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PEON ST eee:

DEPCRTNZ * LSULOGY

THE

CAMBRIDGE NATURAL HISTORY

EDITED BY

S. F. HARMER, Sc.D., F.R.S., Fellow of King’s College, Cambridge ;
Superintendent of ie coe Museum of Zoology

AND

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

VOLUME VI

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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.

dace &

ee

+ SBI Ah NEI PE FEIT
DEPeRi ie Ly ZeULObY |

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2 teed Os OE

London
WACMILLAN AND CO. LIMITED

NEW YORK: THE MACMILLAN COMPANY

IQOI

All rights reserved

‘Men are poor things; I don’t know why the world thinks so
much of them.”—WMrs. Bee, by L. & M. Wintle.

First Edition 1899
Reprinted 1gor

CONTENTS

PAGE
SCHEME OF THE CLASSIFICATION ADOPTED IN THIS Book ‘ E ‘ oe aval
CHEATER
HYMENOPTERA PETIOLATA CONTINUED—SERIES 2. TUBULIFERA OR CHRYsI-
DIDAE.—SERIES 3. ACULEATA—GENERAL—CLASSIFICATION— DIVISION I.
ANTHOPHILA OR BEES. : ; ° : é c : A it
(OUSAIZAMID ey, JUL
HYMENOPTERA ACULEATA CONTINUED—DIVISION II. DIPLOPTERA OR WASPS
—EuMENIDAE, Sonirary TruE Wasps—VESPIDAE, SoctAL WaAsps—
MASARIDAE. 7 , = : : : : , : a é es
CJEPAGEZ Rave El
HYMENOPTERA ACULEATA CONTINUED-——DIviIsion III. Fossorgs or
FossortaL SoLirARyY WASpPS—FAMILY SCOLIIDAE OR SUBTERRANEAN
FossoRES— FAMILY POMPILIDAE 0R RUNNERS—FAMILY SPHEGIDAE OR
PERFECT-STINGERS : ; : ‘ E ; 5 é - A 90
CHVAGR RA Rava
HYMENOPTERA ACULEATA CONTINUED — DIVISION IV. FORMICIDAE OR
ANTS : i : ; F 3 : 5 5 : : y uusil
CCAS AE eave
COLEOPTERA OR BEETLES—STREPSIPTERA ‘ : = : ; = 84

CHAPTER VI

LEPIDOPTERA, OR BUTTERFLIES AND MOTHS . : F ; . ‘ . 304

v1 CONTENTS

CHAPTER Vil

PAGE
DIPTERA, OR FLIES—APHANIPTERA, OR FLEAS—THYSANOPTERA, OR THRIPS 438

CHAPTER VE

HEMIPTERA, OR BUGS—ANOPLURA : A : : a é é a bey

NoTEs . : ! . : : 4 : ; : é : 3 - 602

INDEX . : ; : A é ‘ : 4 < : = (0B:

SCHEME OF THE CLASSIFICATION ADOPTED
IN THIS BOOK

Order Sub- oe aa
HYMENOPTERA ( Petiolata.
(continued from - (continued
[Mall VN \ from Vol. V).

Sub-Family or

Family. Thais. Group.

tee ir { CHRYSIDIDAE (p. 1).
( Archiapides (p. 21).
Obtusilingues (p. 22).
ANTHOPHILA Andrenides (p. 23).
(p. 10) - Denudatae (p. 29).
APIDAE (p. 10) | Seopulipedes (p. 32).
Dasygastres (p. 35).
Sociales (p. 53).
DIPLOPTERA
(p. 71)
| Vise ( (p. 72).
VESPIDAE (p. 78).
| MASARIDAE (p. 88).
Mutillides (p. 94).
FossorEs (p. 90) | Thynnides (p. 96).
| ScoLIIDAE Scoliides (p. 97).
(p. 94) | Sapygides (p. oh
Rhopalosomides ( p- 100).
PoMPILIDAE (p.101).
( Sphegides (p. 107).
Ampulicides (p. 114).
HYMENOPTERA 4 Larrides (p. 116).
(continued) Aculeata 4 Trypoxylonides (p. 118).
(p. 4) SPHEGIDAE Astatides (p. 119).
(p. 107) Bembecides (p. 119).

Nyssonides (p. 123).
Philanthides (p. 124).
Mimesides (p. 127).
| Crabronides (p. 128).

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

Myrmicini
(p. 159).
Attini (p. 165).
Hekenpeee Myrmicides Reaaarate ag
(p. 131) (p. 158) (p. 168).
ig ae
| FoRMICIDAE (p. 169).
| (p. 131) E

Ponerides (p. 170).

(p. 175).
Dorylini

(p. 177).

Dorylides (p. 174)-

|
( Ecitonini

( ( | Amblyoponides (p. 189).

vill SCHEM

BOR (LAS SHALE AION

Sub-Order, Division

Order, or Series,

Lamelli-
es
(p. 190) °

Adephaga or
eae
(p. 200)

|
|

(

COLEOPTERA
(p. 184)

a

Polymorpha
(p. 213)

a

Sub-Family or
Tribe.

5)

Family.

‘ PASSALIDAE (p. 192).

LUCANIDAE p. 193).

( Coprides (p. 195).
f Melolonthides (p. 198).

Sead) 4 Rutelides (p. 198).
Ip? Dynastides (p. 199).
Cetoniides (p. 199).

CICINDELIDAE (p. 201).

ee (p. 206).

Harpalides (p. 206).
Pseudomorphides (p. 206).
AMPHIZOIDAE (p. 207).
PELOBIIDAE (p. 207).

Mormolycides (p. 206).
HALIPLIDAE (p. 209).

DYTISCIDAE (p. 210).

CARABIDAE
(p. 204)

PAUSSIDAE (p. 213).
GYRINIDAE (p. 215).
HYDROPHILIDAE (p. 216).
PLATYPSYLLIDAE (p. 219).
LEPTINIDAE (p. 220).
SILPHIDAE (p. 221).
SCYDMAENIDAE (p. 223).
GNOSTIDAE (p. 223).
PSELAPHIDAE (p. 223).
STAPHYLINIDAE (p. 224).
SPHAERIIDAE (p. 227).
TRICHOPTERYGIDAE (p. 227).
HyYDROSCAPHIDAE (p. 228).
CORYLOPHIDAE (p. 228).
SCAPHIDIUDAE (p. 229).
SYNTELIDAE (p. 229).
HISTERIDAE (p. 230).
PHALACRIDAE (p. 231).
NITIDULIDAE (p. 231).
TROGOSITIDAE (p. 232).
COLYDIIDAE (p. 233).
RHYSODIDAE (p. 234).
CUCUJIDAE (p. 234).
CRYPTOPHAGIDAE (p. 235).
HELOTIDAE (p. 23).
THORICTIDAE (p. 236).
EROTYLIDAE (p. 236).
MYCETOPHAGIDAE (p. 237).
COCCINELLIDAE (p. 237).
KNDOMYCHIDAE (p. 239).
MYCETAEIDAE (p. 239).
LATRIDIIDAE (p. 240).
ADIMERIDAE (p. 240).
DERMESTIDAE (p. 241).
BYRRHIDAE (p. 242).
CYATHOCERIDAE (p. 248).
GEORYSSIDAE (p. 243).
HETEROCERIDAE (p. 243).
PARNIDAE (p. 248).

DERODONTIDAE (p. 244).

(Continued on the next page.)

SCHEME OF CLASSIFICATION

ete eae aint Sub-Family
Soe nee rity ae
, CLOIDAE a ae
| SPHINDIDAE (p. 245).
BOSTRICHIDAE (p. 246).
PTINIDAE Ptinides (p. 246).
(p. 246) Anobiides (p. 246).
Lycides (p. 248).
| MALACODER- | Diatides ae a
MIDAE
(p. 248) Lampyrides (p. 248).
Bs | Telephorides (p. 248).
MELYRIDAE (p. 252).
| CLERIDAE (p. 253)
| LYMEXYLONIDAE (p. 254).
DASCILLIDAE (p. 255).
RHIPICERIDAE (p. 256).
Throscides (p. 260).
[ ksenemides (p. 260).
w| ELATERIDAE j Elaterides (p. 260).
(p. 256). | Perottoni (p. 260).

Order.

Polymorpha
(continued) ~

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

BUPRESTIDAE (p. 261).

TENEBRIONIDAE (p. 263).
CISTELIDAE (p. 264).
LAGRIIDAE (p. 264).
OTHNIIDAE (p. 265).
| AEGIALITIDAE (p. 265).
COLEOPTERA } MoNOMMIDAE (p. 265).
continued) } NILIONIDAE (p. 265).
ar eeeronae Tees (p. 265).
(P. ) PYTHIDAE (p. 265).
PYROCHROIDAE (p. 266).
ANTHICIDAE (p. 266).
OEDEMERIDAE (p. 266).
MORDELLIDAE (p. 267).
CANTHARIDAE (p. 269).
TRICTENOTOMIDAE (p. 275).

( BRUCHIDAE (p. 276)
Eupoda (p. 280).

CHRYSOMEL- (eee ae (p. 281).

Phytophaga IDAE (p. 278 gi ies (p. 282).

Cryptostomes As 282).

(p- 285) Lamiides (p. 287).

ANTHRIBIDAE (p. 290).
CURCULIONIDAE (p. 290).
SCOLYTIDAE (p. 294).
BRENTHIDAE (p. 295).

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

Rhyncho- /

(p. 276)
; fom Cima waver { Prionides (p. 287).
phora (p. eo)

Strepsiptera i
(p. 298) { Sryzopipar (p. 298).

Cerambycides (p. 287).

xX
Order. °
LEPIDOPTERA
(p. 304)

Sub-Order, Division,

4

———

SCHEME OF CLASSIFICATION

or Series.

Rhopalocera

(p. 341)

Heterocera
(p. 366)

Sub-Family or
Tribe.

‘ Danaides (p. 344).
Ithomiides (p. 346).
Satyrides (p. 347).
NyMPHALIDAE | Morphides (p. 348).

(p. 343) Brassclides (p. 349).
Acraeides (p. 350).
Heliconiides (p. 351.
Nymphalides (p. 352).
ERYCINIDAF Erycinides (p. 355).
(p. 354) Libytheides (p. 355).
LYCAENIDAE (p. 356).
PIERIDAE (p. 357).
PAPILIONIDAE (p. 359).
HESPERIIDAR (p. 363.

( CASTNIIDAE (p. 371).

NEOCASTNIIDAE (p. 372).

SATURNIIDAE (p. 372).

BRAHMAEIDAE (p. 374).

CERATOCAMPIDAE (p. 375).

BOMBYCIDAE (p. 375).

EUPTEROTIDAE (p. 376).

PEROPHORIDAE (p. 377).

SPHINGIDAE (p. 380).

CocyTUDAE (p. 382).

NoroDONTIDAE (p. 383).

CYMATOPHORIDAE (p. 386).

SESIIDAE (p. 386).

TINAEGERIIDAE (p. 387).

SYNTOMIDAE (p. 388).

ZYGAENIDAE (p. 390).

HIMANTOPTERIDAE (p. 392).

