fam. 45. Tineidae, see p. 428. 



VI 



HETEROCERA CASTNIIDAE 



371 



04. Wings divided into plumes (Cat, 65, 66). 

65. Fore wing divided into at most two, hind wing into three 
plumes . . . Fam. 42. Pterophoridae, see p. 426. 

66. Fore wing and hind wing each divided into three plumes 

Fam. 43. Alucitidae ( = Orneodidae ;, see p. 426. 

Fam. 1. Castniidae. The Insects of this family combine to a 
extent the characters of Imtterjiies and moths. The antennae 
are knvlilm/ or hooked at t/ie tip, there is a large precostal area to 
tlte hind u-intj. The nervules of the front icing are complex ami 
<t itastomose so as to form one or more accessory cells (Fig. 1 6 2). This 
important, but not extensive, family consists chiefly of forms found 
in tropical America and Australia. The diversity of size, form and 
appearance is very great, and it is probable that the members of the 
family will be separated ; indeed, taxoiiomists are by no means 
in agreement as to the limits of the family. The Castniidae 
are diurnal Insects, and the Xorth American genus Megathymus 

is by many con- 
sidered to belong 
to the Ehopalo- 
cera. Euschemon 
rafflesiae (Fig. 186) 
is extremely like a 
large Skipper with 
long antennae, but 
has a well-marked 
frenulum. The 
members of the 
Australian genus 
Sync mo it are much smaller, but they also look like Skippers. 
Their habits are very like those of the Hesperiidae ; they 
Hit about in the hot sunshine, and when settling after their 
brief flights, the fore wings are spread out at right angles to 
the body, so as to display the more gaily coloured hind 
wings : at night, or in cloudy weather, the Insect rests on 
blades of grass with the wings erect, meeting vertically over 
the back, like a butterfly. Hecatesia, another Australian genus, 
is now usually assigned to Agaristidae ; its members look like 
moths. The male of H. fenestrata is provided with a sound- 
producing organ similar to that of the Agaristid genus Aegocera. 
The Cx tin i of South America are manv of them like 




FIG. 186. Euschemon rafflesiae. Australia. (After 
Doubleday. ) 



3/2 LEPIDOPTERA CHAP. 

Nymphalid butterflies, but exhibit great diversity, and resemble 
butterflies of several different divisions of the family. 1 

The species are apparently great lovers of heat and can 
tolerate a very dry atmosphere. 2 The transformations of very 
few have been observed; so far as is known the larvae feed in 
stems ; and somewhat resemble those of Goat-moths or Leopard- 
moths (Cossidae) ; the caterpillar of C. tliempon lives in the stems 
of Brazilian orchids, and as a consequence has been brought to 
Europe, and the moth there disclosed. The pupae are in general 
structure of the incomplete character, and have transverse rows 
of spines, as is the case with other moths of different families, 
but having larvae with similar habits. 3 Casttiia eudesmia forms 
a large cocoon of fragments of vegetable matter, knitted together 
with silk. These Insects are rare in collections ; they do not ever 
appear in numbers, and are generally very difficult to capture. 

Fam. 2. Neocastniidae. - -The Oriental genus Tusdna 
formerly placed in Castniidae has recently been separated by 
Sir G. Hampson and associated with Neocastnia nice rill el, from 
East India, to form this family. These Insects have the appear- 
ance of Xymphalid butterflies. They differ from Castniidae by 
the want of a proboscis. 

Fam. 3. Saturniidae. This is a large and varied assemblage 
of moths ; the larvae construct cocoons ; the products of several 
species being used as silk. These moths have no frenulum and 
no proboscis. The hind wings have a very large shoulder, so 
that the anterior margin or costa stretches far forward beneath 
the front wing, as it does in butterflies. The antennae of the 
males are strongly bipectinated and frequently attain a magnifi- 
cent development. The family includes some of the largest 
and most remarkable forms of the Insect-world. Coscinocera 
lie /rules, inhabiting North Australia, is a huge moth which, 
with its expanded wings and the long tails thereof, covers a 
space of about 70 square inches. One of the striking features 
of the familv is the occurrence in numerous forms of remarkable 

V 

transparent spaces on the wings ; these window - like areas 
usually occur in the middle of the wing and form a most remark- 
able contrast to the rest of the surface, which is very densely 

1 See Westwood, Tr. Linn. Soc. London (2), i. 1877, p. 165, etc. 
- For lial 'its ol' some Brazilian Gastuia see Seitz, Knt. Z<-it. Stettin, li. 1890, p. 258. 
3 For pupa see Chapman. Ent. Hcc. vi. 1895, pp. 286, 288. 



VI 



HETEROCERA SATURNIIDAE 



373 



scaled. In Attacas these attain a large size. In other species, 
such as the South African Ludia delegorguei, there is a small 
letter-like, or symboliform, transparent mark towards the tip of 
each front wing. We have at present no clue to the nature 
or importance of these remarkable markings. In the genus 
Automeris, and in other forms, instead of transparent spaces 
there are large and staring ocellate marks or eves, which are 
concealed when the Insect is reposing. In Arceina, Copiopteryx, 
E/tdaemonia and others, the hind wings are prolonged into very 
long tails, perhaps exceeding in length those of any other moths. 




FIG. 187. Larva of Attacus atlas. India. A, at end of 1st instar, profile ; B, 4th 
instar, dorsal view ; C, full-grown larva, in repose. (After Poujade.) 

The cocoons are exceedingly various, ranging from a slight 
o] icn network to a dense elaborate structure arranged as in our 
Emperor moth ; in this latter case an opening is left by the 
larva for its exit after it has become a moth, but by an ingenious, 
chevaux-de-frise work, this opening is closed against external 
enemies, though the structure offers no resistance at all to the 
escape of the moth. Fabre has recorded some observations and 
experiments which seem to show that the instinct predominating 
over the formation of the cocoon is not cognoscent. The Insect, 
if interfered with, displays a profound stupidity. Its method is 
blind perseverance in the customary. 1 The cocoon of Saturniidae 
is more often continuous, i.e. entirely closed. Packard says that 
1 Souvenirs entomologiques, quatrieme serie, 1891, pp. 39-46. 



374 LEPIDOPTERA CHAP. 



Actias In an effects its escape by cutting through the strong 
cocoon with an instrument situate at the base of the front wing. 
Other species were examined and were found to possess the in- 
strument ; but Packard is convinced that the majority of the 
species possessing the instrument do not use it, but escape by 
emitting a fluid that softens the cocoon and enables the moth 
to push itself through. 1 The cocoons of the species of Ceranclna. 
have a beautiful appearance, like masses of filagree-work in silver. 
The pupa in Ceranchia is very peculiar, being terminated by a 
long, spine-like process. In Locpa newara the cocoon is of a 
green colour and suspended by a stalk ; looking like the pod, or 
pitcher, of a plant. The silk of the Satu.rni.idae is usually coarse, 
and is known as Tusser or Tussore 2 silk. 

The larvae of this family are as remarkable as the imagos, being 
furnished with spine-bearing tubercles or warts, or long fleshy 
processes ; the colours are frequently beautiful. The caterpillar of 
Attacus atlas (Fig. 187) is pale olive-green and lavender, and has a 
peculiar, conspicuous, red mark on each flank close to the clasper. 

About seventy genera and several hundred species are already 
known of this interesting family. They are widely distributed 
on the globe, though there are but few in Australia. Our 
only British species, the Emperor moth, Saturn i a pavonia, is 
by no means rare, and its larva is a beautiful object ; bright 
green with conspicuous tubercles of a rosy, or yellow, colour. It 
affects an unusual variety of food-plants, sloe and heather being- 
favourites ; the writer has found it at "\Vicken flourishing on the 
leaves of the yellow water-lily. Although the Emperor moth is 
one of the largest of our native Lepidopterous Insects, it is one 
of the smallest of the Saturniidae. 

The family Hemileucidae of Packard is included at present in 
Saturniidae. 

Fam. 4. Brahmaeidae. The species forming the genus 
lii-uli iiiiica, have been placed in various families, and are 
now treated by Hampson as a family apart, distinguished 
from Saturniidae by the presence of a proboscis. They are 
magnificent, large moths, of sombre colours, but with complex 
patterns on the wings, looking as if intended as designs for 



r. Xntural. xii. 1878, p. 379. 
2 Cotes, "Wild Silk Insects of India," I ml. Mus. Xotcs, ii. No. 2, 1891, 
15 plates. 



vi HETEROCERA BRAHMAEIDAE^-BOMBYCIDAE 375 

upholstery. About fifteen species are recognised ; the geo- 
graphical distribution is remarkable ; consisting of a comparatively 
narrow belt extending across the Old World from Japan to West 
Africa, including Asia Minor and the shores of the Caspian Sea. 
Little has been recorded as to the life-histories of these Insects. 
The larva is said to have the second and third segments swollen 
and armed with a pair of lateral spines projecting forwards. A 
cocoon is not formed. 

Fam. 5. Ceratocampidae. This is a small family. They are 
fine moths peculiar to the New World, and known principally by 
scattered notices in the works of North American entomologists. 
Seven genera and about sixty species are known. The chief 
genus is Citlieronia. Some of the larvae are remarkable, being 
armed with large and complex spines. A cocoon is not formed. 

Fam. 6. Bombycidae. - In entomological literature this 
name has a very uncertain meaning, as it has been applied to 
diverse groups ; even at present the name is frequently used for 
the Lasiocampidae. We apply it to the inconsiderable family 
of true silkworm moths. They are comparatively small and 
uninteresting Insects in both the larval and imaginal instars : 
1 tut the cocoons formed by the well-known silkworm are of great 
value, and some other species form similar structures that are of 
more or less value for commercial purposes. The silkworm has 
been domesticated for an enormous period, and is consequently 
now very widely spread over the earth's surface ; opinions differ 
as to its real home, some thinking it came originally from 
Northern China, while others believe Bengal to have been its 
native habitat. The silkworm is properly called Bombyx mori, 
but perhaps it is as often styled Sericaria mori. Besides being 
of so great a value in commerce, this Insect has become an 
important object of investigation as to anatomy, physiology and 
development. Its domestication has probably been accompanied 
by a certain amount of change in habits and instincts, the 
creature having apparently lost its appreciation of freedom and 
its power of flight ; it is also said to lie helpless in certain 
respects when placed on trees in the larval state ; but the import- 
ance of these points has been perhaps somewhat exaggerated. 1 

Although the family Bombycidae is very widely distributed 
in the warmer regions of the world, it includes only 15 or 20 

1 See on this subject Perez, Act. 'Soc. Bordeaux, xlvii. 1894, p. 236, etc. 



3/6 LEPIDOPTERA 



CHAP. 



genera, and none of them have many species. The Mustiliidae 
of some entomologists are included here. Like the Saturniidae, 
the Bombycidae are destitute of proboscis and of frenulum to the 
wings, but they possess two or three internal nervures on the 
hind wing instead of the single one existing in Saturniidae. 

Fam. 1. Eupterotidae (Striphnopterygidae of Aurivillius).- 
This family has only recently been separated from Lasiocampidae ; 
its members, however, possess a frenulum ; while none is present 
in the larger family mentioned. Its limits are still uncertain, 
but it includes several extremely interesting forms. The larvae 
of the European processionary moth, Cn<'f/ii><'<///ipa2}rocessionea,&Te 
social in habits ; they sometimes occur in very large numbers, and 
march in columns of peculiar form, each band being headed by a 
leader in front, and the column gradually becoming broader. It 
is thought that the leader spins a thread as he goes on, and that 
the lateral leaders of the succeeding files fasten the threads they 
spin to that of the first individual, and in this way all are 
brought into unison. The hairs of these caterpillars are abun- 
dant, and produce great irritation to the skin and mucous mem- 
brane of any one unlucky enough to come into too close contact 
with the creatures. This property is, however, not confined to the 
hairs of the processionary moths, but is shared to a greater or 
less extent by the hairs of various other caterpillars of this 
division of Lepidoptera. In some cases the irritation is believed 
to be due to the form of the hair or spine, which may be barbed 
or otherwise peculiar in form. It is also thought that in some 
cases a poisonous liquid is contained in the spine. 

The larvae of other forms have the habit of forming dense 
webs, more or less baglike, for common habitation by a great 
number of caterpillars, and they afterwards spin their cocoons 
inside these receptacles. This has been ascertained to occur in 
the case of several species of the genus Aiutplie, as has been 
described and illustrated by Dr. Fischer, 1 Lord Walsiugham, 2 and 
Dr. Holland. 3 The structures are said to be conspicuous objects 
on trees in some parts of Africa. The common dwelling of this 
kind formed by the caterpillars otHypsoidcs radama in Madagascar 
is said to be several feet in length ; but the structures of most 
of the other species are of much smaller size. 



1 Iln-Iiii. cnt. Zeitschr. xxvii. 1883. p. 9. 
2 7>. Linn. Soc. ser. 2. ii. 1885, \>. 421. 3 Psyche, vi. 1893, p. 385. 



vi MOTHS EUPTEROTIDAE PEROPHORIDAE 377 



The larvae of the South American genus Pahistra, though 
hairy like other Eupterotid caterpillars, are aquatic in their 
habits, and swim by coiling themselves and making movements 
of extension ; the hair on the back is in the form of dense 
brushes, but at the sides of the body it is longer and more 
remote : when the creatures come to the surface which is but 
rarely the dorsal brushes are quite dry, while the lateral hairs 
are wet. The stigmata are extremely small, and the mode of 
respiration is not fully known. It was noticed that when 
taken out of the water, and walking in the open air, these 
caterpillars have but little power of maintaining their 
equilibrium. They pupate beneath the water in a singular 
manner : a first one having formed its cocoon, others come suc- 
cessively and add theirs to it so as to form a mass. 1 Another 
species of P/dnstra, P. ljunneisteri , Berg, 2 is also believed to 
breathe by means of air entangled in its long clothing ; it 
comes to the surface occasionally, to renew the supply ; the 
hairs of the shorter brushes are each swollen at the extremity, 
but whether this may be in connexion with respiration is not 
known. This species pupates out of the water, between the 
leaves of plants. 

Di/jili i/i tarquinia is remarkable on account of the great 
difference of colour and appearance in the two sexes. In the 
Australian genus Marane the abdomen is densely tufted at the 
extremity with hair of a different colour. 

Fam. 8. Perophoridae. The moths of the genus Peropliora 
have for long been an enigma to systematists, and have been 
placed as abnormal members of Psychidae or of Drepauidae, 
In it Packard now treats them as a distinct family. The larvae 
display no signs of any social instincts, but, on the contrary, each 
one forms a little dwelling for itself. Some twenty species of 
!' i-ujilnn'ii are now known; they inhabit a large part of the 
Xew World, extending from Minnesota to Buenos Aires. The 
habits of P. melslieimeri have been described by Harris, 
Packard '' and Xewman, and those of P. batesi by Xewrnan. 4 The 
larva is very peculiar ; there is a flexible pair of appendages on the 

1 Bar and Laboulbene, Ann. Soc. ent. France, (v. ) iii. 1873, p. 300. 
- "r >>. ."' . vii. 1877, p. 181 ; and Ent. Zeit. Mett-in, xxxix. 1878, p. 221 ; and 
xliv. 1883, p. 402. 

3 Ami. New }~rk Ac. viii. 1893, p. 48. 

4 Tr. cat. Soc. Loitdu/i, n.s. iii. 1854, p. 1. 



3/8 LEPIDOPTERA CHAP. 

head, the use of which is unknown ; 1 they arise by slender stalks 
behind and above the eyes, are about as long as the head, and 
are easily broken oft'. After hatching, the young larva, when it 
begins to feed, fastens two leaves together with silk threads, and 
so feeds after the fashion of a Tortricid, rather than a case-making, 
larva. Subsequently, however, the caterpillar entirely detaches 
two pieces of leaves and fastens them together at the edges, thus 
constructing a case that it lives in, and carries about ; it can 
readily leave the case and afterwards return to it. When at 
rest, the larva relieves itself from the effort of supporting this 
case by the device of fastening it to a leaf with a few silken 
threads ; when the creature wished to start again, " it came out 
and bit off these threads close to the case." Subsequently it 
changes inside the case to a pupa armed with transverse rows of 
teeth, like so many other pupae that are capable of a certain 
amount of movement. The larva is of broad, short, peculiar 
form, and is said to be very bold in defending itself when at- 
tacked. The moth is somewhat like the silkworm moth, though 
df a more tawny colour. Newman does not allude to any 
cephalic appendages as existing in the larva of P. batesi. 
If we accept the eggs figured and described by Snellen, 2 as those 
of P. Itatesi, it is possible that this Insect possesses a peculiar 
mode of oviposition, the eggs being placed one on the other, so 
as to form an outstanding string ; but we think this example 
probably abnormal; the mode is not shared by P. mclsheimcri. 
The genus Lacosoma is considered by Packard to be an ally of 
rr/'iiji/toi-tt. The caterpillar of L. chiridota doubles a leaf at the 
mid-rib and fastens the two edges together, thus forming an un- 
symmetrical case. Many larvae of Microlepidoptera do something 
like this, but the Lacosoma cuts off the habitation thus formed and 
carries it about. Packard says it may have descended from 
ancestors with ordinary habits and that certain peculiar obsolete 
markings oil the body of the caterpillar may be indications of this. 3 
The Argentinian Insect Mam illo curtisea 4 is also probably an ally 

1 Dyar says, "We may surmise that it is to present a terrifying appearance to- 
ward small enemies." He calls the Insect both Perophora and Cicinnus, melshei- 
uti'i'i, and states that it belongs [according to the larva] to Tineidae ; the appendages 
he considers to be enormously developed setae. /. X. )'ork ent. Soc. iv. 1896, p. 92. 

: TijJs^tr. Ent. xxxviii. 1895, p. 56, PI. 4. 

3 Ann. X<:v York Ac. viii. 1893, p. 48. 

4 Weyenbergh, Tijdschr. Ent. xvii. 1874, p. 220, PL xiii. 



VI 



HETEROCERA PEROPHORIDAE 



379 



of Lacosoma. The caterpillar of this moth spins a dwelling 
for itself, and is remarkable from the bright colour of the 
thoracic segments, the following somites being colourless ; the 
head bears a pair of large processes, quite different from 
those figured bv Harris. The moth itself is very Geometric!- 

.~* i/ 

like in colour and form. This species is now assigned to Pero- 




FIG. 188. Larvae of Hammock-moth, Perophora sanguinolenta, projecting from 
Hammocks, built from their o\vu excrement. South America. (After Jones.) 