HETEROGYNIDAE (p. 392).

PSYCHIDAE (p. 392).

COSSIDAE (p. 395).

ARBELIDAE (p. 396).

CHRYSOPOLOMIDAE (p. 396).

HEPIALIDAE (p. 396).

{ CALLIDULIDAE (p. 400).

DREPANIDAE (p. 400).
LIMACODIDAE (p. 401).
MEGALOPYGIDAE (p. 404).
THYRIDIDAE (p. 404).
LASIOCAMPIDAE (p. 405).
ENDROMIDAE (p. 406).
PTEROTHYSANIDAE (p. 406).
LYMANTRIIDAE (p. 406).
HyYPSIDAE (p. 408).
ARCTIIDAE (p. 408).
AGARISTIDAE (p. 410).
GEOMETRIDAE (p. 411).
NocTumpAE (p. 414).
EPICOPEIIDAE (p. 418).
URANIIDAE (p. 419).
EPIPLEMIDAE (p. 420).
PYRALIDAE (p. 420).
PTEROPHORIDAE (p. 426).
ALUCITIDAE (p. 426).
TORTRICIDAE (p. 427).
TINEIDAE (p. 428).
ERIOCEPHALIDAE (p. 433).

| MICROPTERYGIDAE (p. 485).

Family.

SCHEME OF CLASSIFICATION

Sub-Order, Division,

Orde1 on Series Family Sub venily or
CECIDOMYIIDAE (p. 458).
MYCETOPHILIDAE (p. 462).
3LEPHAROCERIDAE (p. 464).
CULICIDAE (p. 466).
CHIRONOMIDAE (p. 468).
Orthorrha- ORPHNEPHILIDAE (p. 470).
pha Nemo-} PsycHODIDAE (p. 470).
| Hae DIxIDAE (p. 471).
| (p. 455) Toran ave {aie (p. 472)
(p. 471) imnobiinae (p. 473).
| Tipulinae (p. 475).
BIBIONIDAE (p. 475).
| SIMULIIDAR (p. 477).
| _RHYPHIDAE (p. 478).
|

( STRATIOMYIDAE (p. 478).
LEPTIDAE (p. 479).
TABANIDAE (p. 481).
ACANTHOMERIDAE (p. 483).
THEREVIDAE (p. 484).

Orthorrha- SCENOPINIDAE (p. 484).
pha Bra- NEMESTRINIDAE (p. 484).
chycera BoMBYLIIDAE (p. 485).
(pp. 455, | ACROCERIDAE (p. 489).
DIPTERA 478) LONCHOPTERIDAE (p. 490).

pe es) MyYDAIDAE (p. 491).

ASILIDAE (p. 491).
APIOCERIDAE (p. 492).
EMPIDAE (p. 492).

| DoLICHOPIDAE (p). 493).

Cyclorrha- ( PHORIDAE (p. 494).
pha As- PLATYPEZIDAE (p. 496).
ama PIPUNCULIDAE (p. 496).

(pp. 455, ) Conoprpas ( (p. 497).
494) SYRPHIDAE (p. 498).
Muscipak ACALYPTRATAR (p. 503).
Cyclorrha- ANTHOMYIIDAE (p. 506).
pha Schi- Lee (p. 507).

(pp. 456, | SARCOPHAGIDAE (p. 510).
503) MuscIpDAE (p. 511).
OESTRIDAE (p. 514).

HIpPpoposcIDAE (p. 518).
| BravuLtpas (p. 520).
STREBLIDAE (p. 521).
NYCTERIBIIDAE (p. 521).

eee a
(pp. 456,
517)

pee 3 XIIDAE (p. 510).

APHANIPTERA / z

(pp. 456, 522) 1 PULICIDAE (p. 522)

THYSANO- f Terebrantia (p. 531).
PTERA (p. 526) \ Tubulifera (p. 531).

Xl

SCHEME OF CLASSIFICATION

Order. Sub-Order. Series.

GYMNOCER-
INGUIN (oh
544)

Heteroptera
(pp. 548, 4
544)

HEMIPTERA
(p. 532)
ee

ATA (p.
562)

TRIMERA
544)

(p.

Homoptera
(pp. 543,

568) DIMERA

544)

(p-

| MoNOMERA
| (p. 544).
| Anoplura

(p. 599.)

Family.
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).

a

GALGULIDAE (p. 562).
NEPIDAE (p. 563).

} NAUCORIDAE (p. 565).
BELOSTOMIDAE (p. 565).
NOTONECTIDAE (p. 567).

CoRIXIDAE (p. 567).

CICADIDAE (p. 568).

[ Fence RIDAE (p. 574).
MEMBRACIDAE (p. 576).

| CERCOPIDAE (p. 577).
JASSIDAE (p. 578).

{ PSYLLIDAE (p. 578).
APHIDAE (p. 581).

ALEURODIDAE (p. 591).

{ CoccIDAE (p. 592).

{ PEDICULIDAE (p. 599).

CHAPIER 1
HYMENOPTERA PETIOLATA CONTINUED

SERIES 2.' TUBULIFERA OR 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.

Trochanters undivided ; the hind-body consisting of from three
to five visible segments; the female with an ovipositor,
usually retracted, transversely segmented, enveloping a fine,
pointed style. The larvae usually live in the cells of other
Hymenoptera.

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.!- 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

' Systematic monograph, Moesary, Budapest, 1889. Account of the European
Chrysididae, R. du Buysson in André, Spee. gen. Hym. vol. vi. 1896.
VOL. VI £ B

bo

HYMENOPTERA 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
Cleptes the abdoren
is, however, sinilar
in form to that of
the Aculeate Hymen-
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 Ruby-
flies have the same
number of segments
as other Hymenoptera Petiolata. The difference in this re-
spect 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 be
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 fine rods or spiculae.
There are no poison glands, except in Cleptes, which form appears
to come very near to the Aculeate series. Some of the Chrysi-
didae on occasions use the ovipositor as a sting, though it is only
capable of inflicting a very minute and almost innocuous wound.

Although none of the Ruby-flies attain a large size, they are
usually very conspicuous on account of their gaudy or brilliant
colours. They are amongst the most restless and rapid of Insects ;

Fic. 1.—Chrysis ignita, 9. England.

I CHRYSIDIDAE

Oo

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. He noticed Chrysis
ignite flying about the cell of Odynerus parietum, a solitary
wasp that provisions its nest with caterpillars; in this cell the
Chrysis 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 Chrysis 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 1s, 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 Chrysis 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 ot
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 Chrysis 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 Ent. Mag. vi. 1869, p. 153.

A HYMENOPTERA CHAP.

after emerges a perfect Chrysis of the most brilliant colour, and
if it be a female indefatigable in activity. It is remarkable
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 Chrysis 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—Jonema 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. shang-
haiensis really lives on a Hymenopterous parasite in the cocoon
of the Lepidopteron.

Parnopes carnea frequents the nests of Lembex rostrata, a
solhtary 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
Parnopes takes advantage of this habit by entering the cell and
depositing there an egg which produces a larva that devours that
of the Bembex. 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 5. Hymenoptera Aculeata.

The females (whether workers or true females) provided with a
sting: trochanters usually undivided (monotrochous). Usually
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 Tubulfera, 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 great series, characterised by the fact that the ovipositor is

I ACULEATA

un

withdrawn into the body in a direct manner so as to be entirely
internal, whereas in the -Parasitica 1t 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 two divisions characterised by
the fact that the ovipositor in one retains its function, the egg

oo

ce
Y VA = —— Fic. 2.—Diagram of upper sur-

O SS face of Priocnemis affinis ¢,

}) eC & Pompilidae. 0, ocelli; J,

A pronotum ; 4”, mesonotum ;
0° @) re Be, scutellum of mesonotum ;

Be? Me paises ee. B4, post-scutellum or middle

part of metanotum ; B?, propo-
deum or median segment (see
vol. v. p. 491); b°, combing
hairs, pecten, of front foot: C',
first segment of abdomen, here
not forming a pedicel or stalk +
D*, coxa; b?, trochanter; LD,
femur ; D®, calearia or spurs
of hind leg: 1 to 15, nerv-
ures of wings, viz. 1, costal ;
2, post-costal ; 3, median ;
4, posterior; 5, stigma ;
6) mancinaly- 7, wpper
basal ; 8, lower basal’; 9, 9,
cubital; 10, the three sub-
marginal ; 11, first recurrent ;
12, second recurrent; 1:3,
anterior of hind wing; 14,
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.

y

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."

We have dealt with the external anatomy of Hymenoptera in

1 For new views on this subject see note on p. 602.

6 HYMENOPTERA CHAP.

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’ 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
Fic. 3.—Sting of bee. A, One of the needles chamber, through Which the

Separated ; a, the barbed point ; 6, piston; needles play ; the needles are

¢, arm. 8B, Transverse section of the 5 :

sting: dd, the two needles; e, bead for Cach provided with a pro-

ad af Sa eee: jpceing eee wane oe bas

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 projections on the needles, acting after the manner of a piston,
carry the poison before them. The needles are so arranged on

1 Ann. Sci. Nat. (7) ix. 1890, 105 Ie

I ACULEATA—STING—LARVA jf

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 ' 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 ¢. R. Ac. Paris, exviii. 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 Hymenoptera.
Sir S. Saunders gives the following statement as to the larva of
a wasp of the genus Psiliglossa,’ just before it assumes the pupal
form: “ The respective segments, which are very distinctly indi-
eated, may be defined as follows:—The five anterior, mcluding
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 Ratzeburg.’
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 Forel’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” 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. ent. Soc. London, 1873, p. 408. 2 Zeitschr. wiss. Zool. xxv. 1875, p. 184.

I ACULEATA—_DEVELOPMENT 9

(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
body 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 C

of the pair that form the director are yy¢, 4 Development of sting of

differentiated secondarily from the
primary pair of these masses of hypo-
dermis. A good deal of discussion
has taken place as to whether the

the bee: A and C, ventral ; B,
side view. A, End of abdomen
of adult larva: a, 6, c, d, the
last four segments, c being the
eleventh body segment, 11 ;
6 bearing two pairs, and ¢ one
pair, of rudiments. B, Tip of

) nye ~ Ss = I
oe onenT iia ts of the : suing abdomen of adult bee: 9, the
gonapophyses—are to be considered as ninth, d, the tenth body seg-

ment. C, Rudiments in the
early condition as seen within
the body: ¢, first pair ; 4, the
second pair not yet divided into
two pairs ; 6”, ec’, commence-
ment of external growths from
the internal projections. (After
Dewitz.)

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." 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.” 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.

1 Morph. Jahrb. xxiv. 1896, p. 192. ? Zeitschr. wiss. Zool. xxx. 1878, p. 78.

10 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 ' 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 back to tegulae. 2. Entomophila

[ = Fossores part]
Pronotum reaching back 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 invegular 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 Ashmead’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, usually 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 great 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. Washington, iii. 1896, p. 334.

I ANTHOPHILA——BEES II

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! 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 fune-
tion of this kind of hairs is
unknown; Saunders suggests’ 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 plumose, some are simple, Tt,5-—Hals of Bes Ay simple bat
as shown in Fig. ap A, and this is hair from abdomen of Megachile ; C,
specially the case with the hairs Deda ihre ae ee
that are placed at the edges of dorsata ; E, from thorax of Prosopis.
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, eg. 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. ent. Soc. 1878, p. 169.

12 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 (rig. 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.' The legs also assist in arrang-
ing the pollen on the other parts of

the body. The males do not carry
Fic. 6.—A, Worker of the honey-bee pollen, and though their hind-legs

(Apis mellifica), with pollen plates ; 9 :

laden ; B, basal portions of a are also highly modified, -yet the

Sone il a eee modifications do not agree with

hairs and grains of pollen; C, one those of the female, and their func-

eee cg tes Cueerans: tions are in all probability sexual.
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, where 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

* The mode of wetting the pollen is not clear. Wolff says it is done by an exu-
dation from the tibia; H. Miiller by admixture of nectar from the bee’s mouth.
The latter view is more probably correct.

1 BEES—PROBOSCIS I

Oo

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’ is a very complex organ, and in its
extremely developed forms exhibits a comphcation of details and
a delicacy of structure that elicit the admiration of all who study
it. In the lower bees, however, 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. he 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 be 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 that 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.

14 HYMENOPTERA CHAP.

submentum is articulated with the mentum in 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 submentum. The lorum is a
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 ylo-
copa, Where the submentum is unusually short, this part reposes

Z

Fic. 7.—Side view of basal portions of proboscis of Bombus. a, Epipharyngeal sclerites ;
6, arrow indicating the position of the entrance to pharynx, which is concealed by
the epipharynx, ¢ ; d, hypopharyngeal sclerites ; e, vacant space between the scales
of the maxillae through which the nectar comes: J, lobe ; f’, stipes; g, cardo of
maxilla: h, encephalic pillar on which the cardo swings ; 7, angle of junction of lores
and submentum lorum ; %, mentum ; /, 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 there 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-

I BEES——PROBOSCIS m5

pharynx; one pair of these strap-like pieces extends backwards
from the two sides of the base of the epipharynx; Huxley called
them sclerites of the oesophagus; a better name would be
epipharyngeal sclerites (Fig. 7, @): the other pair pass from the
terminations of the epipharyngeal sclerites, 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 sclerites, 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.

Returning 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.’ 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 invaginated; 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.”

16 TIYMENOPTERA 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 will 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 Huglossa (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.
Fic. 8.—Transverse section of ligula The whole of the proboscis 1s usually

of honey-bee, diagramatic. A, called the tongue; this, however, is

With the long sac invaginate. B, : S : .

evaginate: a, chitinous envelope @dmittedly an erroneous application
with the bases of the hairs; 6, of this term. The terminal deli-
rod ; c, groove of rod; d, lumen j :

due in A to invagination of the Cate, elongate, flexible organ 18 by

rol, in B to its evagination; x, some called the tongue; but this
nerve ; ¢7, trachea, 2

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.! The application
of the terms mentum, submentum and hypoglottis is open to the
same doubts that exist with regard to them in so many other

’ For figures and descriptions of the proboscides of British bees, refer to E. Saun-
ders, Jowr. Linn. Soc. xxiii. 1890, pp. 410-432, plates III.-X.: and for details of
the minute structure and function to Cheshire, Bees and Bee-keeping, vol. 1.

F BEES—PROBOSCIS 17

Insects, and we have omitted the term hypoglottis 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
sheht 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, ¢) so that a
cup or cavity 1s 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

Fic. 9.—A, Proboscis of a “long-tongued”’ bee,

the cup ) ; tb Is between Anthophora pilipes ; B, lower, C, upper view

these two parts that the of proboscis of an “obtuse-tonguéd” bee,

Prosopis pubescens. a, Labrum ; 4, stipes ;

fluid passes to reach the c, palpiger; d, scale: f, lobe; g, pa)pus;
pharynx. It is no doubt h, cardo, of maxilla: 4, lorum ; #, submentum ;
Med 7, mentum ; m, labial palp ; 7, paraglossa ;
to shght movements of the o, ligula; p, tip of ligula (with “spoon ”’ at tip
membranous parts of the Ae some of the hairs more magnified) ; g,
1ypopharyngeal sclerites.
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,
MOIS Vir C

18 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 ' 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,” 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 :* 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 lp, 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

' Breithaupt, Arch. Naturges. lii. Bd. i. 1886, p. 47.
2 See Fig. 26, p. 71. 3 Bull. Mus. Paris, i. 1895, p. 38.

1 BEES—HABITS 19

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 Pérez and Bellesme,!
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 Hymenoptera, 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
of 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 O.R. Ac. Paris, \xxxvii. 1878, pp. 378 and 535.

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 eases 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 Hymenopterists 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.
Kftorts have recently been made to establish an improved classifi-
cation, but as they relate solely to the European bees they are
insufticient 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)
Dasygastres, those that carry it under the abdomen; some of the
parasitic and other forms have been separated as (5) Denudatae
(or Cuculinae); the Bombi and the more perfectly social bees
forming another group, viz. (6) Sociales. A group Andrenoides,
or Panurgides, was also proposed for certain bees considered to
belong to the Apidae though exhibiting many points of resem-

2)

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* recognises no less
than fourteen primary divisions, but is far from satisfactory.

The two genera Prosopis and Sphecodes have been recently
formed into a special family, ARCHIAPIDAE, by Friese,” 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 hgula 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 Fic. 10.— Prosopis signata, Cam
Prosopis must be parasites. This ee A, SEES B, front of

read of female ; C, of male.

is, however, known not to be the

ease 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 im 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 lquid than usual, and has. been
supposed to be entirely honey, since they have no apparatus for
earrying pollen. Mr. R. 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 Hymenopterorum, Leipzig, 10 vols. 1892-96 ; Bees, vol. x.

2 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 HYMENOPTERA CHAP.

has ascertained that a few of the very numerous Hawauan 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, by some of the non-
parasitic forms, enables the bee to sweep the pollen towards its
mouth. These observations show that the structural peculiarities
of Prosopis 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 Col/etes 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, ine 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 Obtusilingues
may be apphed 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
difference 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 Sphecodes
examples of industrial bees extremely ill endowed for their work.
tecent observations tend, however, to prove that Sphecodes are to
a large extent parasitic at the expense of bees of the genera
Halictus and Andrena. Breitenbach has taken S. rubicundus out
of the brood-cells of Halictus quadricinctus ; and on one of the few

I 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
Andrena nigroaenea. Marchal’ has seen S. subquadratus fight
with Halictus 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
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.
Friese states that in Sphecodes the first traces of collecting
apparatus exist; and, accepting the condition of affairs as being
that mentioned above, itis 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

Fie. 11.—Sphecodes gibbus 2. Britain.

1 Bull. Soc. ent. France, 1894, p. exv.
* Marchal, Rev, Sci. 15th February 1890, and Ferton, f.c. 19th A’pril.

24 HYMENOPTERA CHAP.

of great interest and variety. Fabre observed //. Jineolatus and
H. sexcinctus' 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. Pérez, 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 up to the end of

USCOR. Ae, pParis. exe 01879) p. 1079, and Ann. Sci. Nat. (6), ix. 1879, No. 4.
* Act. Soc. Bordeaux, xlviii. 1895, p. 145.

I BEES—ANDRENIDES 25

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
generations of the same species may vary.
Verhoeff has described the burrows of
Halictus quadrieinctus 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
em., 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 Fic. 12.—Nesting of Malictus
the shaft. The larvae that are present ee Hie aes
in the brood-chambers at any one _ thereto; m, retreat or con-
: 6 b tinuation of the burrow ; w,
moment. differ much in their ages, a the vaults; s, the accumula-
fact that throws some doubt on the tion of cells, (After Ver-
F : hoeff, Verh. Ver. Rheint. xlviii.
supposed parthenogenetic generation. 1891; scale not mentioned. )
No cocoons are formed by these Ha/ictus,
the polished interior of the cell being a sufficiently refined resting
place for metamorphosis. Verhoeff states that many of the
larvae are destroyed by mouldiness; this indeed, he considers to
be the most deadly of the enemies of Aculeate Hymenoptera.
The nest of Halictus maculatus has also been briefly described
by Verhoeff, 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
eround, preferring sandy places, but frequently selecting a gravel
path as the locality for their operations; they nearly always live

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 Stylops, by which their form is more or less altered.
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 hght 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. R. C. L. Perkins
recently captured a specimen of Halictus
rubicundus, from 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
ran cae ee Lh the bee then maintains the parasite in its
drena aterrima. (After position, and at the same time supplies it
vane 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.

I BEES——-ANDRENIDES Oh

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
earries loads of pollen,each about
half its own weight, to its nest.
The habits of this insect have been described by Hermann
Miiller.’ 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

Fic. 14.—D. hirtipes 2. Britain.

l Verh. Ver. Rheinland, xii. 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 bee
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 Chalicodoma,
referred to on a subsequent page. If interrupted while storing a
ehamber 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
not 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 growth the larva becomes red and increases in weight from °0025
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 condi-

I BEES——DENUDATAE 29

tion of the integument, for this last exuvium is thick and rigid.
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
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 Ste/is larva emerges from the
ego 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 Stelzs at the lower,
end thereof. By the consumption of the provisions the two larvae
are brought into proximity, and by this time the S/e/is 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. Verhoetf
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.

30 HYMENOPTERA CHAP,

Fabre has made us 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 difticult 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 Colletes daviesanus
and its parasite Hpeolus variegatus
both specially affect the flowers of
Tanacetum vulgare, Some of the
parasitic bees have a great resem-
blance to their hosts ; Stelis signata,
for instance, is said to be so like
; Anthidium strigatum that for many
Fig. 15.--Nomada sex-fadciata 9, ‘yeate ab owas considered: ombeua
erat 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 Andrena have no 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 Vomada flies about unmolested ; indeed, according
to Friese, it is treated as a welcome guest. He says he has often
seen, and in several localities, Vomada lathburiana and Andrena
ovina flying peacefully together. The Nomada 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 1s really not
so, for, as we have before had occasion to observe, there 1s not the
slightest reason for believing that host Insects have any idea
whatever that the parasites or inquilines are injurious to their

I PARASITIC BEES—-DENUDATAE

W
_—

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 Momada, 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? Ifthe species of Vomada
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
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

Fic. 16.—Melecta luctuosa ?. Britain.

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 Anthophora,
when the two bees meet in the burrows of the Anthophora.

We shall have occasion to remark on some of the habits of
Dioxys cineta 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 cyanoxantha, 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 Diovys are
produced, offering another example of the fact that the quantity
of food ingested during the lifetime of the larva does not intlu-
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 Anthophora. It has
recently been merged by Friese with Andrenides into a single
group called Podilegidae. Four British genera, Ceratina, Antho-
phora, Eucera and Saropoda (including, however, only seven

1 It is impossible for us here to deal with the question of the origin of the para-

sitic habit in bees. The reader wishing for information as to this may refer to
Prof. Pérez’s paper, Act. Soc. 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 Bombi is exhibited, indeed the
female of one of our species of Anthophora is so very like the
worker of Lombus 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. personata 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, J/eloe, as well as by a bee of the genus Melecta.
The genus Yylocopa* 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, 1. violacea 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. A. violacea 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 Réaumur, 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 Réaumur 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

' Refer to p. 70 postea, note, as to a recent discovery about Xylocopa.
VOLS val 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. We have figured the larva and pupa of this species in
the previous volume (p. 170).