/i/mrrt, but it seems to be very doubtful whether many of the 
species placed in this genus really belong to it. The diversity 
of habits and instincts evinced by these moths of exceptional 
modes of life, but considered to be closely allied, is very interest- 
ing. The most remarkable of all is the Hammock-moth, Pero- 
phora sanguinolenta, of the centre of South America, the larva of 
which constructs its portable habitations out of its own excre- 



38o 



LEPIDOPTERA 



CHAP. 



ment, which is of peculiar form, specially suitable for the 
purpose. The caterpillar, when wishing to enlarge its case, builds 
it up from excrement " flattened at the sides, so as to adapt it 
for building purposes." 

Fam. 9. Sphingidae (Hawk -moths}. A very important 
family of moths of large or moderate size. They have a pro- 
boscis which is frequently very long ; there is a frenulum ; the 
body is stouter than in most other Lepidoptera, and the wings 
are of small superficies in comparison with it ; the antennae are 
somewhat peculiar, having a thick, solid appearance, pointed at 
the tip. This is usually somewhat hooked, and bears a few hairs. 
In the males the antennae are formed in a manner specially 
characteristic of the family. In section, each joint shows a 
chitinous process on the under side (Fig. 189, A), forming with 
that of the other joints a continuous ridge, and on each 

Fie. 189. Antennae 
of Sphingidae. A, 
One joint of an- 

tenna Of (.'ItlH-fll- 

caiiijin >! ft' inn- $, 
enlarged ; B, three 
joints of antenna of 





seen from one side, 
and enlarged. 



side of this ridge there exists a series of short, delicate " cilia " 
arranged in a very beautiful manner (Fig. 189, B). This structure, 
with some modifications, appears to be usually present in the 
family ; it attains a very perfect development in cases where 
the tips of two rows of cilia bend towards one another, meeting 
so as to form an arched cavity. This structure is different from 
what occurs in the males of other families of Lepidoptera, for 
though cilia are very common, they are usually placed either on 
two projections from the body of the antennae (instead of on the 
two sides of a single projection), or there is but a single whorl, or 
set, of them on each joint (Catoccda, etc.). The front wings are 
usually pointed at the tip, and are long in proportion to their 
width: but in the Smerinthini they are of different form, with 
the outer margin scalloped ; the hind wings are remarkably 
small : the abdomen is frequently pointed, but in the Macro- 
glossini, or Humming-bird hawk-moths, it is furnished at the 

1 Jones. P. Liverpool Hoc. xxxiii. 1879, \>. Ixxvii. 



VI 



HETEROCERA HAWK-MOTHS 



tip with a tuft, or with two tufts, of dense, long scales, capable 
of expansion. 

The larvae are remarkable for their colours and form. The 
anterior segments are attenuated, but are capable of great retrac- 
tion, so that in repose (Fig. 190, A) this shape is concealed by 
the curious attitudes that are assumed. There is in nearly 
all cases a conspicuous horn on the eleventh segment, and the body 
at the extremity behind the horn is so much modified that the 
terminal two segments look like little more than a pair of large 
claspers. In the Choerocampini, the thoracic segments are 
retractile, and can be withdrawn into the more or less inflated 
fourth segment, and give the creature somewhat the appearance 





FIG. 190. Larva of 
the Poplar Hawk- 
inoth, Smerinthits 
populi. x 1. A, 
in repose ; B, iu 
movement. 



of a miniature hooded snake. The larvae of Sphingidae do not 
bear any conspicuous hairs except during the first instar. They 
do not spin cocoons, but bury themselves in the earth. The 
pupa is remarkable from the deep cleft that exists to admit air 
to the first spiracle, and for a deep depression on each side of the 
anterior part of abdominal segments 5-7 ; in some cases the 
proboscis projects on the breast somewhat like the handle of a 
pitcher. 

A great deal has been written on the colours, markings, and 
attitudes of Sphingid larvae, and many interesting facts h;i\c 
been brought to light. We may refer the reader to the writings 
of Weismann J and Poulton, 2 without, however, recommending 
him to place an implicit confidence in their somewlmt 
metaphysical disquisitions ; for the views there shadowed will 

1 Studies in the Theory of Descent, part 2, London, 1881. 
1 Tr. ent. Soc. London, 1885 and 1886. 



382 LEPIDOPTERA CHAP. 



necessarily became much modified with the advance of exact 
knowledge. It is certain that the position assumed by the same 
individual varies much according to age, and to the interval 
since the last moult ; sometimes the attitude is much more remark- 
able than that shown in Figure 190, A, for the anterior seg- 
ments are held erect, as well as contracted, the front part 
of the body being curled, and the Insect supported by the 
claspers and two pairs only of the abdominal feet. There is, 
too, a considerable difference in colour before and after an 
ecdysis. Piepers, who has had a long experience among Sphingid 
larvae in Java, considers that much of what has been written as 
to the protective value of their colours and attitudes, is mere 
fancy, and wild generalisation. 1 

Sphingidae have been recorded as capable of producing sounds 
in the larval and pupal, as well as in the perfect, instars ; but 
the method in which this is done has not been ascertained, 
except in the case of the imago of the Death's-head moth, 
which is well known to emit a very audible cry when not on the 
wing; in this case it is highly probable that the method is the 
friction of the palpi against the proboscis, as stated by Reaumur 
and Landois ; the inner face of the palp is said to be marked in 
this case with fine ridges or striae. 

Fam. 10. Cocytiidae. A single genus constitutes this family, 
and there are only three or four species known ; they come from 




FIG. 191. Cocytin i/ni-rillii. New Guinea. (After BoisJuval. ) 

the region of Xew Guinea, whence the first was brought by 

D'Urville nearly a hundred years ago. They are still amongst 

1 Tijdschr. Ent. xl. 1897, pp. 127-103, 4 plates. 



VI 



HETEROCERA COCYTIIDAE NOTODONTIDAE 



333 



the rarest of Insects. Nothing is known as to their life-histories. 
In appearance they somewhat remind us of the Bee-hawk moths and 
Zygaenidae. Butler says l the family is characterised by the palpi, 
which differ much in the two sexes, and by the antennae resem- 
bling those of Castniidae or Hesperiidae. The form, transparency, 
and coloration of the wings reminds one vividly of the Sphingid 
genus Hemaris ; the nervuration is somewhat like that of Hemaris, 
but has certain features of Zygaenidae. Butler places the family 
between Agaristidae and Zygaenidae. 

Fam. 11. Notodontidae (Prominents, Puss-moths, etc.).- This 
is one of the most extensive of the families of Bombyces ; it con- 
sists in larger part of obscure-coloured moths, somewhat like the 
ordinary Noctuidae of temperate regions ; to which family the ISToto- 
donts are indeed considered to be very closely allied. The family 
contains, however, some very remarkable forms. Tarsolepis has 
an elongate body, 
terminated (in the 
female of T. ful- 
! I a i- if era) by a very 
conspicuous tuft 
of enormously 
long, battledore 
scales ; while in 
the male of T. 
sommeri the hind 
legs are provided 
with an append- 
age of beautiful, 
roseate hairs. A 
few of the larger 
kinds bear a con- 
siderable resem- 
blance in form and 

proportions to the Sphingidae. Some of the larvae are most inter- 
esting objects ; the Puss-moth caterpillar, the Lobster, and t he- 
Dragon larvae are of such strange forms that they have already 
interested several generations of observers. The Puss-moth is 
common in the southern half of England ; its caterpillar (Fig. 192) 
has, instead of the claspers, a pair of tubes in which are concealed 

1 Tr. ent. >'oc. London, 1884, p. 351. 




FIG. 192. Centra vinitJa (Pnss-moth) caterpillar. A, Moult- 
ing ; B, the same individual a i'e\v hours after the moult. 



384 LEPIDOPTERA CHAP. 

two long, flexible whips, capable of being thrust out, and with- 
drawn, with rapidity. The structure and the mode of action of 
these flagella have been well elucidated by Professor Poulton. 1 
The flagella are to be considered as actual prolongations of the 
receptacle in which each is placed, though they are of very 
different texture therefrom ; they are everted by blood-pressure 
and drawn in by muscular action ; this latter function is very 
perfectly accomplished, the amount of relaxation and contraction 
of the muscle being very great. It has been maintained that 
the whips have arisen as arms of protection against the attacks 
of Ichneumon flies ; observation shows, however, that the pro- 
portion of these " protected " Insects destroyed by enemies of this 
sort is quite as large as it is in the case of forms that are not so 
protected. The Puss-moth larva is also believed to be protected 
by terrifying attitudes, 2 as well as by ejection (like so many other 
larvae and insects generally) of fluid. There is no reason for 
believing that these larvae are less eaten than others, and con- 
sequently a further hypothesis has been proposed, to the effect 
that if they had not acquired these means of defence they would 
have been exterminated altogether. This supposition is con- 
sidered to account for their acquiring the defence by means of 
natural selection ; realising the dictum of D. O'Phace, Esq. 

Some flossifers think that a fakkilty's granted, 
The ininnit it's proved to be thoroughly wanted. 

When the Puss-moth caterpillar is full grown it spins a peculiar 
cocoon of a solid and impervious nature, which it manages to make 
look very like the spots, crevices, or other places amongst which 
it is located ; in this prison the creature remains fur nine or ten 
months by far the larger part of its existence. When it has 
changed to a moth it has to escape from the cell in which it so 
effectually confined itself. This is effected by the cocoon being 
tli inner in front of the head of the moth, and by the emission 
from the alimentary canal of a fluid that softens the cocoon at 
the spot alluded to. Mr. Latter has ascertained 3 that this fluid 
is strongly alkaline, and contains potassium hydroxide. The 
front of the head of the moth is provided with a shield, consist- 
ing of a portion of the pupa shell, which enables the moth to 

1 Tr. ent. Soc. London, 1887, p. 297, PI. x. 
2 See Poulton, Tr. cut. Soc. London, 1886, etc. 3 Op. tit. 1895, p. 399. 



vi HETEROCERA NOTODONTIDAE 385 

push through in safety, and at the same time protects the head 
from the emitted fluid. Figure 192 shows the great change that 
occurs in the period of a few hours in the size of the head of the 
larva, as well as in that of the spiracles : in A the old spiracles 
are seen surrounded by the much larger new orifices, which are 
at the moment of moulting quite visible through the skin that is 
about to be cast off. 

The caterpillar of the Lobster-moth, Stauropus fayi, is more 
remarkable than that of the Puss-moth, but is unfortunately 
very rare. It has remarkably long thoracic legs, the abdomen is 
swollen at the tip, and instead of the terminal claspers has two 
long slender processes. The effect of these peculiarities is greatly 
enhanced by the extraordinary attitude assumed by the cater- 
pillar, which holds the first five segments erect, with the second 
and third pairs of thoracic legs outstretched ; the swollen terminal 
segment is also held erect. Hermann Muller states l that when 
seen from the front this caterpillar looks like a spider, and also 
that when alarmed it moves the long legs after the fashion of 
an Arachnid. He believes that it is thus effectually protected 
from the attacks of Ichneumons. Birchall says 2 that the young- 
larva, when at rest, closely resembles, in colour and outline, 
one of the twigs of beech with unopened buds, on which it 
frequently stations itself; and that, when feeding, its likeness 
to a great earwig or to a 8ta})hylinus is very striking. Others 
say that this caterpillar resembles a dead and crumpled beech 
leaf. 

The larva of Hybocampa milhauseri the Dragon of old Sepp 
is highly remarkable. When young it has grand lateral 
horns in front, and a dorsal row ; as it grows the lateral 
horns disappear. Dr. Chapman says 3 that he could not under- 
stand at first why any larva should have such remarkable angular 
outlines, curiously conspicuous corners and humps. But he after- 
wards found that the creature exactly resembled a curled oak 
leaf, eaten and abandoned by a Tortrix larva. This caterpillar 
also constructs an elaborate cocoon from which the moth escapes 
by an operation performed by the pupa, which is provided with 
two hard spines, called by Dr. Chapman sardine-openers. " By 
a lateral rotatory movement of the pupa, which obtains its fulcrum 

1 P. eat. Sue. London, 1880, p. iii. - Ent. Monthly May. xiii. 1877, p. 231. 

3 Entomologist, xxiii. 1890, p. 92. 
VOL. VI 2 C 



386 LEPIDOPTERA 



CHAP. 



from the tightness with which it is grasped by the cocoon, it 
traverses over and over again " the same part of the cocoon till 
it is cut through ; at the same time the spines act as guides to 
a fluid which is emitted so as to soften the part that has to lie 
sundered. 

Though many other larvae of Xotodontidae are of most 
curious form and assume remarkable attitudes, yet this is not 
the case with all, and some are quite ordinary and like the cater- 
pillars of common Xoctuidae. This is the case with the species 
Rlicgmatophila alpina we have selected to illustrate the meta- 
morphosis of the Order (Fig. 157). Those who wish to form an 
idea of the variety of larval forms in this family will do well 
to refer to Packard's beautiful volume on the Xorth American 
forms. 1 The family has a very wide distribution, but is absent 
from Xew Zealand and Polynesia, and appears to be but poorly 
represented in Australia. In Britain we have about two dozen 
species. 

Fam. 12. Cymatophoridae. A small family of nocturnal 
-noths that connect the Bombyces with the X 1 octuids ; they are 
usually associated with the latter, but are widely separated in 
Hampson's arrangement because of a slight difference of nervura- 
tion, nervule 5 being nearer to 6 than to 4, whereas in Xoctuidae 
the reverse is the case. The Insects, however, in certain respects 
approach the Xotodontidae, and are of interest if only as showing 
that the linear sequences we adopt in books are necessarily con- 
ventional, and to some extent deceptive. We have three genera 
in Britain ; our pretty Peach-blossom, Tkyatira batis, and the very 
different Buff-arches, T. derasa, being among them. Meyrick 
denies any connexion of this group with Xoctuidae, and in his 
nomenclature Cymatophora becomes Polyploca, and the family, 
consequently, Polypi ocidae. 

Fam. 13. Sesiidae or Aegeriidae (Clear-lyings'). A family 
of comparatively small extent ; its members have frequently one 
or both pairs of wings in large part free from scales, the tip of 
the body tufted, the hind legs of one sex peculiar. The size is 
usually small, but in the largest forms the measurement may be 
but little less than two inches across the expanded wings. 
The pupa is of the kind classed as " incompletae " by Chapman, 
the appendages not being firmly glued to the body, and much 

1 Mem. Ac. IJ'ashinyton, vii. 1895, 290 pp., 49 plates. 



VI 



HETEROCERA SESIIDAE TINAEGERIIDAE 



337 



mobility existing ; an " eye-collar " is present, and the segments 
of the abdomen are armed with series of teeth. The larva 
is a concealed feeder, nearly naked and colourless, but with 
the legs normal in number three thoracic, four abdominal pairs 
of feet, and the terminal claspers ; these are sometimes but 
poorly developed ; the larvae have a greater or less resemblance 
to those of Longicorn beetles, the habits of which they share. The 
family was formerly associated with the Sphingidae, with which 
it has no true relationship ; it is more closely allied to the 
Tineidae. Some of the species have a certain resemblance to 
Hymenoptera, which is probably in most, if not in all cases 
merely incidental. The proper position of the family was pointed 
out by Butler, 1 but he did not distinguish it from Tinaegeriidae. 
Meyrick calls the family Aegeriadae, and places it in his series 
Tineina. 

We have two genera of these Clear -wings in Britain. 
They are Trocliilium (called variously Sesia, S-plti:<'i<', and 
Aegeria), with two species of comparatively large size, and 
(called variously Trocliilium and 
Aegeria), with nearly a dozen species 
of smaller size. A third genus, 
Sciapteron, is doubtfully native with 
us. They are much prized by col- 
lectors on account of the rarity of 
the Insects and their great differ- 
ence in appearance from our other 
native Lepidoptera. 

Fam. 14. Tinaegeriidae. This 
is one of the least known of the 
families of Lepidoptera, and has only 
recently been distinguished from 
Sesiidae. It is entirely exotic, and 
our knowledge of it is principally FIG. 

.f i- Africa. (After Walsingham.) 

due to Lord Walsingham.- Nothing 

is known as to the life-histories, except that it has been stated 
by Staintoii that a larva feeds in webs on shoots of a shrub of 
the genus Clerodendron. The family is widely distributed, but 
its metropolis will probably prove to be the tropics of Africa. It 
is of considerable interest as showing that the Sesiidae really 
1 Tr. cat. Soc. London, ]878, p. 121, PI. v. " Op. cit. 1SS9, pp. 1-40, 6 plates. 




388 LEPIDOPTERA 



CHAP. 



belong to the Tineid series of moths. ' The species we figure (Fig. 
193) has a character otherwise peculiar to Sesiidae in the wings 
being inserted 011 the thorax remote from the head a feature we 
do not find in the Tineidae proper ; while on the other hand it 
has the lung wing-fringes, and the shape of the wings that are 
characteristic of Tineidae. It is worth mentioning that though 
these Insects are of excessive rarity and very peculiar, there exists 
in the Solomon Islands l a species distinct from, though at first 
sight excessively similar to, the S. African one we figure. 

Fam. 15. Syntomidae. This family has usually been asso- 
ciated with the Zygaenidae. It includes a large number of 
moths having, as a rule, in external appearance little to distin- 
guish them from the family named. Many of them are of gaudy 
colours, and probably of diurnal, but somewhat sedentary, habits. 
The wings are less ample than usual, the hind pair frequently 
very small, so that the Insects have somewhat the proportions of 
Hymenoptera. In some cases the resemblance is made more 
remarkable by the fact that the wings are transparent and bare 
of scales, or have scales only at the margins, so as to be like the 
wings of Hymenoptera. Not less remarkable is the fact that 
these Insects use the body itself for the purposes of adornment 
or display ; thus adopting a system prevalent in the Hymenop- 
tera, rather than that of their own Order, where the rule is that 
the wings are more ornamented than the body. In many 
cases the shape of the body is so very different from the normal 
that the disposition of the organs of life in the interior of the 
body must be materially affected. In some genera, such as Andre- 
nimorpha, the form, colour and attitude of the body and some 
of the limbs are plainly similar to Hymenoptera. These Insects 
have a highly-developed frenulum, retinaculum, and proboscis ; 
bipectinate antennae in the male, a complex organ at the base 
of the abdomen on each side, and are in fact highly-developed 
forms, except perhaps as regards the structures in connexion 
with flight. 

Unfortunately little or nothing is known as to the habits 
and metamorphoses of these extraordinary creatures, but it is no 
doubt to them Seitz referred in saying, " How far one may be 
deceived by appearances of a mimetic nature can only be com- 
prehended by visiting the tropics; in this part of the world 

1 \Valsinghaui, Op. at.. 1839. c. p. 21. 



vi SYNTOMIDAE MIMICRY 389 



[Europe] one is prepared by knowledge gained from books for 
the appearance Sesia presents. Had one no knowledge of this 
sort as to Sesiidae he would actually in the field [in Brazil] over- 
look dozens of these little creatures without being aware of his 
deception. The surprise at finding a quite different being in the 
net from what one believes he has caught occurs daily in Brazil, 
so rich in Lepidoptera." The same intelligent observer says 2 
that a species of Macrocneme was observed by him to be exactly 
like one of the blue wasps of the genus Pepsis. 