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

Fic. 17.—Xylocopa (Koptorthosoma), sp. near flavonigrescens, §. 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 Eneyrtus: 300 specimens of the parasite
have been reared from a single larva of the bee; two-thirds of
the larvae of this bee that Horne endeavoured to rear were
destroyed by the little Chalcid.:

The most beautiful and remarkable of all the bees are
the species of Huglossa. 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 Huglossa 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

I DASYGASTRES—MASON-BEES 35

social; but this is doubtful, Bates having recorded that 2. swr-
inamensis forms a “solitary nest.” Lucas concluded that
cordata is social, on the authority of a nest containing “a
dozen individuals.” No workers
are known. The species of
Hulema have a shorter tongue
than Fuglossa, and in form
and colour a good deal re-
semble our species of Lombus
and Apathus.

The group DASYGASTRES 1n-
cludes seven European genera,
four being British (Chelostoma
being included in Heriades).
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 (Megachile,
e.g.) the labrum is very large,
and in repose is inflected on
to the lower side of the head,
and closely appled to the
doubled-in tongue, Which it Fic. 18. — Euglossa cordata, 6. Amazons.
serves to protect ; the man- A, The Insect with extended proboscis ;
dibles then lock together out- B, outer face of hind tibia and tarsus.
side the labrum, which is thus completely concealed. This group
includes some of the most interesting of the solitary bees.

The genus Chalicodoma 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 Réeaumur 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 Bombus ; 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

30 HYMENOPTERA CHAP.

suitable sort of earth with the material secreted by its own
salivary glands; the amount of cement used is reduced hy 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, lke
other Dasygastres, carries the
polen 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
Frc. 19.—Chalicodoma muraria. Greece. a fresh cargo; after a few loads

Bete eee cma. 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 days,
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 or 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 Leucospis gigas succeeds
in perforating the masonry and depositing therein an egg, so that

DD?

a Leucospis is reared in the cell instead of a Chalicodoma.

I DASYGASTRES—MASON-BEES O7/

This Insect has been the object of some of J. H. Fabre’s
most instructive studies on instinct.’ 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 Chalicodoma is rooted
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. Réaumur 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 away ; 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

T Souvenirs entomologiques. 4 vols. Paris, 1879 to 1891.

HYMENOPTERA CHAP.

Ww
oe

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 Pérez 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 do 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 Réaumur. 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 1s
specially interesting, because 1t would appear that the two com-
batants did not seek to injure one another, but were merely
engaged in testing, as it were, which was the more serious in its
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 1s 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 always 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 hberated 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; andit 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

.

l The ‘‘ Chalicodome des galets”’ or C. ‘‘des murailles” of the French writer ; in
some places he speaks of the species as being C. muraria, in others as C. parietina.

40 HYMENOPTERA CHAP.

its nests to the small boulders brought down and left by the
Rhone 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 Lembex. A cross experiment was made by taking
away the stone with the attached nest of the bee while the latter

I DASYGASTRES

-MASON-BEES

INSMTINGL 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 Chalei-
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 1s 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, Ste/is nasuta, breaks
open the cells after they have been completely closed and places
its own eggs in them, and then again closes the cells with
mortar. The larvae of this Ste/is 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 Dioxvys, 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 1s much the
same as in the case of the Sée/is, 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 Dioxys 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.
Leucospis gigas and Monodontomerus nitidus ; the habits of which
we have already discussed (vol. v. p. 543) under Chalcididae.
Lampert 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 the 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

I DASYGASTRES—CARDER-BEE 45

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

The bees of the genus Anthidiwm 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; 1t 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 nidification.
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, Fic. 20.—Anthidium manicatum, Carder-
almost as large as itself, it flies pees ged toate 9 Batemele
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 Anthidiwm 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. manicatum 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 stems,
has been specially observed by Fabre; it will take the cotton for

46 HYMENOPTERA 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 1t 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
Leucospis, which insinuates its eggs through the sides, and
frequently destroys to the last one the inhabitants of the
fortress. Fabre states that these Anthidiwm,as well as Megachile,
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 Osmia,
place the excreta in a vacant space. The Anthidiwm adopts,
however, a most remarkable system: about the middle of its

1 DASYGASTRES—ANTHIDIUM AV,

larval life it commences the expulsion of “frass” in the shape of
small pellets, which it fastens together with silk, as they 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 theseare Anthidium septemdentatumand

A, bellicosum. 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. bellicosum, 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 oceupied by
the Anthidium ; the result of this is that when the metamorphoses
are completed, the latter bee is unable to effect its escape, and

48 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.

Osmiw 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
Fic. 21.—Osmia tricornis, 9. Algeria. our disposal, and we must
content ourselves with a con-

sideration of one or two of the more instructive of the traits

of Osmia life. O. tridentata forms its nest in the stems 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

ae

I DASYGASTRES—OSMIA AQ

thus made is stored with pollen and honey, and an egg is
deposited. Then a barrier has to be 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 (Odynerus 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 was 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 be 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
\WAQIE, WIL 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. Solenvus 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 51

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 suftice 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 Megachile it is well to state that
the bee, the habits of which were described by Réaumur under
the name of “Vabeille 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 Réaumur’s work, founded the
genus Anthocopa for the bee in question. Megachile 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 orgam 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

52 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. JZ. albocincta 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 lke
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 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.

Horne has given particulars as to the nest of Megachile anthra-
cina (fasciculata), an Kast Indian species." 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 ld is made for the
receptacle. The cells are about
the size and shape of a common
thimble; in one specimen that
Horne examined no less than
thirty-two pieces of leaf disposed
in seven layers were used for one
cell, in addition to three pieces for
the round top. The cells are
varefully prepared, and some kind
of matter of a gummy nature is
Fic. 22.—Nidification of leaf-cutting believed to be used to keep in

bee, Megachile anthracina. A, one place the pieces forming the in-

eae Gan aa? terior layers. The cells are placed

part of astring of the cells. (After end to end, as shown in Fig. 22, B;
Horne. ) 3 ( :

five to seven cells form a. series,

and four or six series are believed to be constructed by one pair

' Trans. Zool. Soc. London, vii. 1870, p. 178.

I BEES—DASYGASTRES—SOCIALES 53

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 (Aeniceros bicornis).

Megachile lanata is one of the Hymenoptera that in Kast
India enter houses to build their own habitations. According
to Horne 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.” JZ proxima, a
species almost exactly similar in appearance to JZ Janata, makes
its cells of leaf-cuttings, however, and places them in soft soil.

Fabre states that Jf 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 wp 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 (Socrates) 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 Bombus (Fig.
3931, vol. v.), with its commensal genus or section, Psithyrus
(Fig. 23); Melipona (Fig. 24), in which T7rigona 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 Bombus 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.

Hoffer has given a description of the commencement of a
society of Bombus lapidarius.’ 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
was lined with honey-saturated pollen; then several eggs were
laid in this receptacle, and it was entirely closed. Hoffer 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 Bombus agrorum he was able to describe
the subsequent operations ; these are somewhat as follows :—The
first 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, Hoffer 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.

I SOCIALES——BUMBLE-BEES 55

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-hke 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 worker-cells with food before the
egos 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 (Bombus) 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 we 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
BLombus may attain the number of 300 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
bumble-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 Lombus 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, yet 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. Fea 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 oclock 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 Bombus 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.
Cholodkoysky,' on whose authority this statement as to the sting
is made, has not examined it in 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 Zool. Anz. vii. 1884, p. 312.
2 SB. Ges. Wien. xxxviii. 1888, p- 34.

I SOC S— SAV IIE MAO'S 59

species of Lombus 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. arenicola, B. pratorum and B. scrimshiranus. 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 ; £. terrestris, B. hortorum, B. lapidarius
and £. pomorum are examples of these variable species. The
yariation 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. With 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 (Apathus 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. Psithyrus
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

60 HYMENOPTERA CHAP.

no harm beyond appropriating a portion of their food supplies.
Schmiedeknecht says they are commensals, not parasites; but 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
more definite, the
vague ideas that
have been generally
prevalent | among
entomologists. He
found and _— took
home a nest of Bom-
bus variabilis, which

Fic. 23.—Psithyrus vestalis, Britain. A, Female, x 3 ; anoalee tale
B, outer side of hind leg. containec also a

female of Psithyrus
campestris, so that he was able to make observations on the two.
The Psithyrus was much less industrious than the Lombus, 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 Psithyrus 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 Psithyrus itself, for 1t can scarcely
be supposed that the Bombus carries its complaisance so far as
to construct a cell specially adapted to the superior stature of its
uninvited boarder. When a Psithyrus 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 Psithyrus to a nest that had not previously had one as an in-
mate. The new arrival caused a great hubbub among the Lombus,
which rushed to it as if to attack it, but did not do so, and the
alarm soon subsided, the Psithyrus taking up the position in
the nest usually affected by the individuals of the species. On

li

I SOCIALES—STINGLESS BEES 61

introducing a female Psithyrus to a nest of Bombus in which a
Psithyrus 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 Psithyrus 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 Lombus, who pulled it about but did not attempt to sting it.

When Psithyrus 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 BLombus 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 Ist of September the assemblage consisted only of a
queen Lombus and fifteen workers, together with eighteen speci-
mens of the Psithyrus, eight of these being females.

The nests of Lombus 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, Merops apiaster, 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 be 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! 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 Zrigona 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

Rem i Melipona interrupta (fasei-
Pig. 28.—Afelipona sp. @.--AINaZONs i latan) se alteeie aloo no Nesp hier
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 Ent. Nachr. xii. 1886, p. 177. 2 Tr. ent. Soc. London, 1868, p. 133.

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.

Hockings* 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 1s supposed, 77rigona
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 ina 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. ent. 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
Trigona 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 Ganella 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 MMelipona, 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 two communities
continued to form its comb in the same way, thus showing the
continuity that prevails in these cases as long as circumstances
permit. Miller 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 Trigona were imported
from Brazil to France, and kept there for some time in captivity
by M. Drory. Girard has published’ 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
queen-bee of JZ scutellaris seems to display more intelligence
than the corresponding sex of 4. mellifica. The mode of feeding
the larvae apparently differs from that of 4. mellifica, a provision
of pollen being first placed in the cell, then some honey ; when

1 Ann. Soc. ent. France (5), iv. 1874, 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
Trigona crassipes has the very peculiar habit of always locating
its brood-comb in the nest of a species of Zermes.

The honey-hee, Apis mellifica (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

Oy
o

ViOIsa vel EF

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-
surgitated 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 supphed, 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 ege 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-

eas

I SOCIALES—-HONEY-BEE 67

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 ege 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.’ 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 ege.

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 mellifica 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 down 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 Pérez, Act. Soc. Bordeaux, xxxiii. 1880, p. Ixv. ; and Cameron, 7’r. Soc.
Glasgow, n. s. ii. 1889, p. 194.

68 . HYMENOPTERA 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 be 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 Lombus.

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 httle 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 fotnd in southern Europe called A. ligustica; this is
certainly a variety of A. mellifica, and the same remark apphes
toa 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 Ray; the A. ligustica and A.
fusciata 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.
ligustica 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. mellifica is also

I SOCIALES—-HONEY-BEE 69

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.
adansonit, 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-
tinetive 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 Fic. 25.—Portions of hind-feet,
to be distinguished from the correspond- - Leah geen ae
ing sex of allied species of the genus. series on the left, outer

We are indebted to Horne for some = 3? a ae ee
particulars as to the habits of A. dorsata, first joint; ¢ second joint
an allied East Indian species. He informs cS aa
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

7O 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 Ramsay 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.

Nore 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.
INSKDOS Tos Be

Nore 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 Von Ihering.! 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. Nat. Hist. (6), xix. 1897, p. 186.

Norte to Vor. 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 toa
chain of embryos, instead of a single embryo.—@.R. Ac. Paris, cxxvi. 1898,
p-. 662, and translation in Ann. Nat. Hist. (7), 1. 1898, p. 28.

CHAPTER, - di

HYMENOPTERA ACULEATA CONTINUED—DIVISION II. DIPLOPTERA
OR WASPS——EUMENIDAK, SOLITARY TRUE WASPS—VESPIDAE,
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 tegula,; the basal seg-
ments of the hind body not
bearing nodes or scales; the
hind tarsi formed for simple
walking. The species either
solitary or social in their

Biiires ae ¢ ie Fic. 26.—Upper aspect of pronotum
Labits; some existing in three and mesonotum of a wasp, Eume-

forms, males, females, and es coarctata. a, Angle of prono-
tum ; 6, tegula; c, base of wing ;

workers. d, mesonotum.
Tuts division of Hymenoptera includes the true wasps, but not
the fossorial wasps. The name applied to it has been suggested
by the fact that the front wings become doubled in the long direc-
tion when at rest, so as to make them appear narrower than in
most other Aculeata (Fig. 27). This character is unimportant
in function so far as we know,’ 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 Leucospis
—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 has suggested that the folding is done to keep the delicate hind-margins
of the wings from being frayed.

V2 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. Eumenidae, Vespidae and Masaridae. We anticipate that
Eumenidae and Vespidae will ultimately be found to constitute
but one family.

Fam. i. Eumenidae—Solitary True Wasps.

Claws of the feet toothed or bifid; middle tibiae with only one
spur at tip. Social assemblages are not formed, and there is
no worker-caste, the duties of nest-construction, ete., being
performed solely by 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 flavopicta 9. 4-6 : F
Burma. The wines on the lett Gifference in habits there seems to be

in the position of repose, to no satisfactory way of distinguishing

show folding. ae
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 Hwmenes,
probably £. pomiformis. This Humenes 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 are
believed to be stung by the parent-wasp (as is the case in the

M1 DIPLOPTERA—WASPS—EU MENIDAE 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 Humenes or the deli-
eate 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 1s 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 Fic. 28.—Nidification of solitary wasps:
the food grows less by consumption, aoe Bag as aR = ae
the little larva is still enabled to arbustorum. a, The suspended egg
reach it by the fact that the egg- aie reer s ae aoe
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 Hwmenes construct vase-
like nests; 4. wnguiculata, 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, L. coarctata, L. of this genus occurs in Britain. The clay

7A HYMENOPTERA CHAP.

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 <
partition, a sort of dust-heap in which
it accumulates the various débris re-
sulting from the consumption of its
stores.
Eumenes conica, according to Horne,
constructs in Hindostan clay-nests with
very delicate walls. This species pro-
Fic. 29.—Nest of Humenes _- - : :
‘ouretate: A. the nest Visions its nest with ten or twelve green
attached to wood: B, de- caterpillars; on one occasion this ob-
tached, showing the larva. 5
i. thelarva: 6, the parti. SChWel a uOOkee iron s@ne cell eight green
nes, the cell. (After caterpillars and one black. It is much
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 Humenes were reared.

Odynerus, 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 Hwmenes, 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 Hoplopus 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 Réaumur, of another by
Dufour; and recently Fabre has added to the accounts of these
naturalists some important information drawn from his own

1L WASPS—-EU MENIDAE IS

observations on O. reniformis. This Insect provisions its celi
with small caterpillars to the number of twenty or upwards (Fic.
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,
Réaumur, 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

Fic. 30.—Odynerus antilope 2. 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.
Kabre 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 HYMENOPTERA 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, O. (Pterocheilus) spinipes
have been observed by Verhoeff; the facts as regards the con-
struction and provisioning of the cell are almost the same as in
O. 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, Argyromoeba sinuata, takes advantage of the habit of
the Odynerus of leaving its nest open during the process of pro-
visioning, and deposits also an egg in the nest; the Odynerus
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
Argyromoeba hatches, and the maggot produced feeds on the
caterpillars the wasp intended for its own offspring. Verhoeff
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. Rt. C. L. Perkins once observed several individuals of our
British O. 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 Odynerus.
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 lke the pot of an Humenes, 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 oh

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.

Horne has recorded that the East Indian 0. punctum is fond
of availing itself of holes in door-posts where large screws have
been; 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 Rhygehivm 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 J.
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. Horne 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, R. nitidulum. This latter Insect, however, does
not nidificate in the stems of plants, but constructs clay cells
similar to those of Huwmenes, and fixes them firmly to wood.
Rhygchium brunneum is said by Sir Richard Owen to obliterate
hieroglyphic inscriptions in Egypt by its habit of building mud

78 ’ WWYMENOPTERA CHAP.

nests amongst them. An individual of this wasp was found
by Dr. Birch when unrolling a mummy—‘There being every
reason to believe that the Insect had remained in the position in
which it was found ever since the last rites were paid to the
ancient Egyptian.”

Fam. 2. Vespidae—Social Wasps.

Claws of the feet simple, neither toothed nor bifid, middle tibiae
with two spurs at the tip. Insects living in societies, form-
ing a common dwelling 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 cells, 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 1s
desirable; de Saussure, the most important authority, bases 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 1s
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 woody fibre—amalgamated by means of
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 Vespidwe than they do in
that of the bees and fossorial Hymenoptera.

In most of the nests of Vespide 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 cells 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
imago taking place in the cells after they have been entirely

II SOCIAL WASPS—VESPIDAE 7a

closed. The food supplied is of a varied nature according to
the species, being either animal or vegetable, or both.

Although the nests of the social wasps are very elaborate con-
structions, yet they serve the purposes of the Insects for only a
single season. This is certainly the case in our own country.
Here each nest is commenced by a single female or queen; she
at first performs unaided all the duties for the inauguration of

oy

r a Fie. 31.—Section of the
2 me Wy N subterranean nest of the
iN \ common wasp, Vespa
germanica, in position.
(After Janet.) a, One
of the chambers of an
ant’s nest, Lasius flavus,
placed above the wasps’
nest; 6, 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 ; 7,the chief suspen-
sory pillar of the second
layer of comb ; g, lateral
galleries ; h, one of the
secondary pillars of
suspension between two
layers of comb; 7, the
layers of wasp-paper
forming the envelope of
nest; 7, vacant space
onl = round the nest ; /, flints
Z Ly ~ : vl {_A Hi; pa ee to tee
7) RAE: tZ during the work of ex-
3 eo a LL. cavation ; 2, numerous
larvae of a fly, Pegomyia
inanis (2 ) placed vertically in ground beneath the nest ; m1 to m’*, the layers of comb,
in m* the cells are indicated , in m® (above the main figure) the arrangement of the
three cells forming the natimeenanen of the new layer of comb, m’, is shown ; 7,
gallery of access from surface ; 0, burrow of a mole; p, interval of 90 mm. between top
of nest and surface ; ¢, height of the nest, 163 mm.

the colony; she lays the foundation of the cells, deposits the
eggs in them, feeds the young, and thus rears a brood of workers
that at once assist her, and for the future relieve her of a con-
siderable portion of her former occupations; the nest is by them
added to and increased, till the cold weather of the autumn is at
hand; at this time many males and females are produced; the
cold weather either destroys the inhabitants of the nést, or re-
duces their vitality so that it is impossible for them to pursue
successfully the avocations necessary for their subsistence, and

80 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. We 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 * 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-maché, 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 Zool. Anz. xix. 1896, p. 449. See also note, antea, p. 70.

II VESPIDAE—WASPS’ NESTS 81

which is probably chiefly glandular products. Our British
Vespidae 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. rufa form
subterranean nests, while V. arborea, V. sylvestris and V. norvegica
suspend their habitations from the branches of trees, bushes, or
strong annual plants. Vespa erabro, 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 occastonally make subter-

Ny

PEP ON)

ranean nests. It is ascertained that

V. austriaca, the eighth species, is an

inquiline. a
De Saussure,’ the monographer of the 4

social wasps, classifies them according to Z

the architecture of their nests. He estab- 4 ;

lishes three groups: (1) Stelocyttares, in 4,

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
Fig, 32.

: : Nest of (7) Polybia
of a tree, round, or on, which the envelope ~ “5,” The eae nore

is placed (Fig. 32); (38) Phragmocyttares, cut. open; 9, entrance.

; 5 After de Saussure. )
in which the layers of 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. 53).
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 Monographie des quépes sociales, Geneva, 1853-1858, pp. cc. and 356, plates
1.-XXXV11.

VOL. VI qi

82 HYMENOPTERA 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 Jschnogaster (Fig. 54) make layers of comb, con-
nected by a pedicel, but without any envelope; these Insects
form a section of Stelocyttares called Gymnodomes.

Most of the nests of the Poecilocyttares have only a single
layer of comb. 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 Polybia (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 Phragmocyttares, 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-maché 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.

I VESPIDAE—-WASPS’ NESTS 83

A very remarkable wasp’s nest 1s preserved in the British
Museum of Natural History; it is considered to be the work of
Montezimia dimidiata 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 Fic. 33.—Section of nest of
skin, so that the nest looks somewhat eee a AN Can oa

South America. 0, 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 Polybia—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 is 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 JV. 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. crabro. 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
egos soon hatch and produce larvae that grow rapidly, the labours
of the queen-wasp are chiefly directed to feeding the young. At
first she supphes 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 innutritious 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

ioe)
UL

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 upper 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 two 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,’ 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

' 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 HYMENOPTERA CHAP.

destroyed and reformed, as the combs they contain increase in
size. Marchal’s observations! relate chiefly to the production of
the sexes and worker-forms, in the subterranean species, Vespa
germanica and V. vulgaris. 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 parthenogenetically
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.
Polistes 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.

Next to Polistes, Vespa is the most numerous in species,
about 150 being known, and it is to this genus that all our
British social wasps belong. No 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. When 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 CR. Ac. Paris, cxvii. 1893, p. 584; op. cit. exxi. 1895; p. 781 5; Arch. Zool.
exper. (3) iv. 1896, pp. 1-100.