One remarkable point in these Hyrnenopteroid Syntomids 
is their complete dissimilarity from their immediate allies. 
They resemble very different Hymenoptera ; and not only 
stinging Hymenoptera ; the Sessiliventres have a large share 
of their attentions ; the numerous species of Dycladia partaking 
the appearance of the South American Sawflies in a wonderful 
manner. Bees, Wasps of the most different kinds, and a variety 
of Sawflies are beautifully paralleled, if one may use such an 
expression, by these Syntomids. That shown in Fig. 194 has the 
abdomen formed like that of a Petiolate Hymenopteroii ; the 
base of this part, moreover, resembles in a remarkable manner 
the " median segment " of that Order. The constriction is, how- 
ever, placed not at the base of the abdomen but beyond the second 
segment. Thus the structure is not morphologically similar to 
that of the Hymenoptera, for the median segment of Aculeate 
Hymenoptera consists of only one abdominal segment, while in 
this moth the corresponding part is formed of two segments. 
Though anatomically inexact, the resemblance is, as to propor- 
tions, correct ; and those who delight in the use of the imagination 
will see that had the moth used only one segment for the imita- 
tion, the result would have been less successful owing to insufficient 
size. In his very interesting account of some Brazilian Syn- 
tomids, 3 Seitz describes a species of Trichura provided with a 
long appendage that is held straight backwards during life ; . and 
he informs us that this creature resembles a female Ichneumon, 
the long process looking like the elongate ovipositor of the 
Hymenopteron. Possibly the species from Demerara we figure 
may resemble an Ichneumon we are not acquainted with, though 
its colour and form rather suggest a likeness to an Aculeate. 

1 Ent. Zcit. Stettin, Ivi. 1895, p. 233. - Op. tit. li. 1890, p. 261. 

3 Ent. Zcit. Stettin, li., 1890, p. 263. 



390 LEriDOPTERA CHAP. 

This case of resemblance is of the most noteworthy character, 
for an appendage of this kind in a Lepidopterous Insect is 
without parallel, and is almost equivalent to the production of 
a new structure. An interesting feature of the case is that 

O 

Ichneumonidae do not sting, and there is no evident reason why 
the enemies of the moth should be particularly afraid of an 
ovipositor. 

The larvae appear to be in form somewhat like those of 
Zygaenidae ; but with the same sort of remarkable clothing, in 
the form of tufts and brushes, that we find in Lymantriidae. A 




FIG. 194. Trichura, sp. x j. Demerara. 

cocoon is formed. In Britain no member of this family is to 
be met with, but Nudia ancilla may formerly have been a 
native; Syntomis phegea has occurred here; probably an escaped 
example that had been introduced in one of its earlier stages. 

Fam. 16. Zygaenidae (-Bur net-Moths). This family is one 
about the limits and characters of which much difference of 
opinion prevails. As exemplified by our Burnet- moths it is 
characterised (in addition to the points given in the table) by 
the peculiar, flexible antennae; these are a little thicker before 
the tip, but are curved and pointed at the extremity, and without 
pectinations in the male. There is an elongate proboscis ; 
1 dadder-like organs at the sides of the first abdominal segment 
are not present. The pupa is softer than is usual in the Macro- 
lepidoptera, and the parts are less firmly fixed together, so that 
unusual mobility exists; six of the intersegment al mrmbranes 



vi HETEROCERA ZYGAENIDAE 391 

are free, and the abdomen has much power of movement ; there 
is no eye-collar ; the antennae, hind legs, and proboscis -tips 
stretch backwards as far as the fifth or sixth abdominal seg- 
ment, the tips being quite free ; on the dorsal plates of the 
abdomen there are rows of minute elevations reminding one of 
the teeth existing in pupae that live in stems or galleries. This 
is altogether a peculiar pupa ; it lives closely enclosed in a small 
hard cocoon, and its great capacity for movement is perhaps con- 
nected with the fact that the pupa itself manages to force its 
way through the cocoon in anticipation of the emergence of the 
moth. This cocoon is fastened tightly to a stem, and is covered 
with a substance that gives it a glazed appearance. The larvae 
are objects of a baggy nature, with inferior coloration, consisting 
of large dark blotches on a light ground, and without any 
remarkable development of their somewhat feeble system of 
hairs. Numerous small moths from the tropics are assigned 
to the family ; they are most of them conspicuously marked 
and coloured, and like our Burnets are probably diurnal. 

The family Chalcosiidae is reduced by Hampson to the 
position of a sub-family of Zygaenidae. It consists of a large 
variety of diurnal moths of varied and brilliant colours, with an 
expanse of wing large in 
comparison with the typi- 
cal Zygaenae, and with 
the antennae pectinate or 
flabellate to the tip. Some 
of these Insects (which are 
as conspicuous as possible 

in appearance, at any rate . , , ~^r~ "" w . 

J FIG. 195. Hampsoma putchermma. Wings on 

ill a Cabinet, the East right side detached and denuded to show 

Indian Cadphises moorei ^miration. India. (After Swinhoe.) 

e.g.} are considered to be destitute of any special " protection." 
Histia- is a genus of remarkable cruciform moths, of a mixture 
of black and metallic colours, with carmine - tinted bodies. 
Hu.mpsonict, pulcherrima (East India) is a curious moth of butter- 
fly form and coloration, red and black with yellow patches, and 
with some of the nervules distorted, as if they had been forced 
a] tart in certain spots in order to accommodate these patches. 

Two or three hundred species of Chalcosiidae are recorded. 
They are specially characteristic of the Indo-Malayan region. 




39 2 LEPIDOPTERA CHAP. 

Fam. 17. Himantopteridae (Thymaridae of some autho- 
rities) are placed by Hainpson in the sub-family Phaudinae of 
Zygaenidae characterised by the absence of the mouth-parts. 
The Himantopteridae are small moths, and have the scales on 
the wings very imperfect and hair-like; the hind wings form 
long slender tails, so that the Insects scarcely look like moths. 
They are peculiar to India and Africa. In the South African 
genus Dianeura (belonging really to rhaudinae) also the wings 
are scaleless and nearly transparent. 

Fam. 18. Heterogynidae. Consists of the single genus 
Hcterogynis which has hitherto been found only in the south of 
Europe. This is an important form connecting Zygaenidae and 
Psychidae. The larvae resemble those of Zygaena, and construct 
an oval cocoon for their metamorphosis. The male issues as a 
small moth of smoky colour, the scales being but imperfect ; 
the female chrysalis show's no trace of any appendages, and the 
imago is practically a maggot, and never leaves the cocoon ; in it 
she deposits her eggs, and the young larvae hatch there. 1 

Fam. 19. Psychidae. Small, or moderate-sized moths, with 
imperfect scales, and little or no colour beyond certain shades of 
duskiness ; the sexes very different, the female being wingless and 
sometimes quite maggot-like ; the male often with remarkable, 
bipectinate antennae, the branches sometimes very long and 
flexible. Larva inhabiting a case that it carries about. This 
family consists of Insects unattractive in appearance but present- 
ing some points of great interest. It is frequently stated that 
the Psychidae are destitute of scales, but Heylaerts states' 2 that, 
in addition to hairs, scales of a more or less imperfect formation 
are present in all, but that they are, like those of some Sphingidae 
(Macroglossa), very easily detached. There is much difference in 
the females, some having well-developed legs, while others are 
not only apterous, but are bare and destitute of appendages 
like a maggot, while in certain cases (Fig. 196, G), the head 
is reduced in size and is of peculiar form so as to make the 
Insect look really like the larva of one of the parasitic Diptera. 
These females never leave their cases, but deposit their eggs 

1 For details as to habits, etc., see Ramlmr, Ann. Soc. cut. France, v. 1836, 
p. 577 ; and Graslin, <>p. <//. xix. 1850, p. 396. 

2 Monograph of European T.sycliidae, Ann. Soc. cut. Bclyiqnc, xxv. 1881, 
p. 29, etc. 



VI 



HETEROCERA PSYCHIDAE 



393 



therein, and inside, also, their former pupa-skin ; and here the 
young hatch ; the peculiar little larvae are very numerous, and it is 
suggested that they make a first meal on the body of their parent, 
but this we believe has not been satisfactorily ascertained. 
Great differences as to the condition of the legs, antennae, etc., 
are said to exist in species of the same genus. There is also a 
remarkable diversity in the pupae of the females ; the male sex 
being normal in this respect. Some of the female pupae are 
destitute of wing-sheaths and all other appendages, while others 
are said to possess them, though there are no wings at all in 
the imago (Fumea, e.g.'). 1 Great difficulties attend the study of 
these case-bearing Insects, and several points require careful 




FIG. 196. Metamorphosis of Monda rhabdophora. Ceylon. A, Larva in case, nat. 
size ; B, larva itself, magnified ; C, case of female during pupation ; D, case of male 
during pupation ; E, female pupa, magnified ; F, male moth, nat. size ; G, female 
moth, magnified. ( From unpublished drawings by Mr. E. E. Green). 

reconsideration, amongst them the one we have just mentioned. 
The males fly rapidly in a wild manner, and may sometimes 
be met with in swarms ; their lives are believed to be very brief, 
rarely exceeding a couple of days, and sometimes being limited 
to a few hours. 

The larvae are called basket-worms, and their baskets or 
cases are w T ell worthy of attention. Their variety is remark- 
able ; the most extraordinary are some of the genus Apterona 
Fig. 197, B, which perfectly resemble the shells of Molluscs 
such as snails ; indeed, the specimens in the collection at 
the British Museum were sent there as shells. This case is not, 
like those of other Psychidae, constructed of earth or vegetable 
matter, but is of silk and is in texture and appearance exactly 

1 Heylaerts, op. cit. p. 55. 



394 



LEPIDOPTERA 



CHAP. 




like the surface of a shell. Psyche helix is, according to 
Ingenitzky, 1 found in great numbers near Lake Issyk-kul in 

Central Asia, where the 
larvae feed, in their snail- 
shell-like cases, on a grass, 
just like snails. Only 
females could be reared 
from these larvae. The 
case of Chalia hockingii 
(Fig. 197, C) consists of 
little pieces of wood cut 
to the proper lengths, and 
spirally arranged, so as to 
A c form a construction that 

FIG. 197. Baskets, or cases, of Psychidae. A, WOllld be quite a Credit t> 
Amictaquadrangularis; B Apterona(orCoch- sneeies Til some 

lophora) val-vata; C, Chalm hockingi. 

of the Canephorinae we 

meet with long cylindrical cases, like those of Caddis-worms, or 
of Tineid larvae. 

lliley has given an account of several points in the struc- 
ture and natural history of one of the North American basket- 
or bag- worms, Thyridopteryx ephemeraeformis ; one of his points 
being the manner in which the newly hatched larva forms its 
case.' 2 This question has also been discussed by Packard.' 1 The 
larvae when hatched in unnatural conditions will make use of 
fragments of paper, cork, etc., for the case ; the act of construc- 
tion takes one or two hours, and the larva does not eat till the 
case is completed. It walks in a peculiar manner, the legs of 
the third pair being moved forwards together, as if they were 
the prongs of a fork. 

This family is already one of considerable extent, but its 
study, as already renmrked, is but little advanced. Some 
naturalists are inclined to place it among the Tineidae, but it 
is connected with Zygaenidae by means of Heterogynidae. Mr. 
Meyrick divides it, placing Psyche and Sterrhopteryx (the forms 
representing, according to his ideas, the family Psychidae in 
Britain) in the series Psychina which includes Zygaenidae. He 



l. An;, xx. 1897, p. 473. This is probably Aptcrona crenulclla, or one 
of its varieties. " Hull. U.S. Dcp. Aijric. Ent. x. 1S37, p. 22. 

3 Ann. New ]">/ ./-. viii. IS'i:',, p. 54. 



vi HETEROCERA PSYCHIDAE- COSSIDAE 395 

removes the other British genera, Fumea and Epichnopteryx, to 
Tineidae near Solenobia and Taleporia. The group Canephorinae, 
to which the two genera in question belong, was long since separ- 
ated from Psychidae by Herrich-Schaffer, but this course was 
condemned by Heylaerts. Parthenogenesis has been thought by 
some to occur in numerous species in this family, but Heylaerts 
says that it is limited to Apterona crenulella var. helix, and even 
of this species males are found in certain localities. 

Fam. 20. Cossidae (Goat -Moths, or Carpenter -Worms}. 
Moths of moderate, or rather large size, without proboscis, fre- 
quently with a dense covering of matted, imperfect scales ; the 
pattern being vague. The larvae bore into trees in which they 
often make large burrows, leaving holes from which sap exudes. 
Our common Goat-moth is a good specimen of this family, which 
is a very widely distributed one. The Australian genus Ptilo- 
macra has very large, pectinated antennae in the male. The 
larvae of Cossidae are nearly bare of clothing and are unadorned ; 
they form a slight cocoon of silk mixed with gnawed wood. 
The pupa of the Goat-moth is remarkable for the great develop- 
ment of the rows of teeth on the dorsal aspects of the segments 
of the abdomen, and for the absence of consolidation in this 
part, six of the intersegmental incisions being free, and the 
ventral aspect almost membranous. Very little is known as to 
other pupae of the family. It is believed that the generations 
of these Insects are fewer than usual, the growth of the larva 
occupying a period of two or three years. The larva of Zeuzera 
aesculi forms a temporary cocoon in which it passes a winter- 
sleep, before again feeding in the spring. 1 It is a moot question 
whether the Zeuzeridae should be separated from the Cossidae or 
not. The group includes our Wood-leopard moth, which, like 
many other Zeuzerids, is spotted in a very striking but inartistic 
manner. The position the family Cossidae should occupy in an 
arrangement of the Lepidoptera is a very difficult question. Some 
consider the Insects to be allied to Tortricidae. The wing-nervura- 
tion of Cossus is very peculiar and complex, there being four or 
five cells on the front wing, and three on the hind one. Meyrick 
places Zeuzeridae as a family of his series Psychina, but separ- 
ates Cossidae proper (he calls them Trypauidae) as a family of 
the series Tortricina. 

1 Kalender, Ent. Zcit. Stettin, xxxv. 1874, p. 203. 



396 LEPIDOPTERA CHAP. 

Fam. 21. Arbelidae. Closely allied to Cossidae, but with- 
out frenulum, and with less complex wing-nervures. A small 
family believed to be similar to Cossidae in the life-history. 
The tropical African Arbelidae are considered by Karsch to be 
a distinct family, Hollandiidae. 

Fam. 22. Chrysopolomidae. This family has been estab- 
lished by Dr. Aurivillius l for an African genus, allied in wing- 
nervuration to Cossidae ; the Insects are like Lasiocampidae. 

Fam. 23. Hepialidae (Ghost- and Svrift-Moths). Moths of 
very diverse size, some gigantic ; wings not fitting together well 
at the bases ; without a frenulum ; no proboscis ; the scales 
imperfect ; the nervures complex. The Hepialidae are extremely 
isolated amongst the Lepidoptera ; indeed, they have really no 
allies ; the conclusion that they are connected with the Micro- 
pti'rygidae being certainly erroneous. Although but small in 
numbers only about 150 species being known they exhibit a 
remarkable variety in size and colour. Many are small obscure 
moths, while others are of gigantic size six or seven inches 
across the wings and are amongst the most remarkably coloured 
of existing Insects. The great Charagia of Australia, with 
colours of green and rose, bearing white spots, are remarkable. 
The South African Leto venus is of large size, and has an 
astonishing supply of glittering metallic splashes on the wings, 
making a barbaric but effective display. The South Australian 
Zelotypia staceyi, of enormous size, is also a handsome ninth: 
but the majority of species of the family are adorned only in 
the feeblest manner. 

Very little is known as to the larvae ; they are either sub- 
terranean, feeding on roots, or they live in the wood of trees and 
shrubs. They are nearly bare, and are apparently the lowest type 
of Bombycid larva. At the same time, it would appear there is 
considerable variety amongst them. Packard says 2 the young 
larva of Jfi-ju'/tfi/s mustelines has the arrangement of setae that is 
normal in Tineidae. The larva of H. hurnali seems to be a very 
simple form, but H. liectus shows a considerable amount of 
divergence from it. They probably live for several years; the 
larva of //. (irgenteo-iii<-nl<it us in North America lives for three 
years, ;it first eating the roots of Alder and then entering the 

1 Ent. Ti-dsTcr. xvi. isnr,. ),. n r>. 

- On lurvnr (.]' Hrjiialidau, J. JW: York c nf. Soc. iii. 1895, p. 69, Plates III. IV. 



VI 



HETEROCERA HEPIALIDAE 



397 



stems. The pupae are also peculiar. They are of unusually 
elongate, cylindrical form, with comparatively feeble integument, 
Luc with a considerable development of chitinous, elevated, toothed 
ridges, on the dorsal aspect, and a very strong ridge of this kind 
on the ventral surface of 
the seventh segment ; the 
wing-sheaths are short; it 
is very difficult to distinguish 
the full number of abdominal 
segments. These pupae are 
remarkably agile, and by 
wriggling and kicking are 
able to move a considerable 
distance ; it is said that they 
can force themselves to the 
surface even when the super- 
ficial soil is quite hard. 
We cannot consider this 
pupa naturally placed 
either the pupae 




amongst 




x-O 




Britain. 



obtectae or incompletae of FIG- 198. ?\\y&ofHepiaiuship'uiini<s. 

A, Ventral ; B, dorsal aspect. 

Chapman. 

We have already remarked that little is known as to the 
life-histories. The species are probably prolific, a female of 
H. thule having been known to deposit more than 2000 eggs. 
Of the Australian forms little more is known : than that they live 
in the wood of trees and shrubs, and are rapidly disappearing ; 
we may fear that some are extinct without ever having been 
discovered, and others, also unknown but still existing, may 
disappear only too soon ; the wasteful destruction of timber in 
Australia having been deplorable. 

The peculiar habits of the Hepialidae are not likely to bring 
the Insects to the net of the ordinary collector, and we believe 
they never fly to light, hence it is probable that we are 
acquainted with only a small portion of the existing species ; 
their distribution is very wide, but Australia seems to be their 
metropolis, and in Xew Zealand twelve species are known. The 
genera as at present accepted are remarkable for their wide 
distribution. Leto is said to occur in South Africa and in the 

1 Olliff, Australian Hepialidae, Ento-i>K>!o<jist, xxviii. 1895, p. 11-L 



398 LEPIDOPTERA CHAP. 

Fiji Islands ; but we must repeat that the study of these 
interesting Insects is in a very primitive state, and our present 
knowledge of their distribution may be somewhat misleading. 

The habits of the European Hepialus in courtship have been 
observed to a considerable extent and are of great interest, an 
astonishing variety and a profound distinction in the methods 
by which the sexes are brought together having been revealed. 

H. humuli, our Ghost-moth, is the most peculiar. Its 
habits were detected by Dr. Chapman. 1 The male is an 
Insect of exceptional colour, being white above, in consequence of 
a dense formation of imperfect scales ; the female is of the 
brownish tints usual in Swift -moths. In the month of June 
the male selects a spot where he is conspicuous, and hovers 
persistently there for a period of about twenty minutes in the 
twilight ; his colour has a silvery- white, glistening appearance, 
so that the Insect is really conspicuous notwithstanding the 
advanced hour. Females may be detected hovering in a some- 
what similar manner, but are not conspicuous like the male, 
their colour being obscure; while so hovering they are oviposit- 
ing, dropping the eggs amongst the grass. Females that have 
not been fertilised move very differently and dash about in 
an erratic manner till they see a male ; they apparently have 
no better means of informing the hovering male of their presence 
than by buzzing near, or colliding with him. Immediately this 
is done, the male abandons his hovering, and coupling occurs. 
There can be little doubt that the colour of the male attracts 
the female; but there is a variety, hethlandica, of the former 
sex coloured much like the female, and in some localities 
varieties of this sort are very prevalent, though in others the 
species is quite constant. This variation in the colour of the 
males is very great in Shetland, 2 some being quite like the 
females. In H. kcctus the two sexes are inconspicuously and 
similarly coloured. The male hovers in the afternoon or evening 
in a protected spot, and while doing so diffuses an agreeable 
odour said by Barrett to be like pine-apple and this brings 
the female to him, much in the same manner as the colour 
of If. li a in nil brings its female. The hind legs of the male 



1 Ent. Minj. xiii. 1876, p. 63 ; ami xxiii. 1886, p. 164. 

' Weir, J'Jii/ini/ii/iii/is/, xiii. 1S80, p. 219, plate ; King, Ent. llcconl, vii. 1895, 
p. 111. 



vr IIETEROCERA- HEPIALIDAE 399 

are swollen, being filled with glands for secreting the 
odorous matter. 1 This structure has led to the suggestion of 
the generic name Phymatopus for the Insect. Turning to 
other species of the genus, we find that the normal rela- 
tive roles of the sexes are exhibited,, but with considerable 
diversity in the species. In H. lupulinus the males fly about 
with rapidity, while the female sits on a stem and vibrates 
her wings ; she thus attracts the males, but they do not 
perceive her unless happening to come within three or four 
feet, when they become aware of her proximity, search for and 
find her. It is doubtful whether the attraction is in this case 
the result of an odour ; it would appear more probable that it 
may be sound, or that the vibration of the wings may be felt by 
the male. 

In H. sylmnus, H. velleda and H. pyrenaicus less abnormal 
modes of attracting the males occur, the individuals of this latter 
sex assembling in great numbers at a spot where there is a female. 
In the first of the three species mentioned the female sits in the 
twilight on the stem of some plant and vibrates the wings with 
rapidity ; she does not fly ; indeed, according to Mr. Eobson, she. 
does not till after fertilisation move from the spot where she 
emerged. In H. pyrenaicus the female is quite apterous, but is 
very attractive to the males, which as we have said, assemble 
in large numbers near her. Thus within the limits of these 
few allied forms we find radically different relations of the 
sexes. 

1. The male attracts the female (A) by sight (H. humuli) ; 

(B) by odour (H. liectus}. 

2. The female attracts the male (A) by vibration of wings 

(H. lupulinus and If. sylvinus) ; (B) without vibration, 
but by some means acting at a distance (H. velleda, 
If. pyrenaicus}. 

Little or nothing is known as to the habits of the great 
majority of the more remarkable forms of the family. The 
gigantic Australian forms are believed to be scarcely ever seen on 
the wing. 

The Hepialidae differ from other Lepidoptera by very im- 
portant anatomical characters. The absence of most of the 

1 Bertkau, S. Vcr. Rheinland, xxxvi. 1879, p. 288 ; and Arch. Natury. 
xlviii. i. 1882, p. 362. 



400 LEPIDOPTERA CHAP. 






mouth-parts is a character common to them and several other 
divisions of Lepidoptera ; but the labial palpi are peculiarly 
formed in this family, being short and the greater portion of 
their length consisting of an undivided base, which probably 
represents some part of the labiuin that is membranous in 
normal Lepidoptera. The thoracic segments are remarkably 
simple, the three differing less from one another than usual, and 
both meso- and nieta-notum being much less infolded aud co- 
ordinated. The wings are remarkable for the similarity .of the 
ncrvuration of the front and hind wings, and by the cell being- 
divided by longitudinal nervules so as to form three or four 
cells. On the inner margin of the front wing there is near 
the base an incision marking off a small prominent lobe, the 
jugmii of Prof. Comstock. Brandt mentions the following 
anatomical peculiarities, 1 viz. the anterior part of the alimentary 
canal is comparatively simple ; the respiratory system is in some 
points like that of the larva ; the heart is composed of eight 
chambers ; the appendicular glands of the female genitalia are 
wanting. The testes remain separate organs throughout life. 
The chain of nerve ganglia consists of the supra- and iufra- 
oesophageal, three thoracic, and five abdominal, ganglia, while 
other Lepidoptera have four abdominal. 

Fam. 24. Callidulidae. -A small family of light - bodied 
diurnal moths having a great resemblance to butterflies. In 
some the frenulum is present in a very rudimentary condition, 
and in others it is apparently absent. Cleosiris and J'f<rodecta are 
very like butterflies of the Lycaenid genus Thccln. Although 
fifty species and seven or eight genera are known, we are quite 
ignorant of the metamorphoses. Most of the species are found 
in the islands of the Malay Archipelago, but there are a few in 
East India. 

Fam. 25. Drepanidae (or Drepanulidae). (Hook-tips}. The 
larger moths of this family are (A moderate size ; many of 
the species have the apex of the front wing pointed or even 
hooked ; some have very much the appearance of Geometrid 
moths; they resemble very different members of that family. 
(h-i'hi. liyalodisca is remarkable on account of the very large, 
transparent patch on each front wing, though the other species 
of the genus have nothing of the sort. In the genus Deroca we 

1 Zoo!. An;, iii. is.SD, p. 18(5. 



VI 



HETEROCERA DREPANIDAE LIMACODIDAE 



4OI 



find Insects with the scales imperfect, they being few and small 
and approximating in form to hairs ; in D. hyalina scales are 
nearly entirely absent. In other genera, e.g. Peridrepana, Stre^>- 
toperas, there is only a very inferior state of scale -formation. 
The few larvae that are known are peculiar ; they are nearly 
bare of hair, without the pair of terminal claspers, while the 
body is terminated by a long tubular process. They form a 
slight cocoon among leaves. 

The members of the family were formerly much misunder- 
stood, and were assigned to various positions in the Order. 
There are now about 30 genera, and 150 species known, the 
geographical distribution of the family being very wide. In 
Britain we have half a dozen species. Cilix glaucata (better 
known as G. spinula) is said " to undoubtedly imitate " the 
excrement of birds. No doubt the Insect resembles that sub- 
stance so as to be readily mistaken for it. This Insect has a 
very wide distribution in North America, Europe and East 
India, and is said to vary so much in the structure of its organs 
as to justify us in saying that the one species belongs to two or 
three genera. 

Fam. 26. Limacodidae (or Eucleidae). These are some- 
what small moths, of stout formation, sometimes very short in 
the body, and with rather small wing - area. The family 
includes however at present many Insects of diverse appearance ; 
there are numerous forms in which 
apple-green is a prominent colour : 
some bear a certain resemblance 
to the Swifts, others to Noctuids ; 
som.e,Hosema and Staetkerinia,are of 
extraordinary shapes ; certain very 
small forms, Gavara, Ceratonema, 
resemble Tortricids or Tineids ; a 
few even remind one of Insects of FlG - 199. Mature larva of Apoda 

testudo, on beech-leaf. Britain. 

other Orders ; so that the group is 

a mimetic one. Nagoda nigricans (Ceylon) has the male some- 
what like a Psychid, while the female has a different system of 
coloration and wing-form. In Scopelodes the palpi are in both 
sexes remarkable ; elongated, stiff, directed upwards and brush- 
like at the tip. Altogether there are about 100 genera and 
400 species known ; the distribution of the family is very wide 
VOL. vi 2 D 




402 



LEPIDOPTERA 



CHAP. 



iu both hemispheres, but these Insects do not occur in insular 
faunas. In Britain we have two genera, Hcterogenea and Apoda 
(better known as Limacodes x ), each with a single species. 

The early stages of these Insects are of great interest. The 
eggs, so far as known, are peculiar fiat oval scales, of irregular 
outline and transparent ; we have figured an example in Vol. V. 
Fig. 83. The eggs of the same moth are said to vary much 
in size, though the larvae that emerge from them differ little 

from one another in this respect. The 
latter are peculiar, inasmuch as they 
have no abdominal feet, and the 
thoracic legs are but small ; hence the 
caterpillars move in an imperceptible 
gliding manner that has suggested for 
some of them the name of slug-worms. 
The metamorphoses of a few are 
known. They may be 




A V- 



arranged in 

two groups ; one in which the larva 
is spinose or armed with a series of 
projections and appendages persisting 
throughout life ; while in the members 
of the second group the spines have 
only a temporary existence. At the 
moment the young larva of Apoda 
testudo emerges from the egg it 

FIG. 200. Larva of Ajx>d<( tcxtndo ] lag nQ CO nspicuOUS SpillCS 01' processes, 
just hatched. A, Dorsal view 

of larva ; B, C, D, a spine in and is an extremely soft, colourless 
different states of evagination creature 2 but it a l most immediately 

All magnified. (After Chapman.) 

displays a remarkable system of com- 
plex spines. These really exist in the larva when it is 
hatched, and are thrust out from pits, as explained by 
Dr. Chapman. In the succeeding stages, the spines become 
modified in form, and the colour of the body and the nature of 

1 It is much to be, regretted that, as in so many other Lepidoptera, no satis- 
factory agreement as to names has been attained'; -our British A. tcstudo is 
variously styled Limacodes testudo (by Chapman and most naturalists), Apoda 
liiiiKi-oifcn' C\<y Meyrick), or Apoda ai'cllana (Kirby, Catalogue of Motlis}. The 
family is called either Limacodidae, Apodidae, Cochliopodidae, or Heterogeneidae. 

2 See Chapman, Tr. ait. Soc. London, 1894, p. 345, Plate VII., for our British 
species ; for North American forms, Dyar, Life-histories of the New York Sluy-cater- 
pillars (in progress, with numerous plates), J. Neiv York cut. Soc. iii. etc., 1895. 



vi HETEROCERA LIMACODIDAE 403 

the integument are much changed, so that in the adult 
larva (Fig. 199) the spines have subsided into the condition 
of mere prominences, different in colour from the rest of 
the surface. These larvae appear to be destitute of a head, 
hut there really exists a large one which is retracted, except 
during feeding, into the body ; the five pairs of abdominal feet of 
the larvae of allied families are replaced by sucker-like structures 
on the first eight abdominal segments. The spinneret of the 
mouth is not a pointed tubular organ, but is fish-tailed in shape, 
and hence disposes the silky matter, that aids the larva in mov- 
ing on the leaves, in the form of a ribbon instead of that of a 
thread. It has been stated that these peculiar larvae " imitate " 
the coloured galls frequently found on the leaves of trees. The 
North American forms of this family have very varied and most 
extraordinary larvae. 1 In the pretty and conspicuous larva of 
Empretia stimulea, the tubercles or processes of the body are, in 
the later stages, armed with hairs, that contain a poisonous or 
irritating fluid, said to be secreted by glands at the bases of the 
processes. These hairs are readily detached and enter the skin 
of persons handling the caterpillars. The larva of the North 
American Hag-moth, Phobetron pithecium, is a curious object, 
bearing long, fleshy appendages covered with down. Hubbard 
makes the following statement as to the instincts of this larva : 2 - 
' The hag-moth larvae do not seek to hide away their cocoons, 
but attach them to leaves and twigs fully exposed to view, with, 
however, such artful management as to surroundings and har- 
monising colours that they are of all the group the most difficult 
to discover. A device to which this Insect frequently resorts 
exhibits the extreme of instinctive sagacity. If the caterpillar 
cannot find at hand a suitable place in which to weave its 
cocoon, it frequently makes for itself more satisfactory surround- 
ings by killing the leaves, upon which, after they have become 
dry and brown in colour, it places its cocoon. Several of these 
caterpillars unite together, and selecting a long and vigorous 
immature shoot or leader of the orange tree, they kill it by 
cutting into its base until it wilts and bends over. The 
leaves of a voung shoot in drying turn a light tan-color, which 

*/ O v O O 



1 See Packard, P. Amcr. Phil. Sue. xxxi. 1893, pp. 83, 108, Plates. (He uses 
the term Cochliopodidae instead of Limacodidae) ; also Dyar, as above. 
- Insects affectiny the Oranyc, "Washington, 1885, p. 143. 



404 LEPIDOPTERA CHAP. 

harmonises most perfectly with the hairy locks of the caterpillar 
covering the cocoon. The latter is, consequently, not easily 
detected, even when placed upon the exposed and upturned 
surface of the leaf." 

The cocoons of Limacodidae are unusually elaborate, the 
larva forming a perfect lid in order to permit itself to escape 
when a moth. Chapman states that the larva lies unchanged 
in the cocoon, all winter, moulting to a pupa in the spring, and 
that the pupa escapes from the cocoon previous to the emergence 
of the moth. 1 Both Chapman and Packard look on the family 
as really nearer to Microlepidoptera than to Bombyces ; Meyrick 
(calling it Heterogeneidae) places it at the end of his series 
Psychina next Zygaenidae. 

We may allude here to the little moths, described by West- 
wood under the name of Epipyrops? that have the extraordinary 
habit of living on the bodies of live Homopterous Insects of the 
family Fulgoridae in India. What their nutriment may be is 
not known. The larva exudes a white flocculent matter, which 
becomes a considerable mass, in the midst of which the caterpillar 
changes to a pupa. Westwood placed the Insect in Arctiidae ; 
Sir George Hampson suggests it may be a Limacodid, and this 
appears prol table. 

Fam. 27. Megalopygidae (or Lagoidae). The American 
genera, Megalopyge and Lagoa, are treated by Berg and by 
Packard 3 as a distinct family intermediate between Saturniidae 
and Limacodidae. The larva is said by the latter authority to 
have seven pairs of abdominal feet instead of live pairs the 
usual number in Lepidoptera, When young the caterpillars of 
Lagoa o]>?irnl<iris are white and resemble a Mock of cotton wool. 
When full grown the larva presents the singular appearance of 
a lock of hair, moving in a gliding, slug-like manner. Under 
the long silky hair there are short, stiff, poison - hairs. The 
larva forms a cocoon, fitted with a hinged trap-door for the 
escape of the future moth. This curious larva is destroyed by 
both Dipterous and Hymenopterous parasites. 

Fam. 28. Thyrididae. A small family of Pyraloid moths, 
exhibiting considerable variety of form and colour, frequently 
with hyaline patches on the wings. They are mostly small 

1 Tr. cut. Sue. Lv,ulii. 1894, p. 348. ' 2 Op. cif. 1876. p. 522 ; and 1S77, p. 433. 
3 P. Amer. Phil. Soc. xxxii. 1894, p. 27u. 



vi HETEROCERA THYRIDIDAE LASIOCAMPIDAE 405 

Insects, and contain no very striking forms. Some of them 
look like Geometrids of various groups. The family is widely 
distributed in the tropical zone, and includes 25 genera, of 
which lihodoneura, with upwards of 100 species, is the chief 
one. The larvae are said to be similar to those of Pyralidae. 
This family is considered by Hampson and Meyrick to be 
ancestral to butterflies. 1 

Fam. 29. Lasiocampidae (Eyycrs, Lappet -moths'). Usually 
large Insects densely covered with scales, without frenulum, 
but with the costal area of the hind wing largely developed, 
and the male antennae beautifully pectinate, Lasiocampids are 
easily recognised. They are well known in Britain, though we 
have but few species. The flight of some of the species is 
powerful, but ill-directed, and the males especially, dash about 
as if their flight were quite 
undirected ; as indeed it 
probably is. The differ- 
ence in the flight of the 
two sexes is great in some 
species. In the genus 
Suana and its allies we meet 
with moths in which the 
difference in size of the FIG. 201. Lappet-moth, Gastropacha querci- 

two sexes is extreme ; the llia > ? ' BritahK 

males may be but 1^- inches across the wings, while the very 
heavy females may have three times as great an expanse. Kirby 
separates these Insects to form the family Pinaridae ; it in- 
cludes the Madagascar silkworm, Boroccra madagascariensis. 
The African genus Hilbrides is remarkable for the wings being 
destitute of scales, and consequently transparent, and for being of 
very slender form like a butterfly. The eggs of Lasiocampidae 
are smooth, in certain cases spotted in an irregular manner like 
birds' eggs. Sometimes the parent covers them with hair. 
The larvae are clothed with a soft, woolly hair, as well as with a 
shorter and stiffer kind, neither beautifully arranged nor highly 
coloured, and thus differing from the . caterpillars of Lyman- 
triidae ; this hair in some cases has very irritating pro- 
perties. Cocoons of a close and compact nature are formed, and 
hairs from the body are frequently mixed with the cocoon. In 
1 Revision of the Thyrididae ; Hampson, P. Zool. Soc. London, 1897, p. 603. 




406 LEPIDOPTERA 



CHAT. 



some species the walls of the cocoons have a firm appear- 
ance, looking very like egg-shell a fact which is supposed to 
have given rise to the name of Eggers. Professors Poulton 
and Meldola have informed us that this appearance is produced 
by spreading calcium oxalate on a slight framework of silk, 
the substance in question being a product of the Malpighian 
tubes. 1 In various families of Lepidoptera it happens that 
occasionally the pupa exists longer than usual before the appear- 
ance of the perfect Insect, and in certain members of this family 
notoriously in Poecilocampa populi, the December moth this 
interval may lie prolonged for several years. There is not at 
present any explanation of this fact. It may lie of interest to 
mention the following case : From a batch of about 100 eggs 
deposited by one moth, in the year 1891 (the Puss-Moth of the 
family Notodontidae), some sixty or seventy cocoons were obtained, 
the feeding up of all the larvae having been effected within 
fourteen days of one another ; fourteen of the Insects emerged 
as moths in 1892 ; about the same number in 1893 ; in 1894, 
twenty-five; and in 1895, eleven emerged. Lasiocampidae is a 
large family, consisting of some 100 genera and 500 or more 
species, and is widely distributed. It is unfortunately styled 
Bombycidae by some naturalists. 