11 VESPIDAE—SOCIAL WASPS 87

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
uritated them. The East Indian V. velutina is specially fierce
when aroused, and is said by Horne to have followed a party

Fic. 34.—Ischnogaster mellyi. Java. A, Female imago (the line at the side shows its
length) ; B, nest, C, maxilla ; D, labium ; 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 unless
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
observed. It is said that wasps will rear the progeny of a neigh-
bour in circumstances where this assistance is necessary. Hess
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 1s 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 Jschnogaster (Fig. 54)
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-
eress of knowledge the present divi-
sion between social and_ solitary
wasps will prove ‘untenable.

Remains of Insects referred to
Fic. 35.-—Masaris vespiformis. A, male; the gener: Polistes and Vespa

B, female. Egypt. (Arter Schaum.) aaa F
have been found in tertiary strata
in various parts of Kurope and in North America.

Fam. 3. Masaridae.

Anterior wing with two complete sub-marginal cells. Antennae
usually incrassate or clubbed at the extremity. Claws dis-
tinctly or obsoletely dentate.

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

II MASARIDAE 89

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,
Quartenia and Coelonites are also members of the Mediterranean
fauna, while Pavagia is Australian, and 7’7rimeria South American.
Several species of the genus J/asaris inhabit North America, and
Cresson has recently described another
Masarid genus from the same country,
under the name of Huparagia.

The little that is known of their
natural history is almost limited to an
account given by Giraud of the habits of
Ceramius lusitanicus, of which species he
found a colony near Briancon. 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 1s 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 Fic. 36.—Cells constructed

j 3 é : by Coelonites abbreviatus.
of the larva is believed to be rapid. (Afiee André}

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 supphed by Ceramius lusitanicus to its young.

CHAPTER i

HYMENOPTERA ACULEATA CONTINUED—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, though very diverse
in form, does not bear prominences on the upper aspect of the
basal segments ; front wing without longitudinal fold along
the middle ; hairs of body not plumose. Only two forms
(male and female) of each species.

Fossor1AL Hymenoptera are distinguished from other Aculeates
at present only by negative characters, 7c. 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, ete.; (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. III FOSSORIAL SOLITARY WASPS QI

forms, they are true individualists, and their lives and instincts
offer many subjects for reflection. Unlike the social Insects they
ean 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 earler: 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

Fic. 37.—Sceliphron nigripes 9 (Sub-Fam. Sphegides). Amazons. x 3.

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-
quaintanee 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.

Q2 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 Mutillides 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. 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. Food or Occurrence.
Fam. Scoliidae.
Sub-Fam. Mutillides _. . As parasites on Hymenoptera Aculeata.
mM Thynnides . . (2) Parasites on Lepidopterous pupae.
5 Scoliides . . . Larvae of Coleoptera [(?) spiders in the case of
Elis 4-notata].
= Rhopalosomides. Unknown.
2 Sapygides . . The provisions stored by bees. Caterpillars

(teste Smith).
sae { Spiders. Rarely Orthoptera (Gryllidae and
Heo ge epee | Blattidae, teste Bingham) or Coleoptera.
Orthoptera (especially Locustidae), larvae
Fam. Sphegidae. {

Sub-Fam. Sphegides

of Lepidoptera, Spiders [(?) same species
(Sceliphron madraspatanum and Sphex
coeruleus), both spiders and caterpillars].

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

Ill FOSSORIAL WASPS 93

Group of Fossores. Food or Oceurrenee.
Fam. Sphegidae.
Sub-Fam. Ampulicides, . Orthoptera (Blattidae only).
re Larrides . . . Orthoptera of various divisions. Aculeate

Hymenoptera, in the case of Palarus.
[Spiders stolen from nests of Pelopaeus by
Larrada. |

Trypoxylonides. Spiders, caterpillars, Aphidae.

Astatides. . . Astata boops uses Pentatomid bugs, cockroaches,
and even Aculeate Hymenoptera (Oxybelus,
teste Smith).

$5 Bembecides . . Diptera and Cicada.

. Nyssonides . . Diptera, Homoptera (Gorytes mystaceus takes
Aphrophora out of its ‘‘ cuckoo-spit ”).

3 Philanthides. . Aculeate Hymenoptera (Philanthus). Hard

beetles, viz. Curculionidae, Buprestidae,
Chrysomelidae (Cerceris).

Mimesides . . Small Homoptera, even Aphidae. Diptera
(Tipulidae) in Hawaii.
A Crabronides . . 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.

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.

O4 HYMENOPTERA CHAP.

Fam. 1. Scoliidae.

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. We adopt five sub-families.
Sub-Fam. 1. Mutillides.—T7he sides of the pronotum reach the

tequlae: the female is destitute of wings and veelli,
frequently having the parts of the 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 two apical spurs. Front
wing with two or three sub-marginal cells. 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-hke
spaces. They have the faculty of stridulating, the position and
nature of the organ for the purpose being the same as 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 J europaea lives in the nests of bees of the
genus Lombus, and Hoffer has since made some observations on the
natural history of the same species in South East Europe, where
this JZfutilla is found in the nests of ten or eleven species of
Bombus, being most abundant in those of Bb. agrorum, and B.
variabilis ; occasionally more individuals of Jutila than of bees
may be found in a nest. He supposes that the egg of the
Mutilla is placed in the young larva of the Bombus, 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

ich.

11 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 Bombus 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

Fic. 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
ditference 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 MJutilla are
described as being apterous in the male as well as in the
female 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

96 HYMENOPTERA CHAP.

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

Sub-Fam. 2. Thynnides.—JMuales and females very different in
form; the male winged, the front wing with three, or only
two, sub-marginal cells ; the female wingless and with the
thorax divided into three sub-equal parts.

The Thynnides are by some entomologists not separated from
the Mutillides; but
the distinction in
the structure of
the thorax of the
females~ is very
striking. In the
Thynnides the
nervuration of the
wing appears
always to extend
to the outer
margin, and in the

Mutillides not to

Fic. 39.— Methoca ichneumonides. A, Male; B, female. do so. This family
Britain. ‘

is represented in
Britain by a single very rare Insect, Methoca ichnewmonides : 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 beheve the two sexes
to have any connection; but the fact that they are so con-
nected has been demonstrated beyond doubt. In very few

Il SCOLIIDAE—_THY N NIDES——-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 with 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 Fic. 40.—Scolia haemorohoidalis 9. Europe.
angles are always co-
adapted with the points of insertion of the front wings. The
nervuration 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
Seolides found in France, and has informed us that their means
of subsistence consists of larvae of the larger Lamellicorn beetles,
Cetonia, Oryctes, Anoxia, and Euchlora; 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 débris 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

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 Cetonza 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 Cefonia 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 régle.
If the head of the Scolia larva be taken out from the victim and
apphed 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
Jetonia larva dies, its death involving also that of the destroyer.
It is necessary, too, that the victim should be paralysed, for if an
intact Cefonia 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 Scolia 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
cue, and the Sco/ia 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 ;

!

11] SCOLIIDAE

SCOLILDES——SAPYGIDES 99

in order to reach this the Seo/ia@ 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 belheved 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.

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. Sapygides.— Closely allied to the Scoliides, but pos-
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,
Peckham on Zhe 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 ; CHAP.

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

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

Fic. 41.—Sapyga 5-punctata @,
Britain.

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

hypermetamorphosis (7.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. We have two
species of Sapyga in
Britain; they differ in
colour, and the sexes of
S. 5-punctata also differ
in this respect; the
abdomen, spotted with
white in both sexes, is
in the female variegate
with red. Smith found
our British Sapyga 5-
punctata carrying cater-
pillars.

Sub-Fam. 5. Rhopalo-
somides. — Antennae
elongate, spinigerous ;

ocelli very prominent ;

y uy

fi

a/

4 .

Fic. 42.—Rhopalosoma poeyi. A, female imago ;
B, front of head. Cuba. (After Westwood.)

tarsi of peculiar structure, their claws bifid.

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

1 P. ent. Soe. Washington, iii. 1896, p. 303.

111 FOSSORES VOW

an extremely rare American Insect that had previously been placed
by Cresson among parasitic Hymenoptera. Westwood classed
Rhopalosoma among Diploptera, saying of it “ animal quoad
affinitates 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 in form,
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

OZ, HYMENOPTERA CHAP.

enraged at the audacity of its persecutor. The Calieurgus 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
ageressor 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 iatten-
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,

4 ; \ which now pounces on its
4 \

i iy 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 may 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 which 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
Pompilid, yet if the two be confined together in a vase, both are

Fic. 43.—Calicurgus hyalinatus 2. Britain.

EEE

111 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
polistoides. 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 flew 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 Pepsis, a genus peculiar to America, whence
upwards of 200 species are already known. Some of them
attain 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. Zeitschr. xxxix. 1894.

\
i
\

|

eee

WW POMPILIDAE 105

due to scales. P. formosus, Say, 1s 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 flles “in cireles 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, MJygale 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 Pepsis, 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 Pepsis. 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. Parapompilus 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 use for flight.
P. gravesii is an inhabitant of Chili.

Agenia carbonaria and A, hyalipennis 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 4. hyalipennis 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 Agenza larva is of peculiar
shape, the head not being inflexed, 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. Pérez 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 ege on 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 hyalinatus, 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
habits of some of the Ichnemonidae, though the perfect Insect
(Fig. 145) 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.

Ill FOSSORES—SPHEGIDAE——-SPHEGIDES NO

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—Prioenemis 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
Jar back as the wing-insertion, they are placed below it and
separated by 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’ 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 Sphea.
1 «Die Gattungen der Sphegiden,” Ann. Hofmus. Wien. 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, ete. of Sphex chrysis. f points
to a division of the mesonotum, not of the metanotum, as there stated.

108 HYMENOPTERA 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. maaillosus.
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 Sphex 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 Sphex, 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
Sphex’s 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-

III SPHEGIDAE——SPHEGIDES 109

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 Sphex, 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 Sphez 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
ericket 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 12 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

110 HYMENOPTERA 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 quite 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 coeruleus 1s 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 Pelopaeus (Sceliphron)
coeruleus, S. coeruleus is no doubt the same as S. (Chlorion)
lobatus, which Rothney 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
Sphexz 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
Kothney repeated the experiment; in due course the same series

190% SPHEGIDAE——SPHEGIDES joa dt

of 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. hirsuta 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 has 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. 4. 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. afinis; 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 mother-Ammophila herself profits by appropriating an exuda-
tion from the victim.

Some species of Sphegides have the curious habit of choosing

Li2 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 flames,
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 Pelopaews 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. Two or three
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. Pelopaeus (Sceliphron)
is a genus including many species;' several of them are known

' Pelopaeus disappears from the new catalogue of Hymenoptera as the name of a
valid genus ; its species being assigned to Sceliphron and 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.

A
ee

III SPHEGIDAE—SPHEGIDES—-PELOPAEUS Lis

to be specially attached to the habitations of human beings.
Roth has given an account of the habits of P. (Sceliphron)
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, hable to the charge of
being very capricious. Roth 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 Leuveospid 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. (Sceliphron) madraspatanus is common in the north-west
provinces of Hindostan, and is called the “ mud-dauber” by the
European residents. According to Horne 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 I

IIl4 HYMENOPTERA: -~ CHAP.

spiders. The species is said to be protected by a pecuhar odour
as well as by its sting; it 1s also stated that it disguises its
edifice when completed by making it look hke 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. (Sceliphron) bilineatus, formerly thought to be a variety
ot P. madraspatanus, builds its nests in hedges and trees.