Fam. 30. Endromidae. - -The "Kentish glory," Enclromis 
versicolo)-, forms this family ; it is a large and strong moth, and 
flies wildlv in the daytime in birch-woods. The larva has but 

U I/ 

few hairs, and is said when young to assume a peculiar position, 
similar to that of saw-fly larvae, by bending the head and thorax 
backwards over the rest of the body. 

Fam. 31. Pterothysanidae. Consists of the curious East 
Indian genus Pterothysanus, in which the inner margins of the 
hind wings are fringed with long hairs. They are moths of 
slender build, with large wing-expanse, black and white in colour, 
like Geometrids. There is no frenulum. Metamorphoses un- 
known. 

Fam. 32. Lymantriidae. (Better known as Liparidae\ 
These are mostly small or moderate-sized moths, without brilliant 
colours ; white, black, grey and brown being predominant : with 
highly-developed, pectinated antennae in the male. The larva 
is very hairy, and usually bears tufts or brushes of shorter hairs, 

1 P. cnt. Soc. London, 1891, p. xv. 



vi HETEROCERA LYMANTRIIDAE 407 

together with others much longer and softer, these being some- 
times also amalgamated to form pencils ; the coloration of these 
larvae is in many cases very conspicuous, the tufts and pencils 
being of vivid and strongly contrasted colours. Some of these 
hairy larvae are poisonous. A cocoon, in which much hair is 
mixed, is formed. The pupae are remarkable, inasmuch as they 
too are frequently hairy, a very unusual condition in Lepidoptera. 
The Lyrnantriidae is one of the largest families of the old group 
Bombyces; it includes some 180 genera and 800 species, and is 
largely represented in Australia. Dasyehira rossii is found in the 
Arctic regions. In Britain we have eight genera represented by 
eleven species ; the Gold-tails, Brown-tails and Vapourer-moths 
being our commonest Bombyces, and the latter being specially fond 
of the London squares and gardens, where its beautiful larva may 
be observed on the leaves of roses. Most of the Lymantriidae are 
nocturnal, but the male Vapourer-moth flies in the daytime. In 
this family there are various species whose females have the 
wings small and unfit for flight, the Insects being very sluggish, 
and their bodies very heavy. This is the state of the female of 
the Vapourer-moth. The males in these cases are generally re- 
markably active, and very rapid on the wing. 

Some of these moths increase in numbers to an enormous 
extent, and commit great ravages. Psilura monacha the Nun, 
" die Nonne " of the Germans, 1 is one of the principal troubles of 
the conservators of forests in Germany, and great sums of money 
are expended in combating it ; all sorts of means for repressing 
it, including its infection by fungi, have been tried in vain. The 
caterpillars are, however, very subject to a fungoid disease, com- 
municated by natural means. It is believed, too, that its con- 
tinuance in any locality is checked after a time by a change 
in the ratio of the two sexes. It is not a prolific moth, for it 
lays only about 100 eggs, but it has been shown that after 
making allowance for the numerous individuals destroyed by 
various enemies, the produce of one moth amounts in five genera- 
tions to between four and five million individuals. The larva 
feeds on Coniferae, and on many leafy trees and shrubs. The young 

1 This moth is known under several generic names Psilura, Liparis, Ocneria, 
Lymantria ; there is now a very extensive literature connected with it. A good 
general account by A\ 7 achtl may be found in Jf'ien. ent. Zeit. x. 1891, pp. 149-180, 
2 Plates. 



408 LEPIDOPTERA 



CHAP. 



larva is provided with two sets of setae, one set consisting of very 
long hairs, the other of setae radiating from warts ; each one of this 
second set of spines has a small bladder in the middle, and it has 
been suggested that these assist in the dissemination of the young 
caterpillars by atmospheric means. 1 These aerostatic setae exist 
only in the young larva. The markings of the moth are very 
variable ; melanism is very common both in the larva and imago ; 
it has been shown conclusively that these variations are not 
connected, as black larvae do not give a larger proportion of black 
moths than light-coloured caterpillars do. In England this 
moth is never injurious. A closely allied form, Ocneria dispar, 
was introduced by an accident into North America from Europe 
about thirty years ago ; for twenty years after its introduction it 
did no harm, and attracted but little attention ; it has, however, 
now increased so much in certain districts that large sums of 
money have been expended in attempting its extirpation. 

DasycMra pudilmnda has occasionally increased locally to an 
enormous extent, but in the limited forests of Alsace the evil was 
cured by the fact that the caterpillars, having eaten up all the 
foliage, then died of starvation. 2 Team melanosticta is said to 
produce columns of processionary caterpillars in Australia. 

Fam. 33. Hypsidae (or Aganaidae). -A family of compara- 
tively small extent, confined to the tropical and sub-tropical 
regions of the Eastern hemisphere. The colours are frequently 
huff and grey, with white streaks on the outer parts of the wings. 
We have nothing very like them in the European fauna, our 
species of Spilosoma are perhaps the nearest approach. In 
Euplocia the male has a pouch that can be unfolded in front of 
the costa at the base of the anterior wing; it is filled with very 
long, peculiar, hair-like scales growing from the costal margin ;. 
both sexes have on each side of the second abdominal segment 
a small, projecting structure that may be a sense-organ. The 
female is more gaily coloured than the male. 

Fam. 34. Arctiidae. With the addition recently made to 
it of the formerly separate family Lithosiidae, Arctiidae has 
become the most extensive family of the old liombycid series of 
moths, comprising something like 500 genera and ,'5000 species. 
Hampson recognises four sub-families Arctiinae, Lithosiinae, 

1 WacLtl and Kornauth, Mitt, for si. Fer.s //<// .sv/v.sc/t Osterreichs, Heft xvi. 1893. 
'-' Crahay, Ann. Hoc. cut. L'clyiijttc, xxxvii. 1893, p. 282. 



vi HETEROCERA -ARCTIIDAE 409 

Nolinae, Nycteolinae, to which may be added others from 
America Pericopiiiae, Dioptiuae, Ctenuchinae; these sub-families 
being treated as families by various authors. The sub-family 
Arctiinae includes our Tiger- and Ermine -moths, and a great 
many exotic forms of very diverse colours and patterns ; the 
species of this division are, on the whole, probably more variable 
in colour and markings than in any other group of Lepidoptera. 
There are many cases of great difference of the sexes ; in the 
South American genus Ambryllis the male is remarkable for its 
hyaline wings with a few spots ; while the female is densely 
scaled, and very variegate in colour. There are some cases (the 
South European genus Ocnogytui) where the female is wingless 
and moves but little, while the male flies with great rapidity. 
Epicansis smithi, from Madagascar, one of the most remarkable 
of moths, is placed in this division of Arctiidae ; it is of a tawny 
colour, variegate with black ; the abdomen of this latter colour 
is terminated by a large tuft of long scarlet hairs; the Insect 
has somewhat the appearance of a Hummingbird -hawkmoth. 
Ecpantheria is an extensive genus of tropical American moths 
(having one or two species in North America), of black and 
white or grey colours, with very complex markings ; the male in 
some species has a part of the hind wing produced as a tail, or 
lobe, of a different colour. 

The sub- family Pericopiiiae are almost peculiar to South 
America (two species of Gnophaela exist in North America) ; 
some of this sub-family bear a great resemblance to Heliconiid 
butterflies. 

The Dioptinae are likewise American moths of diurnal habits, 
and many of them bear a striking resemblance to the Ithomiid 
butterflies they associate with when alive. 

The sub -family Lithosiinae is of great extent ; our native 
' Footmen " give a very good idea of it : the moths are generally 
of light structure, with long, narrow front wings ; a simple system 
of yellow and black colour is of frequent occurrence. Many of 
this group feed in the larval state on lichens. Hainpson includes 
in this group the Nyctemeridae light-bodied diurnal moths, 
almost exclusively of black and white colours, of Geometrid 
form, frequently treated as a distinct family. 

The sub-family Nolinae is a small group of rather insignificant 
Insects, in appearance like Pyralids or Geometrids ; four or five 



4IO LEPIDOPTERA CHAP. 

species are native in Britain. Packard maintains the family 
Nolidae as distinct. 1 

The sub-family Xycteolinae consists of a few small moths the 
position of which has always been uncertain; Nycteola (better 
known as Sarrothripus), If alias, and Earias are all British genera 
that have been placed amongst Tortrices, to which they bear a 
considerable resemblance. Sarrothripus is at present placed by 
Hampsoii in Xoctuidae, by others in Lithosiidae, by Meyrick in 
Arctiidae. The sub-family forms the family Cymbidae of Kirby; 2 
it includes at present only about 70 species, all belonging to the 
Eastern hemisphere. Two types of larvae are known in it : one 
bare, living exposed on leaves ; the other, Earias, hairy, living 
among rolled-up leaves. Halias prasinana is known from the 
testimony of numerous auditors to produce a sound when on the 
wing, but the modus operandi has not been satisfactorily ascer- 
tained. Sound-production seems to be of more frequent occurrence 
in Arctiidae than it is in any other family of Lepidoptera : 
Dionychopus nirr-ns produces a sound by, it is believed, friction 
of the wings. In the case of the genera Setina and Chrlonia 
the process is said to be peculiar to the male sex : Laboulbene 
believes it to proceed from drum-like vesicles situate one on each 
side of the base of the metathorax. 3 

Fam. 35. Agaristidae. An interesting assemblage of moths, 
many of them diurnal and of vivid colours, others crepuscular. 
There is considerable variety of appearance in the family, although 
it is but a small one, and many of its members remind one of 
other and widely separated families of Lepidoptera. The style 
and colour of the Japanese Euscmia villieoides are remarkably 
like our Arctia villica. In some forms the antennae are some- 
what thickened towards the tip and hooked, like those of the 
Skipper butterflies. The family consists at present of about 250 
species, but we doubt its being a sufficiently natural one. It is 
very widely distributed, with the exception that it is quite absent 
from Europe and the neighbourhood of the Mediterranean Sea. 
fn North America it is well represented. The larvae, so far as 
known, are not very remarkable; they have some lateral tufts of 
hair, as well as longer hairs scattered over the body. 

1 Amcr. Natural, xxix. 1895, p. 801. 

- Catalogue of Lepidopterct Heterocera, i. 1892. 

3 Ann. &>c. cut. France (4), iv. 1864, p. 689. 



vi HETEROCERA AGARISTIDAE GEOMETRIDAE 4! I 

The male of the Indian Acyocera tripartita has been noticed 
to produce a clicking sound when flying, and Sir G. Hampson has 
shown : that there is a peculiar structure on the anterior wing ; 
he considers that this is rubbed against some spines on the front 
feet, and that the sound is produced by the friction. Though 
this structure is wanting in the acknowledged Congeners of 
A. fi-ipm-tita, yet it occurs in a very similar form in the genus 
Hccntcsin, already noticed under Castniidae. 

Fam. 36. Geometridae (Carpets, Pugs, efr.)--This very 
extensive family consists of fragile moths, only a small number 
being moderately stout forms ; they have a large wing - area ; 
the antennae are frequently highly developed in the males, 
but on this point there is much diversity. Either the frenulum 
or the proboscis is absent in a few cases. The caterpillars are 
elongate and slender, with only one pair of abdominal feet- 
placed on the ninth segment in addition to the anal pair, or 
claspers. They progress by moving these two pairs of feet up to 
the thoracic legs, so that the body is thrown into a large loop, and 
they are hence called Loopers or Geometers. The family is uni- 
versally distributed, and occurs even in remote islands and high 
latitudes ; in Britain we have about 270 species. The family was 
formerly considered to be closely connected with Noctuidae, but at 
present the opinion that it has more intimate relations with the 
families we have previously considered is prevalent. Packard 
considers it near to Lithosiidae, while Meyrick merely places the 
six families, of which he treats it as composed, in his series Noto- 
dontina. Hampson adopts Meyrick's six families as sub-families, 
but gives them different names, being in this respect more con- 
servative than Meyrick, whose recent revision of the European 
forms resulted in drastic changes in nomenclature.' 2 This 
classification is based almost exclusively on wing-nervuration. 
The number of larval legs and the consequent mode of walking 
is one of the most constant characters of the group ; the few 
exceptions that have been detected are therefore of interest. 
Anisopteryx aescularia has a pair of undeveloped feet on the eighth 
segment, and, according to Meyrick, its allies " sometimes show 
rudiments of the other two pairs." The larva of Him era 

1 P. Zool. Soc. London, 1892, p. 188. 

1 Tr. cnt. Soc. London, 1892, pp. 53-140 ; for critic-ism on the nomenclature, 
see Rebel, Ent. Zeit. Stettin, Hii. 1892, p. 247. 



4 I2 



LEPIDOPTERA 



CHAP. 



/ii'/itiaria is said to have in early life a pair of imperfect feet on 
the eighth segment, which disappear as the larva approaches 
maturity. 

The position of the abdominal feet and claspers throws the 
holding power of the larva to the posterior part of the body, 
instead of to the middle, as in other caterpillars. This, com- 
bined with the elongate form, causes these larvae when reposing 
to assume attitudes more or less different from those of other 
larvae ; holding on by the claspers, some of these Insects allow all 
the anterior parts of the body to project in a twig-like manner. 
The front parts are not, however, really free in such cases, but 
are supported by a thread of silk extending from the mouth 
to some point near-by. Another plan adopted is to prop the 

front part of the body 
against a twig placed 
at right angles to the 
supporting leaf, so that 
the caterpillar is in a 
diagonal line between 
the two (Fig. 202). 
Other Geometers assume 
peculiar coiled or spiral 
attitudes during a whole 
or a portion of their 
lives ; some doing this 
on a supporting object 
leaf or twig while 
others hang down 
(Epliyra pendularici}. 
Certain of the larvae of 
Geometridae vary in 
colour, from shades of 
brown to green; there 
is much diversity in 
this variation. In some 




FIG. 202. Larva of 

oil a rose-twig. 



reposin 



x 1. Cambridge. 



species it is simple variation ; in others it is dimorphism, 
i.e. the larvae are either brown or green. In other cases the 
larvae are at first variable, subsequently dimorphic. In AmpJn- 
dasi* I, I ul ,i ri<i it would appear that when the larva is hatched 
the dimorphism is potential, and that the future colour, whether 



vi GEOMETRIDAE 413 

green or brown, is settled by some determining condition during 
the first period of larval life and cannot be subsequently modified. 1 
According to Poulton, the dark tint is due in A. betularia to 

o 

colouring matter in the skin or immediately below it, and the 
green tint to a layer of fat between the hypodermis and the 
superficial muscles ; this layer being always green, but more 
brightly green in the larvae that are of this colour, externally. 
Much discussion has occurred about these larval attitudes and 
colours, and it seems probable that Professor Poulton has over- 
rated the value of protection from birds, mammals and ento- 
mologists ; the chief destroying agents being other than these, 
and not liable to be thus deceived, even if the vertebrates are. 
In some cases such resemblance as undoubtedly exists is not 
made the best use of. The larva shown in figure 202 bore 
a wonderful resemblance, when examined, to the rose-twigs it- 
lived on, but the effect of this as a concealing agent was entirely 
destroyed by the attitude ; for this, being on different lines to 
those of the plant, attracted the eye at once. This larva, and we 
may add numerous other larvae, could have been perfectly con- 
cealed by adopting a different attitude, but never did so ; the 
position represented being constantly maintained except while 
feeding. 

In some species of this family the adult females are without 
wings, or have them so small that they can be of no use for 
flight. This curious condition occurs in various and widely- 
separated groups of the Geometridae ; and it would be naturally 
supposed to have a great effect on the economy of the species 
exhibiting it, but this is not the case. Some of the flightless 
females affect the highest trees and, it is believed, ascend to their 
very summits to oviposit. It has been suggested that they are 
carried up by the winged males, but this is probably only an 
exceptional occurrence ; while, as they are known to lie capable 
of ascending with rapidity by means of crawling and running, 
it may be taken for granted that this is the usual method with 
them. Some of these wingless females have been found in 
numbers on gas-lamps, and are believed to have been attracted 
by the light, as is the case with very many of the winged forms. 2 

1 See Poulton, Tr. cnt. Soc. London, 1SS4, p. 51 ; op. cit - 1892 > P- 293 5 aUl1 
Bateson, p. 213 ; Gould, p. 215. 

2 Girawl, Ann. Soc. ent. France (4), v. 1865, p. 105 ; Fauvel, I.e. Bull. p. liii. 



414 LEPIDOPTERA CHAP. 

Neither is the geographical distribution limited by this inferior 
condition of the most important of the organs of locomotion, for 
CheimatoMa brumata (the Winter-moth) one of the species with 
flightless female, is a common and widely distributed Insect in 
Europe and North America. 

Although the classification of this family is based almost 
entirely on wing-nervuration, yet there are some divisions of the 
Geometridae in which this character is remarkably variable, 
certain individuals frequently exhibiting considerable abnor- 
mality. 1 Amplildasis ltnlur'u< is believed to have changed its 
variation considerably in the course of the last fifty years. Pre- 
vious to that time a black variety of the species was unknown, 
but it has now become common ; and it is believed that other 
species of Geometridae are in process of exhibiting a similar 
phenomenon. 2 

Fam. 37. Noctuidae (Owlet -Moths, Eulen of the Germans). 
-This very extensive assemblage consists of moths rarely seen 
in the day-time, of generally sombre colours, with antennae desti- 
tute of remarkable developments in the male (except in a small 
number of forms) ; proboscis and frenulum both present ; a com- 
plex sense-organ on each side of the body at the junction of the 
metathorax and abdomen. The number of species already known 
can scarcely be less than 8000 ; owing to their large numbers 
and the great general resemblance of the forms, their classifica- 
tion is a matter of considerable difficulty. Although the peculiar 
structure at the base of the thorax was long since pointed put, 
it has never received any thorough investigation. Few other 
remarkable structures have yet been discovered: the most in- 
teresting is perhaps the peculiarity in the hind wings of the 
males of certain Ommatophorinae recently pointed out by Sir 
G. F. Hampson 3 : in the genera Patula and Arijini the form of 
the hind wings is normal in the females, but in the male the 
anterior one-half of each of these wings is aborted, and the 
position of the nervures changed ; this condition is connected 
with the development of a glandular patch or fold on the wing, 
and is remarkable as profoundly affecting a structure which is 

1 For a table, see Meyrick, I.e. 

- Barrett, "Increasing Melanism in British Geometridae," Ent. Monthly May. 
L895, i>. 198. 

a J'. Zwl. tioc. London, 1892, p. 192. 



VI 



NOCTUIDAE 



415 



otherwise so constant that the classification of the family is 
largely based on it. 

The larvae are as a rule destitute of the remarkable adorn- 
ments of hairs and armatures of spines that are so common in 
many of the families we have previously considered ; they are 
fond of concealing themselves during the day and coming out at 
night to feed ; many of them pass most of their time at, or 
beneath, the surface of the ground, finding nourishment in roots 
or the lower parts of the stems of plants ; this is notably the 
case in the genus Agrotis, which is perhaps the most widely 
distributed of all the genera of moths. Such caterpillars are 
known as Cut-worms in North America. 1 The great resemblance, 
inter se, of certain of these Out-worms, much astonished the 
American naturalist Harris, who found that larvae almost per- 
fectly similar produced very different moths. The majority of 
Noctuid larvae have the usual number of legs, viz., three pairs 
of thoracic legs, four pairs of abdominal feet and the terminal 
claspers. In some divisions of the family there is a departure 
from this arrangement, and the abdominal feet are reduced to 
three, or even to two, pairs. One or two larvae are known e.g. 
Eiididiti mi in which the claspers have not the usual function, 
but are free terminal appendages. When the abdominal legs 
are reduced in number (Plusia, e.g.) the larvae are said to be 
Half-loopers, or Semi-loopers, as they assume to some extent the 
peculiar mode of progression of the Geometrid larvae, which are 
known as Loopers. In the case of certain larvae, e.g. Triphaena, 
that have the normal number of feet, it has been observed that 
when first hatched, the one or two anterior pairs of the abdom- 
inal set are ill developed, and the larvae do not use them for 
walking. This is the case 
with the young larva of our 
British Brephos notha (Fig. 
203). Subsequently, how- 
ever, this larva undergoes 
a considerable change, and 
appears in the form shown 
in Fig. 204. This interesting larva joins together two or three 

1 Although this term is widely used in Xorth America, it is not in use in Eng- 
land, though it may possibly have originated in Scotland. See Slingerlaiid, Bull. 
Cornell University Exp. 8tat. 104, 1895, p, 555. 




FIG. 203. Brephos notha. Larva, newly 
hatched. Britain. 



416 



LEPIDOPTERA 



CHAP. 



leaves of aspen and lives between them, an unusual habit for 
Noctuid larvae. When about to pupate it bores into bark or 
soft wood to change to a pupa, Fig. 205 ; 
the specimen represented closed the hole f 
of entry by placing two separate doors 
of silk across the burrow, as shown at d. 
The anal armature of this pupa is ter- 
minated by a curious transverse process. 
The systematic position of this inter- 





FIG. 20-i. Brephos notha. Adult larva. 



FIG. 205. Brephos notha. A, 
Pupa, ventral aspri-t ; B. 
extremity of body, magni- 
fied ; C, the pupa in word ; 
(/, diaphragms constructed 
by the larva. 



esting Insect is very uncertain : Meyrick and others associate it 
with the Geonietridae. 

The larva of Leucania unipunctata is the notorious Army- 
worm that commits great ravages on grass and corn in Xorth 
America. This species sometimes increases in numbers to a con- 
siderable extent without being observed, owing to the retiring- 
habits of the larvae ; when, however, the increase of numbers 
has been so great that food becomes scarce, or for some other 
cause -for the scarcity of food is supposed not to be the only 
reason the larvae become gregarious, and migrate in enormous 
swarms: whence its popular name. The Cotton-worm, Alcti 
xylinae is even more notorious on account of its ravages. Eiley 
states l that in bud years the mischief it commits on the cotton 
crop causes a loss of 0,000,000, and that for a period 
of fourteen successive years the annual loss averaged about 
3,000,000. This caterpillar strips the cotton plants of all but 
their branches. It is assisted in its work by another highly 
destructive Noctuid caterpillar, the Boll -worm, or larva of 
armigera, which bores into the buds and pods. This 

h l!<i>. U. X E/it. Commission, 1885, p. 3. 



vi HETEROCERA NOCTUIDAE 417 

latter Insect attacks a great variety of plants, and has a very 
wide distribution, being found even in England, where happily 
it is always a rare Insect. 

In Britain, as well as in parts of Northern Europe, a Noctuid 
moth, Charaeas graminis, occasionally increases to an enormous 
extent : its larva is called the Hill-grub and lives on the grass of 
pastures, frequently doing great damage in hill-lands. The in- 
crease of this moth seems to take place after the manner of an 
epidemic ; a considerable number of years may pass during which 
it is scarcely seen, and it will then appear in unusual numbers 
in widely separated localities. This moth lays a large number 
of eggs, and is not completely nocturnal in habits ; sometimes it 
may be seen on the wing in great numbers in the hottest sun- 
shine, and it has been noticed that there is then a great dispro- 
portion of the sexes, the females being ten or twenty times as 
numerous as the males. In Australia, the Bugong moth, Agrotis 
spina, occurs in millions in certain localities in Victoria : this 
moth hibernates as an imago, and it formerly formed, in this 
instar, an important article of food with the aborigines. The 
powers of increase of another Xoctuicl moth Erastria scitula 
are of great value. Its habits have been described by 
Eouzaud. 1 On the shores of the Mediterranean the larva of this 
little moth lives on a Scale-Insect Lecanium oleae that infests 
the peach ; and as the moth may have as many as five genera- 
tions in a year, it commits laudable havoc with the pest. The 
larva is of remarkable form, very short and convex, with small 
head, and only two pairs of abdominal feet. The scale of the 
Lecanium is of larger size than is usual in that group of Insects, 
and the young larva of the Erastria buries itself, as soon as 
hatched, in one of the scales ; it destroys successively numerous 
scales, and after having undergone several moults, it finds itself 
provided, for the first time, with a spinneret, when, with the aid 
of its silk, it adds to and adapts a Coccid scale, and thus forms a 
portable habitation ; this it holds on to by means of the pair of 
anal claspers, which are of unusual form. The case is afterwards 
subjected to further alteration, so that it may serve as a protec- 
tion to the creature when it has changed to a pupa. This moth 
is said to be free from the attacks of parasites, and if this be the 
case it is probable that its increase is regulated by the fact that 

1 Insect Life, vi. 1894 p. 6. 
VOL. VI 2 E 



41 8 LEPIDOPTERA 



CHAP. 



when the creature becomes numerous it thus reduces the food 
supply, so that its own numbers are afterwards in consequence 
diminished. 

One of the most remarkable genera of British Noctuidae is 
Acronycta, 1 the larvae of which exhibit so much diversity that it 
has been suggested that the genus should be dismembered and its 
fragments treated as allied to several different divisions of moths. 
There are many points of interest in connection with the natural 
history of these Acronycta. A. psi and A. tridens are practically 
indistinguishable as moths, though the larvae are easily separated : 
the former species is said to be destroyed to an amazing extent 
by parasites, yet it remains a common Insect. The genus 
Apatela is very closely allied to Acronycta, and Harris says that 
' Apatda signifies deceptive, and this name was probably given to 
the genus because the caterpillars appear in the dress of Arctians 
and Liparians, but produce true owlet-moths or Noctuas." The 
species of another British genus, Brybphila, possess the excep- 
tional habit of feeding on lichens. Some of the American group 
Erebides are amongst the largest Insects, measuring seven or 
eight inches across the expanded wings. 

The Deltoid moths are frequently treated as a distinct family, 
Deltoidae, perhaps chiefly because of their res'emblance to Pyra- 
lidae. At present, however, they are considered to be separated 
from Noctuidae by no valid characters. 

Fam. 38 Epicopeiidae. The genus Epicopeia, consists of 
only a few moths, but they are amongst the most extraordinary 
known : at first sight they would be declared without hesitation 
to be large swallow-tail butterflies, and Hampson states that they 
mimic " the Papilios of the Polijxenus group. Very little is 
known about these extremely rare Insects, but the larva is stated, 
on the authority of Mr. Dudgeon, to surpass the moths themselves 
in extravagance ; to be covered with long processes of snow-white 
efflorescence, like wax, exuded from the skin, and to " mimic " a 
colony of the larva of a Homopterous Insect. Some ten 
'species of this genus are known from Java, India, China, and 
Japan. In this family there is said to be a rudimentary frenu- 
lum, but it is doubtful whether the hairs that have given rise to 
this definition really justify it. 

1 Sc-i- < 'li.'ipniaii, The Genus Acronycta and UK Allies, London, 1893. 
2 Insects Injurious, etc., Ed. 18(52, Boston, p. 437. 



VI 



HETEROCERA URANIIDAE 



419 



Fam. 39. Uraniidae. A family of small extent, including 
light-bodied moths with ample wings and thread-like antennae ; 
most of them resemble Geometridae, but a few genera, Urania 
and Nyctalcmon, are like Swallow-tail butterflies and have 
similar habits. The Madagascar moth, Chrysiridia inadagascar- 
iensis (better known as Urania rhipheus], is a most elegant and 
beautiful Insect, whose only close allies (except an East African 
congener) are the tropical American species of Urania, which 
were till recently treated as undoubtedly congeneric with the 
Madagascar moth. The family consists of but six genera and 
some sixty species. The question of its affinities has given rise 
to much discussion, but on the whole it would appear that these 
Insects are least ill-placed near Xoctuidae. 1 The larva of the 
South American genus Coro- 
nidia is in general form like 
a Noctuid larva, and has the 
normal number of legs ; it 
possesses a few peculiar fleshy 
processes on the back. A 
description of the larva of 
Chi 'ysii 'idia madagasc ari ensis 
has been widely spread ; but 
according to Camboue, 2 the 
account of the metamorphoses, 
first given by Boiscluval, is 
erroneous. The larva, it ap- 8 
pears, resembles in general 
form that of Coronidia, and FI G- 206. Abdomen of ci* -,>/.*; ,->,t>,i mada- 

A, Horizontal section show- 




gascanensis, 

ing the lower part of the male abdomen : 
1, first segment ; 2, spiracle of second 
segment ; 4-8, posterior segments. B, 
the abdomen seen from the side, with the 



segments numbered. The section is that 
of an old, dried specimen. 



has sixteen feet ; it is, how- 
ever, armed with long, spatu- 
late black hairs ; it changes to 
u pupa in a cocoon of open 
network. 

In all the species of this family we have examined, we have 
noticed the existence of a highly peculiar structure that seems 
hitherto to have escaped observation. On each side of the 
second abdominal segment there is an ear-like opening (usually 

1 See "\Vestwood, Tr. Zool. Soc. London, x. pp. 507, etc., for discussion of this 
question and for figures ; also E. Renter, Act. Soc. Sci. Fcnn. xxii. 1896, p. 202. 
- Cony r. Internal. Zool. ii. 1892, pt, 2, p. 180. 



420 LEPIDOPTERA CHAP. 

much concealed by overlapping scales), giving entrance to a 
chamber in the body ; this chamber extends to the middle line, 
being separated from its fellow by only a thin partition. At 
its anterior and lateral part there is a second vesicle-like 
chamber, formed by a delicate membrane that extends as far 
forwards as the base of the abdomen. There can be little doubt 
that this is part of some kind of organ of sense, though it is 
much larger than is usual with Insect sense-organs. 

Fam. 40. Epiplemidae. Under this name Hampson has 
assembled certain Geometroid moths, some of them placed previ- 
ously in Chalcosiidae, some in Geometridae. They form a varied 
group, apparently closely allied to Uraniidae, and having a similar 
peculiar sense-organ ; but are distinguished by the presence of a. 
frenulum. The larva seems to be like that of Uraniidae. 

Fam. 41. Pyralidae. This division is to be considered rather 
as a group of families than as a family ; it includes a very large 
number of small or moderate-sized moths of fragile structure, 
frequently having long legs ; antennae simple, only in a few 
cases pectinate ; distinguished from Noctuidae and all the other 
extensive divisions of moths by the peculiar course of the costal 
nervure of the hind wing, which either keeps, in the middle of 
its course, near to the sub-costal or actually unites with it, 
subsequently again separating. Members of the Pyralidae an- 
found in all lands; in Britain we have about 150 species. The 
larvae are usually nearly bare, with only short, scattered setae, 
and little coloration; they have most varied habits, are fond of 
concealment, and are very lively and abrupt in movement, 
wriggling backwards as well as forwards, when disturbed ; a 
cocoon is formed for the metamorphosis. 

The family as a whole consists of Insects of unattractive 
appearance, although it contains some very elegant and interesting 
moths and numerous forms of structural interest. In the genus 
Thiridopteryx little transparent spaces on the wings occur as a 
character peculiar to the males ; the spaces are correlative with 
a greater or less derangement of the wing-nervures. In some 
other forms there is a remarkable retinaculuin, consisting of large 
si-ales, and this, too, is connected with a distortion of the wing- 
nervures. The Pyralidae Pyralites of Eagonot, 1 Pyralidina of 



t, Ann. Soc. ent. France, 1890 and 1891 : and Meyrick, Tr. ent. Soc. 
London, 1890, p. 429. 



vi HETEROCERA PYRALIDAE 421 

Meyrick have recently been revised by two naturalists of dis- 
tinction almost simultaneously ; unfortunately their results are 
discrepant, Meyrick including Pterophoridae and Orneodidae, 
and yet admitting in all only eight families ; while Eagonot does 
not include the two groups named, but defines seventeen tribes 
of the two families Pyralidae and Crambidae that he admits. 
The Pyraustidae of Meyrick is an enormous division including 
the Hydrocampidae and Scopariidae of many authors, as well as 
the Pyraustinae proper and a small group of Poigonot's, the 
Homophysinae. The division Scopariinae is believed to be 
amongst the " most ancient " of Lepidoptera ; the food of the 
larvae consists of moss and lichens. This group is widely dis- 
tributed, being richly represented in Australia, New Zealand, 
and the Hawaiian Islands, as well as in Europe ; and probably 
really occurs wherever their food-plants exist accompanied by 
a tolerable climate. The statistics of the distribution of this 
group, so far as at present known, have been furnished by 
Mr. Meyrick, as follows : European region, about 2 5 species ; 
Madeira, 3 ; St. Helena, ; South Africa, 2 or 3 ; India, 9 ; 
Malayan region, 3 or 4 ; Australia, 24 ; New Zealand, 64 ; 
Hawaiian Islands, 5 ; North America, 1 7 (one of them Euro- 
pean) ; South America, 10. The Hydrocampinae the China- 
marks are of great interest, as being amongst the few forms 
of Lepidoptera adapted for aquatic life. It is believed that all 
their larvae are aquatic, though of only a few is there much 
known. The diversity amongst these forms is of considerable 
interest. The habits of Jfi/drocampa nymphaeata were long since 
described by Reaumur, and have more recently been dealt with 
by Buckler, 1 W. Miiller 2 and Prof. Miall. 3 Although there are 
some discrepancies in their accounts, due we believe to the 
observations being made at different periods of the life and under 
somewhat different circumstances, yet the account given by 
Miiller is we feel no doubt substantially correct. The larvae 
when hatched mine in the leaves of a water-plant for a short 
time thirty hours to three days according to Buckler and are 
completely surrounded by water, which penetrates freely into 
their burrows ; at this period the caterpillar breathes by its 
skin, the spiracles being very small, and the tubes leading from 

1 Ent. Mag. xii. 1876, p. 210, and xvii. 1881, p. 249. 
2 Zool. Jahrb. Syst. vi. 1892, p. 617. 3 Nut. Hist. Aquatic Insects, London, 1895. 



422 LEPIDOPTERA CHAP. 

them closed and functionless. After this brief period of mining 
life, the larva moults and then constructs a habitation by cutting 
a piece out of a leaf, and fastening it to the under side of another 
leaf; it is thus provided with a habitation, but it is one into 
which the water freely enters, and the respiratory apparatus 
remains in the state we have described. The Insect passes 
through several moults, and then hibernates in the water. On 
its revival in the spring a change occurs, and the larva constructs 
a portable, or we should rather say free, habitation out of two 
large pieces of leaf of lens-shape, fastened together at the 
edges ; but the larva has some method of managing matters so 
that the water can be kept out of this house ; thus the creature 
lives in air though immersed in the water. A correlative change 
occurs in the structure of the skin and tracheal system. The 
former becomes studded with prominent points that help to 
maintain a coat of air round the Insect, like dry velvet immersed 
in water ; the spiracles are larger than they were, and they and 
the tracheal tubes are open. One or two moults take place and the 
creature then pupates. There is a good deal of discrepancy in 
the accounts of this period, and it seems probable that the pupa 
is sometimes aerial, sometimes aquatic. Buckler's account of the 
formation of the case shows that the larva first cuts off, by an 
ingenious process, one piece of leaf, leaving itself on this, as on 
a raft : this it guides to a leaf suitable for a second piece, gets 
the raft underneath, and fastens it with silk to the upper portion, 
and then severs this, leaving the construction free ; afterwards the 
larva goes through a curious process of changing its position and 
working at the two extremities of the case, apparently with the 
object of making it all right as regards its capacity for including 
air and keeping out water. He believes that Reaumur was 
correct in his idea that the larva regulates the admission of air 
or of water to the case in conformity with its needs for respiration. 
Miiller calls special attention to the great changes in habit and 
in the structure of the integument during the life of this larva ; 
1'Ui the reader will gather from what we relate as to various 
terrestrial Lepidopterous larvae, that these phenomena are not 
very dissimilar from what frequently take place in the latter; a 
change of h.iluts ;it some particular moult, accompanied by great 
changes in the integument, and even in the size of the stigmata, 
being of frequent occurrence. 



vr HETEROCERA PYRALIDAE 423 



The larva of Xj/rnpluil */";//<"/", a close ally of H. nymphaeata, 
has aquatic habits of a somewhat similar but simpler nature ; 
while N. (Para-pony?} sti'atintittn. is very different. This larva is 
provided with eight rows of tufts of flexible branchiae, occupying 
the position of the spots or setigerous warts usual in caterpillars, 
and reminding one of the spines of certain butterfly-larvae, though 
they are undoubtedly respiratory filaments. These caterpillars 
protect themselves by forming silken webs or cases, or by adopt- 
ing the case of some other larva, and are in the habit of holding 
on by the anal claspers, and rapidly and energetically moving 
the anterior parts of the body in an undulating fashion. The 
spiracles exist, but are functionless. The pupa lives under water, 
and has no branchiae ; but three of the pairs of abdominal 
spiracles are open, and project from the body. Miiller informs 
us that in a Brazilian Paraponyx these three pairs of spiracles 
were already large in the larva, though the other pairs were very 
small, or absent. He considers that the moth of this species 
descends beneath the water of a rapid stream, and fastens its 
eggs on the stems of plants therein. Cataclysta lemnata lives in 
a case of silk with leaves of duckweed attached to it, or in a 
piece of a hollow stem of some aquatic plant ; it is believed to 
breathe, like H. nympliaeata, at first by the integument and 
subsequently by open stigmata : but particulars as to how it 
obtains the requisite air-supply are not forthcoming : the aquatic 
pupa breathes by three large abdominal spiracles like Paraponyx. 