Sub-Fam. 2. Ampulicides.— Prothorax long and narrow, forming
a neck in front; clypeus 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-
ast Galt 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

Fig. 44.—Ampulex compressa. Male. four times its own size, suc-

East India. 0 : :
ceeds, after a struggle, in sting-
ing it; the cockroach instantly becomes quiet and submissive, and
suffers itself to be led away and placed in confinement in some

III SPHEGIDAE—-AMPULICIDES eS

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 Rothney in reference to the habits of A. (2hi-
nopsis) ruficornis 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 were partially known to Réaumur, 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 Ampulex 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 Dolichurus 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

116 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. haemorrhous lives at the expense
of Loboptera decipiens, 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 body not pedicellate, or with
only a short pedicel ; one spur on the middle tibia ; labrum
inconspicuous. Marginal cell of the front wings appendi-
culate or mandibles excised externally, or both.

This group is by some writers called Tachytides instead of
Larrides, as owing to a change of nomenclature Zachytes 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 “ Miscophus group” of Kohl.

The species of the genus Zachytes seem to have habits very
similar to those of the genus Sphex; they form shafts in the
earth and provision them with Orthoptera; lke the Sphex 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 was
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.

‘When a second cell is more or less perfectly marked out, the cell with which
it is connected is said to be appendiculate. The nervures frequently extend beyond
the complete cells towards the outer margin, forming ‘‘ incomplete” cells ; only
complete cells are counted, except when ‘incomplete ” is mentioned.

111 SPHEGIDAE— LARRIDES ez

One species of Zachytes 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,
‘apable of cutting in two by
a single act the body of an
aggressor like the TZachytes ;
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 Fic. 49.—Tachytes pectinipes 2.
Tachytes at last pounces down chee
and instantaneously stings the Mantis in the nerve centre
between the formidable arms, which at once are reduced to in-
capacity ; subsequently the Zachytes 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 Vachysphex panzeri selects grasshoppers of the family
Acridiidae. Larra pompiliformis ( = Tachytes niger, Fabre) some-
times associates itself with Sphex flavipennis (2? S. mazxillosus,
according to Kohl), forming its burrow amidst the works of a
colony of that species, and making use, like the Spe, of crickets
for provender. This led Fabre to believe that the Zarra stole
its prey from the Sphez, but he has since withdrawn this indict-
ment, and declares that the Zarra 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 (Sceliphron) 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 consequences of the intruder’s operations ;

Toes . 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 be 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 1s 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 Zarrada 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 Pe/opaeus 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 modesta,
in Ternate, it seems probable that its habits may prove to be
similar to those of ZL. australis, for it frequents the nests of
Pelopaeus after they have been completed.

Sub-Fam. 4. Trypoxylonides—JDiffer from Larrides 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 distinet discoidal cell, a second
of each being indicated faintly.)

The nervuration of Zrypoxylon 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 Zrypoxylon are apparently rather
fond of human propinquity, and build clay- or mud-nests in or
near houses. 7. albitarse has this habit, and is well known in
Southern Brazil under the name of “ Marimbouda da casa”;

HI SPHEGIDAE——-ASTATIDES——-BEMBECIDE I19

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 7. 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.—Lyes 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. boops—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 LPentatoma, 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 bugs and _ cockroaches
being their disagreeable char-
acter. According to Smith, Ozybelus, 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 <Asfata, with which the
North American Diploplectron (with one species) is doubtfully
associated.

Fic. 46.—A stata hoops, mate. Britain.

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 emarginate externally ; hind body stout, not pedicellate.

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

120 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
Bembex flies direct to it with-
out any hesitation. How neces-
sary ib is to these Insects to
possess this faculty of finding
their nests will be understood
when we recall that the Bembex
does not provision ifs 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 flies away; to proceed possibly with the formation

Fic. 47.—Bembez rostrata 8. Europe.

III SPHEGIDAE—BEMBECIDES 12%

of other burrows.’ 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 Bembex 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 be 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 ember 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.

Me 2 HYMENOPTERA : 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 Bembex 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 embex 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 lght sand. Fabre informs us that the Bembex
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 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 Motuea 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.

ERS

Ill SPHEGIDAE—NYSSONIDES 1238

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. Riley 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.—JLabrum short; mandibles entire on
the outer edge; hind body usually not pedicellate; wing
with the marginal cell not appendiculate.

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 Me/linus, in which the abdomen is pedi-
cellate (Fig. 48), while others treat that genus as forming a
distinct sub-family, Mellinides. Kohl leaves Me//inus unclassified.
Gerstaecker has called attention to the fact that many of the
Insects in this group have the trochanters of the hindvand middle
legs divided: the division is, as a rule, not so complete as it
usually isin Hymenoptera Parasitica ; but it is even more marked
in some of these Nyssonides than it is in certain of the parasitic
groups.

Mellinus 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
not> deficient™ am, “deyicess ) i
noticed once or twice, what I took to be 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 J/ellinus started into life, and
pounced upon its victim.”

Lucas states that in the north of France JMellinus sabulosus
provisions its nest with Diptera, which it searches for on the
flowers of Umbelliferae, and then carries to its nest. This is ¢
burrow in the earth, and when it is reached the Hymenopteron
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.

Fic. 48.—WMellinus arvensis 2. Britain.

Sub-Fam. 8. Philanthides.—Zabrum small ; anterior wings with
three complete submarginal cells; hind body constricted at
the base but not so as to form a slender pedicel.

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

——————————
————————

HI SPHEGIDAE——PHILANTHIDES L2

UL

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 Cercevis 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 Dutour
that one kind of Cerceris

never in its selection goes ~\ “a
out of the limits of a Se

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,
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. /abiata 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 bupresticida 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.

Fic. 49.—Philanthus triangulum &. Britain.

126 ITYMENOPTERA CHAP.

It has long been known that some species of Cerceris prey on
bees of the genus Halictus, 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
tuberculata, 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 had 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

|
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1II SPHEGIDAR——MIMESIDES M47,

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 Phil-
anthus apparently kills the bee at once, while in the case of the
Cerceris, the Halictus 1t 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 in this case is really dead, not merely in a state of

anesthesia, the Philanthus 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, the 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. atratus
form their nests in hollow stems, and the former provisions its
nest with Homopterous Insects of the family Psyllidae. Little

128 HYMENOPTERA CHAP.

information exists as to their habits; but Verhoeff states that
the species of Psen—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 is
also. the case with Stigmus
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 Cynips, and also of
a peculiar swelling formed by a fly—Lipara lucens—on the
common reed, Arundo phragmites. This species also makes
use of Aphidae, and Verhoeff states that it has only an imperfect
instinct as to the amount of food it stores.

Fic. 50.—Mimesa bicolor 2.
Britain.

Fic. 51.—Pemphredon lugubris 2. Britain.

Sub-Fam. 10. Crabronides —Pronotum short, front wing with
one complete submarginal and two discoidal cells: hind body

IIL SPHEGIDAE—CRABRONIDES I29

variable in form, pedicellate in some abnormal forms, but more
usually not stalked.

The Crabronides (Vespa erabro, 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 chrysostomus have been briefly described by Verhoeff:
the food stored consists of Diptera, usually of the family Syr-

VOL. VI K

Fic. 52.—Crabro cephalotes 2. Britain.

130 HYMENOPTERA CHAP. III

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 Zyphlocyba, 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 wasp 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 Hertonius 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 agaist the feeble
larva. Ferton thinks there are three generations of this species
in a single year.

Notr.—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 Bembex spinolae
with a view of ascertaining whether the female provisions two nests simul-
taneously ; as the result they think this improbable. If the female Bembecid
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.

CEeGr TER Ly

HYMENOPTERA ACULEATA CONTINUED——DIVISION IV. FORMICIDAE
OR ANTS

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 with 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 have no
wings, but the males and females
are 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
and tended by the workers or by
the female.

Fic. 53.—Abdomens of ants.
A, Of Camponotus rubripes
In ants the distinction between the — (Formicides); B, of Hetu-
: : tomma auratuin (Ponerides);

three great regions of the body is very C, of Aphaenogaster bar-

marked. The abdomen is connected %¢7¢(Myrmicides). a, Pro-
: ; ; podeum ; 2, first abdominal
with the propodeum ina peculiar manner, —_ segment forming a scale or
one or two segments being detached node: & second ; d, third
c 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

MWB) 2 HYMENOPTERA CHAP.

greatly in the family: and the Amblyoponides do not differ in
a marked manner from the Scoliidae in fossorial 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 lp
and maxilla assum-
ing an ascending
vertical direction,
while the upper
lp hangs down
and overlaps the
lower lip, being closely applied to it; so that 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 lps 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 winged or wingless. The sutures between the dorsal

Fia. 54.—Front of head of Dinoponera grandis.
A, Mouth closed ; B, open.

IV FORMICIDAE—ANTS 13

Oo

(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 Fic. 55.—Oecodoma cephalotes. South
peduncle, or pedicel, formed by the = America. A, Worker major; B,
5 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

Fic. 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, 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,b); at other times there are two

134 HYMENOPTERA CHAP.

small segments (Fig. 53, B, C, 6, ©) 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
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-
Pa ae proportionately elongate, SO that in use

on front leg of an ant, Dino- 1t acts in many species as if 1b were a

Lae, grandis (tip of tibia, portion of the tibia, the other four joints

searing the comb-like spur, ‘ ; 0

and the base of the first joint forming the functional foot. The front

of the tarsus; cf fig. 75). tibiae are furnished with a beautiful

A, Inner, B, outer ee

combing apparatus (Fig. 57).

Features of Ant- life —In order’ that the en eh 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 eges
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.t After a brief period of

' 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—impertect 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
anunals, 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
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 im 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 137

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 most frequently adopted Fic. 58.—Portion of combined nest of For-
when the soil in which the ‘™cafusca and Solenopsis fugax. (After
i s Forel.) x2. f, 7’, Chambers of Mormica,
nests are placed 1s sandy ; 1t recognisable by the coarser shading ;

is probable that the earth is s, s’, chambers of the Solenopsis (with

finer shading) ; s’”, opening in one of the
t >’ ’ 5

in such eases fastened together chambers, the entrance to one of the
by means of a, cement pro- giles fhat comets the chanbers ofthe
duced by the salivary glands tions of the nest and the limits of the
of the ants, but this has not °@™mers-

been determined with certainty ; vaulted galleries or tunnels of
this kind are constructed by many species of ants in order to
enable them to approach desired objects.

In South America Camponotus rufipes 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 Formica fusca (Fig. 58), in such a manner that its chambers
cannot, on account of the small size of the orifices, be entered by
the much larger Mormica. Hence the robber obtains an easy
living at the expense of the larger species. The Sauba or Sauva
ants ef South America (the genus Atta of some, Oecodoma ot
other authors) appear to be most proficient in the art of sub-

138 HYMENOPTERA CHAD.

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. Camponotus ligniperdus in Europe, and C. pennsyl-
vanicus 11 North America, work in this way.