Musotimidae 1 is a small group of two or three genera found in 
Australia and Polynesia ; and the Tiueodidae also consist of only 
two Australian genera. Siculodidae is likewise a small Antarctic 
group, placed by Meyrick in Pyralidina ; but his view is not 
accepted by Snellen and Bagonot. Epipaschiinae (fornierly 
tivated as a separate family) and Endotrichiinae are, according 
to Meyrick, subdivisions of the family Pyralidae proper, an 
enormous group of more than 100 genera, The Chrysauginae 
consist chiefly of American forms, and have not been treated by 
Meyrick ; some of this group have been classed with Tortricid.u- 
or Deltoidae on account of the undulating costa of the front wings 
and the long, peculiar palpi. The Galleriidae are a small group 
including Insects that live in bees'-nests, and feed on the wax 

1 For Bibliographic references connected Avith the divisions of Pyralidae see 
Ragouot, Ann. Soc. ent. France (6), x. 1890, pp. 458, rt<-. 



424 LEPIDOPTERA CHAP. 

etc. ; others eat seeds, or dried vegetable substances. Three 
out of our five British species of this family occur (usually 
gregariously) in bee - hives, and have the peculiar habit 
of spinning their cocoons together. The mass of common 
cocoons formed in this manner by Aphomia sociella is remark- 
ably tough and enduring ; portions of it are not infrequently 
picked up, and as the cocoons are of a peculiar tubular form 
their nature gives rise to some perplexity. 

Phycitidae : is another very large assemblage of Insects with 
very diverse habits. The frenulum and retinaculum are similarly 
formed in the two sexes : the males frequently have the basal-joint 
of the antennae swollen ; hence the term " Knot-horns " applied 
by collectors to these moths. The larvae of the species of 
Epliestia infest groceries, and most children have become to a slight 
extent acquainted with them amongst dried figs ; that of E. 
kuelmiella has become very injurious in flour-mills, its enormous 
increase being due in all probability, to the fact that the favour- 
able and equable temperature maintained in the mills promotes a 
rapid succession of generations, so that the Insect may increase 
to such an extent as to entirely block the machinery. Many of 
the Phycitidae feed on the bark of trees in galleries or tunnels 
constructed partially of silk. A very peculiar modification of 
this habit in Cecidipta excoecaria has been described by Berg. ~ 
In Argentina this Insect takes possession of the galls formed by 
a Chermes 011 Excoecaria Inglandulosa, a Euphorbiaceous tree. 
The female moth lays an egg on a gall, and the resulting larva 
bores into the gall and nourishes itself on the interior till all is 
eaten except a thin external coat ; the caterpillar then pupates 
in this chamber. The galls vary in size and shape, and the 
larva displays much constructive ability in adapting its home to 
its needs by the addition of tubes of silk or by other modes. Some- 
times the amount of food furnished by the interior of the gall is 
not sufficient ; the larva, in such cases, resorts to the leaves of the 
plant for a supplement, but does not eat them in the usual 
manner of a caterpillar; it cuts off and carries a leaf to the 
entrance of its abode, fastens the leaf there with silk, and then 
itself entering, feeds, from the interior, on the food it lias thus 
acquired. Another Phycitid, Dakruma coccidivora, is very 

1 Monograph, by Ragonot, in Romanoff, Mem. Lep. vii. 1893. 

- J-:,if. Z.it. X/.///V, 1.X7S, p. 2:50. 



vi HETEROCERA PYRALIDAE 425 

beneficial in North America by eating large Scale-Insects of the 
Lecanium group, somewhat after the fashion of Erastria scit-ida ; it 
does not construct a case, but shelters itself when walking from 
one scale to another by means of silken tubes ; it suffers from, 
the attacks of parasites. 1 Oxychirotinae, an Australian group, 
is interesting because, according to Meyrick, it possesses forms 
connecting the Pterophoridae with the more normal Pyralids. 

Crambidae, or Grass-moths, are amongst the most abundant 
Lepidoptera in this country, as they include the little pale moths 
that fly for short distances amongst the grass of lawns and 
pastures ; they fold their wings tightly to their body, and 
have a head pointed in front, in consequence of the form 
and direction of the palpi. They sit in an upright position 
on the stems of grass, and it has been said that this is done 
because then they are not conspicuous. Perhaps : but it would be 
a somewhat difficult acrobatic performance to sit with six legs 
across a stem of grass. The larvae are feeders 011 grass, and 
construct silken tunnels about the roots at or near the surface. 
The Ancylolominae are included in Crambidae by Meyrick and 
Hampson. Schoenobiinae 2 are included by Meyrick in Pyraustidae, 
but this view appears not to meet with acceptance, and the group 
is more usually associated with the Crambidae. Most writers 
place the anomalous genus Acentropus as a separate tribe, but it 
is associated by both Meyrick and Hampson with Schoenobius. 
This Insect is apparently the most completely aquatic of all the 
Lepidoptera, and was for long associated with the Trichoptera 
in consequence of its habits and of the scaling of the wings 
being of a very inferior kind. The males may sometimes be 
found in large numbers fluttering over the surface of shallow, but 
large, bodies of water ; the females are rarely seen, and in some 
cases have no wings, or have these organs so small as to be useless. 
The female, it would appear, comes quite to the surface for 
coupling, and then takes the male beneath the water. The larvae 
have the usual number of Lepidopterous feet, and apparently feed 
on the leaves of plants below water just as Lepidopterous larvae 
ordinarily do in the air. 3 They have no trace of gills, and their 

1 Howard, Insect Life, vii. 1895, p. 402. 

2 Monograph by Hampson, P. Zool. Soc. London, 1895, p. 897-974. 

y Discjue, Eat. Zeit. Stettin, li. 1890, p. 59. Cf. also Rebel, Zool. Jahrl. Syst. 
xii. 1898, p. 3. 



426 LEPIDOPTERA CHAP. 

mode of respiration is unknown. A great deal has Leen written 
about these Insects, but really very little is known. They are 
abundant, though local in many parts of North and Central 
Europe ; some of the females have, as we have said, abbreviated 
wings, but how many species there are, and whether the modifica- 
tions existing in the development of the wings are constant in 
one species or locality, are unknown as yet. 

Fam. 42. Pterophoridae : (Plume-moths'). Elegant Insects 
of small size, usually with the wings divided (after the fashion of 
a hand into fingers) so as to form feathers : the extent of this 
division is diverse, but the hind wings are more completely divided 
than the front, which indeed are sometimes almost entire. The 
group is placed by Meyrick in his Pyralidina, but there are many 
entomologists who look on it as distinct. It consists of two 
sub - families, Agdistinae and Pterophorinae, that have been 
treated as families by many entomologists. The Agdistiuae 
(of which we have a British representative of the only genus 
Aydistes) have the wings undivided. Pterophorinae have the 
hind wings trifid or (rarely) cp:iadrifid, the front wings bifid or 
(rarely) trifid. The larvae of the Pterophorinae are different 
from those of Pyralidae, being slow in movement and of heavy 
form, covered with hair and living exposed on leaves ; the pupae 
are highly remarkable, being soft, coloured somewhat like the 
larvae, and also hairy like the larvae, and are attached somewhat 
after the manner of butterfly- pupae by the cremaster : but 
in some cases there is a slight cocoon. There is, however, 
much variety in the larval and pupal habits of the Ptero- 
phoridae, many having habits of concealment of divers kinds. 
We have thirty species of these lovely Plume-moths in Britain. 
The family is widely distributed, and will probably prove 
numerous in species when the small and delicate Insects existing 
in the tropics are more appreciated by collectors. 

Fam. 43. Alucitidae (Orneodidae of Meyrick and others). 
The genus Alucita includes the only moths that have the front 
and hind wings divided each into six feathers. Species of it, 
though not numerous, occur in various regions. The larva and 
pupa arc less anomalous than those of the Pterophoridae, though 
the imago is more anomalous. Tbe caterpillar of our British A. 
polydactyla feeds on the flower-buds of honey-suckle, and forms a 

1 Classification; Mt-yriek, Tr. nit. Sue. London, 1886, p. 1. 



vi HETEROCERA TORTRICIDAE 427 

cocoon. The moth with wings expanded is about an inch across, 
and is a, lovely object. It is not rare, though seldom numerous. 

Fam. 44. Tortricidae. Moths of small size, with a rather 
ample wing area, with the wing-fringes never as long as the 
wings are wide (long across), the hind wings without a pattern : 
the anterior nervure on the hind wings is simply divergent from 
that next to it, and the internal nervure, Ib, is very evidently 
forked at the base. The larvae inhabit their food, which may be 
rolled up or twisted leaves, or the interior of fruits and herbs, or 
galls, or even roots ; they exhibit less diversity than is usual in 
other large series of moths ; all have the normal complement of 
sixteen legs. This group is a very extensive one, but is much 
neglected owing to the great difficulties attending its study ; 
it is not recognised in Hampson's Table of families given on 
p. 370, being there merged in Tiueidae. It appears, however, 
to be a really natural group, and it is not desirable to merge it 
in the sufficiently enormous assemblage of the Tineidae till 
this has been shown to be necessary by the light of a greater 
knowledge of the external anatomy than we possess at present. 
The term Microlepidoptera is frequently met with in entomo- 
logical literature, and should, we think, be confined to the two 
series Tortricidae and Tineidae. The Pterophoridae, and even 
the I'yralidae, have been, and still sometimes are, included under 
this term, but at present it seems best to limit its application as 
is here suggested. 

Three great divisions are at present recognised ; these were 
formerly called by Meyrick, 1 Tortricidae, Grapholithidae, Conchy- 
lidae ; subsequently/ he has adopted the names Tortricidae, 
Epiblemidae, 1'haloniadae. Lord Walsingham, who has devoted 
a great deal of time and study to the elucidation of this most 
difficult group, has suggested 3 that another change is desirable, 
and if so the nomenclature will be : 1. Tortricidae [or Tor- 
ticinae, according to the view that may be taken as to the group 
being family or sub-family] ; 2. Phaloniidae [ = the formerly 
used name, Conchylidae] ; 3. Olethreutidae [ the formerly used 
name Grapholithinae = Epiblemidae, Meyr.]. AYe have upwards 
of 300 species in Britain, nearly 200 of which belong to the 
last division. The name Tortricidae refers to the habit the 

1 P. Linn. Soc. X. S. Wales (2), vi. 18S1, p. 410. 
- Handbook Brit. Ley. 1895, p. 493. y Tr. cut. Soc. London, 1895, p. 495. 



428 LEPIDOPTERA CHAP. 



larvae of these moths possess of rolling up leaves, or twisting 
and distorting shoots and buds. 

The mode in which leaves and shoots are twisted and rolled 
by the very small larvae has been much discussed and is pro- 
bably the result of two or three distinct causes:--!, the 
immediate operations of the larva ; 2, the contraction of silk 
when drying ; 3, changes in the mode of growth of the parts of 
the vegetable, resulting from the interference of the caterpillar. 
The larvae of this family that live in fruits are only too widely 
(we will not say well) known. Stainton gives as the habitat of 
Epinotia fitncbrana, " larva frequent in plum-pies " ; the cater- 
pillar of Carpocapsa pomondla (the Codling -moth) mines in 
apples and pears, and its ravages are known only too well in 
widely distant parts of the world where fruit-trees of this kind 
are cultivated. C. splendana lives in acorns and walnuts ; C. 
jti.li.niM in Spanish chestnuts. Two, if not more, larvae live in 
the seeds of Euphorbiaceous plants, and have become notorious 
under the name of jumpiug-beans, on account of the movements 
they cause. As these latter show no trace externally of being 
inhabited, the movements are supposed to be a mysterious pro- 
perty of the seed ; they are really due to its containing a large 
cavity, extending, in one direction of the seed, nearly or quite 
In >m skin to skin ; in this the larva makes a movement sufficient 
to alter the point of ecjuilibrium of the quiescent seed, or as 
a free body to strike some part of it. The exact nature of the 
movements of the larva have not, we believe, been ascertained. 
There are, at least, two species of these Insects, and two plants 
harbouring them, known in the United States and Mexico, viz. 
Cwrpoccipsa saltitans living in the seeds of Croton colliguaja and 
Grapliolitlia sebudianii.i.e living in the seeds of Sebastiania 
bicapsidaris. 

Fam. 45. Tineidae. Small moths with the labial palpi 
more flexible and mobile than in other moths ; usually separated 
and pointed. Hind wings frequently with very long fringes, the 
wing itself being proportionally reduced in size, and in con- 
sequence pointed at the tip. Larvae very diverse, almost always 
with habits of concealment. The series of forms included under 
this hr.nl is very numerous, the British species alone mounting 
up to 700, while the total described cannot be less than 4000. 
This number, however, must be but a fragment of what exists, 




vi HETEROCERA TIXEIDAE 429 

ii' Mr. Meyrick be correct in supposing that a single one of 
the divisions of the family Oecophoridae comprises 2000 
species in Australia and Xew Zealand alone. As the study 
of these Insects is attended 
with great difficulty on ac- 
count of their fragility and 
the minute size of the great 
majority, it is not a matter 

for surprise that their classiti- 

4%' yt&M 

cation is in a comparatively *^~^T^ +^T 

rudimentary state. AVe shall 

not, therefore, deal with it here. 

^^ . . FIG. '207. Diplosaralignivora(Gelecbiides). 

Neither can we attempt to give Hawaiian islands. 

any idea of the extreme diversity 

in the colours, forms, and attitudes of these small Insects. The 
one shown in Pig. 207, is remarkable on account of the great 
accumulation of scales on the wings and legs. As regards the 
pointed wings and the long fringes, we may remark that it is 
probable that in many of these small forms the wings are 
passive agents in locomotion ; a similar condition of the wings 
is found in other very minute Insects, e.g. Thysauoptera and 
Trichopterygidae ; in all these cases the framework of the wings is 
nearly absent : in some forms of the Tineidae, Opostega, e.g. the 
nervules are reduced to three or four in each wing. The 
variety in habits is as great as that of the external form, and 
the larvae exceed in diversity those of any other group of 
Lepidoptera. Xo doubt a corresponding amount of diversity 
will be discovered in the details of structure of the perfect 
Insects, the anatomy of but few having been at present investi- 
gated. Tinea pellionella has two very important peculiarities in 
its internal anatomy : the testes consist of four round follicles 

u 

on each side, and, contrary to the condition generally prevalent 
in Lepidoptera, are not brought together in a common capsule : the 
two groups are, however, not <|iiite free (as they are in Hfpialv.s), 
but are connected by a loose tracheal network. Even more 
remarkable is the fact also pointed out by Cholodkovsky ' that 
the adult Insect possesses only two Malpighian tubes instead of 
six, the normal number in Lepidoptera; in the larva there are, 
however, six elongate tubes. The group of forms to which 

1 Zool. ^><c. v. 1882. p. 262. 



430 LEPIDOPTERA CHAP. 

Tinea belongs is remarkable for the diversity and exceptional 
character of the food -habits of the larvae ; species subsist 
on dried camel's dung, various kinds of clothes, furs, and 
hair, and even about horns of deer and horses' hoofs : one 
species has been found in abundance in the hair of a live 
sloth, Braili/piis cuculliger, under circumstances that render it 
possible that the larva feeds on the creature's hair, though it 
may feed on minute vegetable matter found in the hair. 
The larva of Tinea vastella is occasionally found feeding on the 
horns of living antelopes. Several species of Tineidae are known 
to devour Scale-Insects. 

Lita solanella is notorious for the ravages it commits on 
stored potatoes. Quite a number of species live on cryptogamic 
matter, or in old wood ; Oinophila v-flavum feeds on the mould 
on the walls of cellars, and is reputed to be injurious by occasion- 
ally also attacking the corks of bottles containing wine. Oecocecis 
yuyonella is said to be the cause of galls on Limoniastrum 
guyonianum, a plant that, growing in the deserts to the south 
of Algeria, is a favourite food of camels, and is frequently entirely 
covered with sand. The deposition of an egg by this moth is 
believed by Gue"nee l to give rise to a gall in which the larva is 
entirely enclosed (like the larvae of the gall-flies). Of Clothes- 
moths there are at least three species widely distributed. 
Trichopliaya, tapctzdla is perhaps entitled to be considered the 
Clothes-moth ; its caterpillar not only feeds on clothes, but spins 
webs and galleries amongst them. Tinea pellionella is also very 
common ; its larva lives in a portable case, while that of the 
third species, Tineola biselliella, forms neither a case nor definite 
galleries. We have found this the most destructive of the three 
at Cambridge'. Clothes or valuable furs may be completely pro- 
tected by wrapping them in good sound paper in such a way 
that no crevices are left at the places where the edges of the 
paper meet. Garments that have become infested may be entirely 
cleared by free exposure to air and sunshine. 

Two species of Tinea have been recorded as viviparous, viz. 
Tinea vim-para in Australia, and an undetermined species in 
Smith Ann-ricii. The species of the genus Sulcnobia in which 
the female is apterous are frequently parthenogenetic. The 
group Taleporiidae, to which this genus belongs, is by some 

1 Ann. Soc. ent. France (4), x. 1870, p. 1, pi. vii. 



vi HETEROCERA TINEIDAE 431 

classified with Psychidae, in which family, as we have pointed 
out, one or two parthenogenetic forms are also known. 

The larvae of Tineidae, though they do not exhibit the 
remarkable armature found in so many of the larger caterpillars, 
are exceedingly diverse. 1 Some are entirely destitute of feet 
(Phyllocnistis). Others are destitute of the thoracic legs ; 
Nepticula is in this case, but it is provided with an increased 
number of abdominal feet, in the form of more or less imperfect 
ventral processes. Some mine in leaves, others live in portable 
cases of various forms. Some are leaf-miners during their 
early life, and subsequently change their habits by con- 
structing a portable case. The genus Coleopliora affords 
numerous instances of this mode of life ; the habits of these 
case-bearers exhibit considerable variety, and there are many 
points of interest in their life-histories. Change of habit during 
the larval life has already been alluded to as occurring in many 
Lepidoptera and is nowhere more strikingly exemplified than in 
certain Tineidae. Meyrick mentions the following case as 
occurring in an Australian Insect, Nematobola orthotricha ; 2 the 
larva, until two-thirds grown, is without feet, and is almost 
colourless, and mines in the leaves of Persoonia lanceolata ; but 
when two-thirds grown it acquires sixteen feet, changes colour, 
becoming very variegate, and feeds externally, unprotected, on the 
leaves. The cases of the case-bearing Tineids are usually of 
small size, and do not attract attention like those of Psychidae. 
A very remarkable one was discovered by Mr. E. E. Green in 
Ceylon, and was at first believed to be formed by a Caddis-worm. 
It has now been ascertained that the Insect forming it is the 
caterpillar of Pseudodoxia limulus, a Tineid moth of the group 
Depressariidae ; 3 the case is composed of minute fragments of 
moss, sand, and lichens ; the anterior end is dilated into a shield- 
like hood that covers and protects the anterior parts of the 
larva when feeding ; the food is mosses and lichens on rocks and 
trees. Before pupating, the larva folds clown the edges of the 
hood over the mouth of the tube, like an envelope, fastening 
them witli silk. The case is fixed to the rock or other support 
and hangs there until the moth appears. 

1 For table of the larvae, according to number of feet and other characters, see 
Sorhagen, Berlin, ent. Zeit. xxvii. 1883, pp. 1-S. 

2 P. Linn. Soc. N.S. Wales (2) vii. 1892, p. 593. 

3 Durraut, Ent. May., xxxi. 1895, p. 107. 



432 



LEPIDOPTERA 



CHAP. 



The family Frodoxidae consists of some Tineids, the larvae of 
which feed in the pods and stems of the Yuccas of south-western 
North America ; they have the mouth of very unusual form 

(Fig. 208, E), and 
some of them, by 
aid of this peculiar 
month, exhibit a 
remarkable modifi- 
cation of instinct. 
The facts are chiefly 
known from the 
observations of 
Eiley 1 on Pro-nulm 
yuccasella, a moth 
living on Yucca 
filamentosa ; this 
plant has been in- 
troduced into our 
gardens in this 
country, where it 
never, we believe, 
produces seed. The 
Yuccas are not 
fitted for self-fertil- 
isation or for fer- 
tilisation by Insect 
agency of an. ordi- 
nary kind. The 
progeny of the 
moth develops in 
the pods of the 
plant, and as these 
cannot grow until 
the flowers have 
been fertilised, the moth has the habit of fertilising the flowers 
at the time she lays her egg in the part that is to develop into 
the pod, and to be the food for her own progeny. The female 
inoth first visits the stamens, and collects, bv the aid of the 




FIG. 208. Pronuba .tynthefim. North America. A, Larva : 
B, C, pupa, ventral and lateral aspects ; D, female moth ; 
E, head and part of thorax of the female moth : a, 
labial palp ; b, maxillary tentacle ; c, maxillary palp ; 
d, proboscis ; e, base of trout leg. (After Riley.) 



1 "The Yucca moth and Yucca Pollination," Hep. Missouri Botanical Garden, 
1892, pp. 99-158. 



vi HETEROCERA PRODOXIDAE -ERIOCEPHALIDAE 433 

maxillae (which in this sex are very remarkably formed), 1 a con- 
siderable mass of pollen, which she holds by means of the peculiar 
maxillary tentacles ; she then lays an egg in the pistil, usually 
of some flower other than that from which she has gathered the 
pollen ; and after she has accomplished this act she carefully 
applies the pollen she had previously collected to the pistil, so 
as to secure the fertilisation of the flower and the development 
of the pod. 

The species of Prodoxus stand in a very peculiar relation 
to Pronuba. They also live in Yuccas, and have habits similar 
to those of Pronuba, with the important exception that, being 
destitute of the requisite apparatus, they do not fertilise 
the Yucca-flowers, and are thus dependent on Pronuba for 
the steps being taken that are necessary for the rearing of the 
progeny of the two kinds of moth. Hence the name of Yucca- 
moth has been bestowed on Pronuba, and that of " bogus Yucca- 
moth " on the Prodoxus. The Pronuba w r e figure is the largest 

O ' O 

and most remarkable species of the genus and fertilises Yucca 
brevifolia ; the larva is destitute of abdominal feet, and in 
the pupa the spines on the back that exist in nearly all pupae 
that live in stems are developed to an extraordinary extent. 
The Yuccas do not flower every year, and the Prodoxidae have a 
corresponding uncertainty as to their periods of appearance, 
passing sometimes a year or two longer than usual in the pupal 
stage. 

Fam. 46. Eriocephalidae. This family has recently been 
proposed for some of the moths formerly included in the genus 
Micropteryx.' 2 They are small, brilliant, metallic Insects, of 
diurnal habits, but are very rarely seen on the wing, and it is 
doubtful whether they can fly much. These little Insects are of 
peculiar interest, inasmuch as they differ from the great majority 
of the Lepidoptera in at least two very important points, viz. 
the structure of the wings and of the mouth-parts. The mouth 
shows that we may consider that the Lepidoptera belong to the 
mandibulate Insects, although in the great majority of them the 
mandibles in the final instar are insignificant, functionless 
structures, or are entirely absent, and although the maxillae are 
t 

1 The maxillary tentacle is considered by Prof. J. B. Smith to be a prolongation 
of the stipes, cf. antea, p. 309 ; also Insect Life, v. 1893, p. 161. 

2 Chapman, Tr. cut. Soc. London, 1894, p. 366. 

VOL. VI 2 F 



434 



LEPIUOPTERA 



CHAP. 



so highly adapted for the tasting of sweets that it is difficult to 
recognise in them the parts usually found in the maxilla of 
inandibulate Insects. Eriocepliala in both these respects connects 
the Lepidoptera with Mandibulata : the mandibles have been 
shown by Walter 1 to be fairly well developed ; and the maxillae are 
not developed into a proboscis, but have each two separate, differen- 
tiated not elongated lobes, and an elongate, five-jointed, very 
flexible palpus. The moths feed on pollen, and use their 
maxillae for the purpose, somewhat in the style we have men- 
tioned in Prodoxidae. The wings have no frenulum, neither 
have they any shoulder, and they probably function as separate 
organs instead of as a united pair on each side : the modification 
of the anterior parts of the hind wing whereby this wing is 
reduced as a flying agent to the condition of a subordinate to the 
front wing does not here exist : the hind wing differs little from 
the front wing in consequence of the parts in front of the cell being 
well developed. There is a small jugum. These characters have 
led Packard to suggest that the Eriocephalidae should be separated 
from all other Lepidoptera to form a distinct sub-Order, Lepidoptera 
Laciniata. 2 The wing-characters of Eriocepliala are repeated as 
to their main features in Hepialidae and Micropterygidae ; but 
both these groups differ from Eriocepliala as to the structure of 
the mouth-parts, and in their metamorphoses. Although Erio- 
<'>'l>!i<i]<i, caltJiella is one of our 
most abundant moths, occur- 
ring in the spring nearly every- 
where, and being easily found 
on account of its habit of sit- 
ting in buttercup-flowers, y't 
its metamorphoses were till 
recently completely unknown. 
Dr. Chapman has, however, 

been able to give US some FlG - 209. Larva of Eriocephala calthella. 

(After Chapman.) A, Young larva from 
information as to the habits side, x 50 ; B, portion of skin with a bulla 

and Structure of the larvae, 111 or f ^all-like appendage : ,, abdominal foot 

of larva. 

both of which points the crea- 

The eggs and young larvae are " quite 




ture is most interesting. 



1 Walter, Jena. ZcHs,-l<r. Xatunv. xviii. 1885. He did not distinguish Erio- 
lilinln as a ^riiiis, ,-is \\-i- have explained <>n ]>. 308. 

2 Amer. \<//nml. xxix. 1895, pp. 636 and 803. 



VI 



ERIOCEPHALIDAE MICROPTERYGIDAE 



435 



unlike our ideas of a Lepidopterous Insect ; " the former have a 
snowy or mealy appearance, owkig to a close coating of minute 
rods standing vertically on the surface of the egg, and often 
tipped with a small bulb. The larva lives amongst wet moss 
and feeds on the growing parts thereof; it is not very similar to 
any other Lepidopterous larva : Dr. Chapman suggests a simi- 
larity to the Slug-worms (Linmcodids), but Dyar is probably 
correct in thinking the resemblances between the two are unim- 
portant : the larva of Eriocephala possesses three pairs of thoracic 
legs, and eight pairs of abdominal appendages, placed on the 
segments immediately following the thorax ; on the under-surface 
of the ninth and tenth abdominal segments there is a sucker, 
trifoliate in form ; this is probably really situate entirely on the 
tenth segment : the body bears rows of ball-appendages, and the 
integument is beautifully sculptured. The head is retractile 
and the antennae are longer than is usual in caterpillars. This 
larva is profoundly different from other Lepidopterous larvae 
inasmuch as the abdominal feet, or appendages, are placed on 
different segments to what is customary, and are of a different 
form. Unfortunately the pupa has not been procured, but there 
is some reason for supposing that 
it will prove to be more like that of 
Tineidae than like that of Microp- 
terygidae. 

The New Zealand genus Palaeo- 
micra is only imperfectly known. 
Meyrick considers it the " most 
ancient " Lepidopteron yet dis- 
covered ; and it would appear that 
its relations are with EriocepJidla 
rather than with Micropteryx. From 
information he has kindly given to 
us, we are able to say that this moth 

possesses mandibles but no proboscis. 

FIG. 210. 

Fam. 47. Micropterygldae.- 

Small moths of metallic colours, 

without mandibles, with elongate 

maxillary palpi : without frenulum : both wings with a complex 

system of wing-veins : on the hind wings the area anterior to 

the cell is large, and traversed by three or four elongate, parallel 




B 




Larva of Micropteryx sp. 
A, Ventral view of the larva, 
magnified ; B, the same, with 
setae unduly magnified. Britain. 



436 



LEPIDOPTERA 



CHAP. 




veins. There are 110 mandibles, but there is a short, imperfect 
proboscis. Larva (Fig. 210) without any legs, mining in leaves. 
The pupa (Fig. 211) is not a pupa obtecta, but has the head 
and appendages free, and it provided with enormous mandibles. 
Although these Insects in general appearance resemble Erio- 
cephala to such an extent that both have been placed in one 
genus, viz. Micropteryx, yet the two forms are radically distinct. 

The most remarkable 

\ / 

point in Micropteryx 
is the metamorphosis ; 
the female moth is 
furnished with a cut- 
ting ovipositor, by the 
aid of which she de- 
posits an egg between 
the two layers of a 
leaf after the manner 
of a saw-fly ; l the larva 
mines the newly-opened 
leaves in the early 
spring, and feeds up 
with rapidity ; it by 
some means reaches 
the ground, and there 
pupates in a firm but 
thin cocoon, with grains 
of earth fastened to it ; 
in this it passes the 

greater part of its life 
IIG. 211. Papa, of Micropteryx (semipurpurella f). A, ' 

Dorsal aspect ; B, C, D, views of head dissected oft' ; as a larva, changing to 
B, profile; C, posterior, D, anterior aspects; m, .-, -,., -1^1-^7 nvlv 

-i . -i i -.-. . t.1 l_/ LI I..' 1 1 \ Cl. y CCl'l. AT J. J.J. 

mandibles. Britain. L J J 

the following spring. 

The pupa is unlike any other Lepidopterous pupa, but is similar 
to those of Trichoptera ; neither the head nor the appendages 
are glued to the body or to one another, but are free, so that 
the pupa can use the appendages to a considerable extent; it is 
furnished with enormous mandibles (Fig. 211, C, D), which are 
detached and shed after emergence. 2 In the interval between 

1 Wood, Ent. May. xxvi. 1890, p. 148. 
- See Chapman, Tr. cut. Soc. London, 1893, p. 255. 





vi MICROPTERYGIDAE 437 

the larval period of feeding and the imaginal instar, the pheno- 
mena of life are essentially like those of Trichoptera. The larva 
has not been, at all satisfactorily studied ; the spiracles appear 
to be excessively minute, but have been ascertained by Dr. 
Chapman to be normal in number and position. 

All the information we possess points to profound distinctions 
between Micropteryx and Eriocepluda, for whereas in the former 
the mandibles drop off from the pupa, so that the imago has no 
mandibles, in the latter the mandibles exist, as they do in 
several other true Lepidoptera. As the history of the mandi- 
bles is not known in other Lepidoptera (where they are present 
in the larva but wanting in the imago), it is premature to 
conclude that no other Lepidoptera suffer the actual loss of the 
mandibles as Micropteryx does, though there is nothing to lead 
us to believe that in any other Lepidopterous pupa are the 
mandibles specially developed as they are in Micropteryx. This 
pupa is in fact quite unique in this Order of Insects. When the 
history of the pupal mandibles is known, we shall be able to 
decide whether they are secondary structures, like the deciduous, 
supplementary mandibles found in Otiorhynchides (Coleoptera, 
Iihynchophora). 



CHAPTER VII 

DIPTERA OR FLIES ; APHANIPTERA- OR FLEAS ; THYSAXOPTERA 

OR THRIPS 

Order VII. Diptera 

Wings two, membranous, usually transparent and never very lar;/< ; 
behind the wings a pair of small erect capitate bodies 
li alteres frequently concealed under membranous hoods. 
No distinct prothorax, all the divisions of the thorax being 
united to form a large mass. Mouth-parts very variable, 
formed for suction not for biting, frequently assuming the 
form of a proboscis that can be retracted and concealed in a 
cleft of the under side of the head. The metamorphosis is 
very great, the larvae bearing no resemblance whatever to the 
perfect Insects, but being usually footless grubs or maggots ; 
frequently the head is indistinct, small, and retracted. 
Pupa variable, either exposed and rather hard, with the 
appendages of the body more or less adherent ; or enclosed 
in a scaly capsule looking like a seed, and when extracted, 
soft and delicate, with the appendages not fastened to the 
body incapable of movement. 

THIS definition of the Diptera, or two-winged flies, is framed 
without reference to the fleas, which are wingless, or to a few 
other parasitic wingless Diptera, such as the sheep-tick. Although 
the Order is of enormous extent, these exceptional cases are 
remarkably few. About 40,000 species of Diptera have been 
discovered, but these are only a tithe of what are still unknown 
to science. The Order is not a favourite one with entomologists, 
and by the rest of the world it may be said to be detested. 
Flies do not display the sort of intelligence we appreciate, 



CHAP. VII 



DIPTERA OR TWO-WINGED FLIES 



439 



or the kind of beauty we admire, and as a few of the 
creatures somewhat annoy us, the whole Order is only too 
frequently included in the category of nuisances that we must 
submit to. Moreover, the scavenger-habits that are revealed, 
when we begin to study their lives, are very repugnant to many 
persons. It is therefore no wonder that flies are not popular, 
and that few are will- 

TTSl 

T 



ing to study them, or 
to collect them for 
observation. jSTever- 
theless, Diptera have 
considerable claims to 
be classed as actually 
the highest of Insects 
physiologically, for it 
is certainly in them 
that the processes of a 
complete life - history 
are carried on with the 
greatest rapidity and 
that the phenomena 
of metamorphosis have 

, f , FIG. 21 2. A Dipteron (Fam. Syrphidae), Cheilosia chryso- 

beeil most perfected. coma . Britain. A, Adult larva ; B, the pupa ; C, 
A mao-o-ot hatchino 1 nymph, extracted from pupa; D, imago. (From 

WeyeubergL.) 

irom an egg, is able 

to grow with such rapidity that the work of its life in this 
respect is completed in a few days ; then forming an impene- 
trable skin it dissolves itself almost completely ; solidifying sub- 
sequently to a sort of jelly, it in a few days reconstructs itself 
as a being of totally different appearance and habits, in all 
its structures so profoundly changed from what it was that the 
resources of science are severely taxed to demonstrate any 
identity of the organs of the two instars. 

A good study of the comparative anatomy of Diptera has never 
been made ; Baron Osten Sacken, one of our most accomplished 
Dipterologists, has recently stated that " the external characters 
of the Diptera have as yet been very insufficiently studied." 
We shall therefore only trouble the student with a few observa- 
tions on points of structure that are of special importance, or 
that he will find frequently alluded to. The head is remarkable 




44 DIPTERA CHAP. 

for its mobility, and is connected with the thorax by a slender 
concealed neck that permits the head to undergo semi-rotation. 
A large part sometimes nearly the whole of the exposed sur- 
face of the head is occupied by the faceted eyes. It is usually 
the case that the eyes are larger in the male than in the female, 
and the sexual discrepancy in this respect may be very great. 
When the eyes of the two sides meet in a coadapted line of union 
the Insect is said to be " holoptic," and when the eyes are well 
separated " dichoptic," The holoptic condition is specially char- 
acteristic of the male, but in some forms occurs in both sexes. 
There is no definite distinction between holoptic and dichoptic 
eyes. The eyes may be enormous, Fig. 238, without actually unit- 
ing, and in the cases where actual contiguity occurs, it takes place 
in different manners. 2 The eyes are frequently during life of 
brilliant colours and variegate with stripes or spots ; this con- 
dition disappears speedily after death, and it is uncertain what 
the use of this coloration may be. 3 The eyes are frequently 
densely set with hairs between the almost innumerable facets. 
These facets frequently differ in size according to their position 
in the organ. The curious double eye of the male Bibio (cf. 
Fig. 224) is well worth notice. There are usually three small 
ocelli placed very near together on the middle of the summit of 
the head. 

The antennae are of considerable importance, as they offer 
one of the readiest means of classification. The families placed 
by systematists at the commencement of the Order have antennae 
similar to those of the majority of Insects, inasmuch as they 
consist of a series of segments approximately similar to one 
another, and arranged in a linear manner (Fig. 213, A). The 
number of these joints is never very great, but reaches sixteen 
in certain Tipulidae, and falls as low as eight in some Bibionidae. 
In certain cases where the antennae, of the male are densely 
feathered (Chironomus, e.g.), the number of joints is in that sex 
greatly augmented, but they are imperfectly separated. This 
form of antenna gives the name Nemocera to the first series of 
Diptera. The majority of flies have antennae of another form, 

1 Ostcn Sacken, Tr. cut. Soc. London, 1884, p. 501, and Berlin, cnt. Zcitschr. 
xxxvii. 1892, p. 423, etc. 

2 Ostrn Sac/ken has recently discussed the intermediate conditions, and proposed 
the ii.-iiin- " |isfii(lhl(iptic " I'm- si. me of them, llcrliii. cnt. Zcitschr. xli. 1896, p. 367. 

3 Girschner, /' /////. ent. /.< ifschr. xxxi. 1887, p. 155. 



VII 



ANTENNAE 



441 



peculiar to the Order, viz. three segments, the outer one of 
which is of diverse form, according to the genus or species, and 
bears on its front a fine projecting bristle, frequently feathered, 
as in Fig. 213, F ; and often distinctly divided into two or more 
joints. This form of antenna is found in the series Aschiza and 
Schizophora ; it is well exemplified in the common house-fly, where 
the organs in question hang from the forehead, and are placed i