Our British Lasivs 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

ee of delicate paper, or that spin

oY Sosa ae teen mon- together by means of silk the

leaves of trees. One eastern

species—Polyrhachis spinigera—tfabricates a gauze-lke web of

silk, with which it lines a subterranean chamber after the
manner of a trap-door spider.

Some species of ants appear to find both food and shelter

h \: /\ ; ‘\
/ \\ @ ! \ \
NW ! /)
A \ 4

|
|

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. 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 1s 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
pecuhar 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 httle inquired into. The
difference between the thoracic structure in the case of the winged
and wingless females of certain species (Fig. 55, and in vol. v.
fig. 339) 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 ;' 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
and 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 alienus 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 CEpyarns,
a worker).?

' 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 Forel, Verh. Ges. deutsch. Naturf. \xvi. 1894, 2, pp. 142-147 ; and Emery

Biol. Centralbl. xiv. 1894, 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.

i's 3 a © 5
2 zt 5 ws 5 Ac a
& Bs B ses 5 5%
Eo FS) BS Bos ‘z Ps
Name of Ants. pia |g a = <= ihe S pies,
aa(Sos/a2| Ss EET i a aS
2 | | st Ss a g orcs
o) = he) aa eo) 7) = 5
EEE | | = —
|
|
Myrmica, Polyrhachis, \ |
eae Seal bas ae | ii
Camponotus, ‘Atta, \ nt + rs an
Pheidologeton, ete. .f |
Pheidole, subg. Colo- |
bopsis 4 7 = = 4
Eeciton hamatwm, |
E. quadriglume, -| + + =e + 4
E. foreli, ete. . al
Cryptocerus disco- \ | |
cephalus, ete. hee “] a 4 =
Str ongylognathus + + | +
Carebara and Nae
opsis (except + ae +
geminata) |
Solenopsis g geminata . : > + | + +
| ~
}
Formica rufa + + excep-
tionally
Ponera punctatissina 4¢ || oe |) se | st
Ponera ergatandria 2) +] 4+ +
Cardiocondyla emeryti _ + ~ tf
C. wroughtonii and \
+
C. stambulofi . af etd |
Fornicoxenus —niti-\
; -
dulus. . | lie
Tomognathus =F | ae ree ||
| +
Odontomachus hae-\ + rs if prs: i
matodes . if [sonality
Polyergus . : ele sk se | ae
Dorylus, Anomma, \ | |
. | | +
Eciton part. . =| oi ist
Aenictus : 2
Leptogenys, Dia-\ ii
camma . : a Aire | 9 “B
} ( is
Myrmecocystus | mel-\| ae J and |
liger, M. mexicanus J ‘pots |
| pots
+
Ponera eduardi . : + + | : If eyes | eyes
|| large I 3 obsolete
Anergates . ; wey Hee fe
| |
: J

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

142 iy HYMENOPTERA CHAP.

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 Afia)
—one of the species in which polymorphism is greatest—says '
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 he doubts it being
“the character of the food.” There is, however, a considerable
body 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, 1s
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 Lomechusa 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 1t 1s
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. 125.

IV ANTS 143

determined by those “ Anlagen” 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
ean it be, he asked, that one egg may produce either a worker,
nurse, soldier or female ant? To this Wasmann (who continued
the discussion) rephed 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.! 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.” Emery accounts® 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 Biol. Centralbl. xv. 1895, p. 640.

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

* Biol. Centralbl. 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
Amblyopone, to certain Mutillides (e.g. Apterogyna) and to the
Thynnides, two divisions of Scoliidae. Emery considers Dory-
lides 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-familes 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 familes in one
direction. The Myrmicides are also highly differentiated, but
are not allied to the Camponotides and Dolichoderides.'

Sub-Fam. 1. Camponotides.— Hind body furnished with but 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 pecuhar 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. Soc. ent. Belgique
xxXxvii. 1893, p. 161; the very valuable paper by Emery, in Zool. Jahrb. Syst.
viii. 1896, p. 760.

ee

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. 7

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, Kormica rufa, 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 1s
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 truequeen-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
paste, 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
Lasius alienus, ammaccompanied by any workers or other assistants.
and 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.” 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. Soct. ent. France, 1893, p. 467.
2 Ann. Soc. ent. France, 1893, Bull. p. eclxiv.

Iv ANTS—CAMPONOTIDES 127

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
ereedily 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 was 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 we shall subsequently see, are formed with
ants by numerous other Insects.

Occophylla smaragdina, a common ant in Eastern Asia, forms
shelters on the leaves of trees by curling the edges of leaves
and joining them together. In doing this it makes use of an
expedient that would not be beheved 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
for it to form a cocoon
with, and the ants utilise
the larvae to effect their Fic. 60.—Oe¢ecophylla smaragdina. Worker
purpose, Several of them using a larva for spinning.
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

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. |

Formica rufa, the Red-ant, Wood-ant, or Hiull-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 1s 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 Formicozenus

IV CAMPONOTIDES—SLAVE-MAKING ANTS 149

nitidulus lives only with #. rufa. Amongst the most peculiar
of its dependants we may mention large beetles of the genera
Cetonia and Clythra, 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 1700 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 /# 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
Britain. 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

150 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 reheve them of
much of their domestic duties. The species that M sanguinea
utilises in this way in England are /. fusca, F. cunicularia, and
possibly Lasius flavus. Huber and Forel have given graphic
accounts of the expeditions of this soldier-ant. In the mixed
colonies of FY 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.

Polyerqus rufescens, 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 1ts existence, and will starve, it is said, in the
midst of food unless its servitors are there to feed it. Wasmann,
however, states that Polyergus does 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

Fic. 61.—Head of Polyergus act of retreating or making flank move-
rufescens. (After André. ) is

ments when necessary. If a Polyerqus
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 deat

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 Eeiton; 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. rufescens 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 worker (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
Polyerqus for purely mechanical reasons.

M‘Cook states that the North American ant, Polyergus lucidus,
which some entomologists consider to be merely a variety of

i562 HYMENOPTERA CHAP.

the European species, makes slaves of Formica schaufussi, 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. When a migration takes place the servitors deport both
the males and females of P. ducidus. 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 1s 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 Myrmecocystus
hortideorum and IM. melliger, are mode-
rate-sized Insects of subterranean habits,
_the entrance to the nest of JZ hortideorum
being placed in a small raised mound.
The honey is the product of a small gall
found on oak leaves, and is obtained by
Fic. 62.—Myrmecocystus mexi- the worker-ants during nocturnal expedi-

aoe a once Hae ant, tions, from which they return much dis-

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 food
till a time when it is required for
the purposes of feeding the com-
munity. The distension is pro- Fis. Peis eee eR
duced entirely by the overcharging
of the honey-crop, the other contents of the abdomen being

WV ANTS—CAMPONOTIDES 153

forced by the distention to the posterior part of the body.
Lubbock has since described an Australian ant, JJelophorus
inflatus, having a similar pecuhar habit, but belonging to the
allied tribe Plagiolepisu. Quite recently a South African honey-
tub ant belonging to the distinct genus Plagiolepis (Ptriment
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 Lasiuvs are amongst the most abundant
of the ant-tribe in Britain. They are remarkable for their con-
structive powers. JL. 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.

L. 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 LZ. 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 Z. 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 confréres, 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 rufa 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 lying 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 Hormica 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 I

UL
un

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 influence 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
fohage, and some of
these contain only a
few individuals. The
leaf-nests (Fig. 64) of
certain species of Poly-
rhachis 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. rastella 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

Fic. 64.—Nest of Polyrhachis sp. (After Smith.)

' Forel, J. Bombay Soc. viii. 1893, p. 36.

156 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
Fic. 65.— Polyrhachis pandurus, and endings of the musical parties
worker. Singapore. : pees Ee)
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
Camponotjdes. Although the species of Camponotides have no
special organ for the production of sound in the position in which
one is found in Myrmicides and Ponerides, yet it is probable that
they are able to produce a sound by rubbing together other parts
of the abdomen.

IV ANTS I

wn
N

Sub-Fam. 2. Dolichoderides.—Hind body furnished with but
one constriction so that only a single scale or node is formed ;
Sting rudimentary ; 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-familes, 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 Yapi-
noma erraticum, a little ant of about the
size of Lasius niger, and somewhat similar
in appearance, but very different in its
habits. 7. 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 Fic. 66.—Tapinoma errati-
P % : : : : cum, worker. Britain.
in 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 TZapi-
noma has no power of ejecting the fluid to a distance, but 1s
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

158 HYMENOPTERA CHAP.

Insect is subterranean in its habits, and is said to change its
abode very frequently. 7. 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
burrows 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.
No 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 Miiller 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 1t 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.
This Insect is thought to protect the plant from the attacks of
leaf-cutting ants of the genus Atta.

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.’

Sub-Fam. 2. Myrmicides.—/Pedicel of abdomen formed of two
well-marked nodes (knot-like segments). Sting present (absent
in the Cryptocerini and Attini). (It should be noted that the

1 See von Ihering, Berlin. ent. Zeitschr. xxxix. 1894, p. 364; and Forel, Ann.
Soc. ent. Belgique, xl. 1896, p. 170.

IV ANTS—-MYRMICIDES 159
workers of the genera Eciton and Aenictus of the sub-family
Dorylides have, like the Myrmicides, two nodes in the pedicel.)

This sub-family consists of about 1000 species, and includes
a great variety of forms, but, as they are fh: si
most of them of small size, they are less
known than the Camponotides, and much A
less attention has been paid to their J
habits and intelligence. Forel, until re-
cently, adopted four groups: Myrmicini,
Attimi, Pseudomyrmini and Cryptocerini ;
but he is now disposed to increase this

number to eight.' They are distinguished ;
by differences in the clypeus, and in the I

form of the head; but it must be noted

that the characters by which the groups p67 pres, Toten ABR:
are defined are not in all cases fully riosus, large and small
applicable to the males) The Crypto- “°™e"s Bast India.
cerini are in external structure the most highly modified of
Hymenoptera, if not of all the tribes of Insecta.

i. The MyRMICINI proper are detined 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
Fic. 68.—Formicoxenus 10 these extraordinary life-histories.

eee male. (After It has long been known that the little

Formicoxenus nitidulus lives as a_ guest
in the nests of Formica rufa, the wood-ant; and another
similar ant, Stenamma westwoodi, which shares the same life,

1 Ann. Soe. ent. Belgique, xxxvii. 1893, p. 163.

160 HYMENOPTERA CHAP.

was declared by Nylander and Smith to be its male; it was
however shown some years ago by André that this is a mistake,
and that S. westwoodi is really the male of another ant that
had till then been called Asemorhoptrum lippulum. This correc-
tion left the workers and females of Formicoxenus nitidulus
destitute of a male, but Adlerz 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

Fic. 69.—Anergates atratulus. Europe. A. male, with found in this country.

part of hind leg broken off ; B, female, with wings: C, HER fees
female, after casting the wings and becoming a queen. The eouue Anei gates 1s
allied to Kormicoxenus,

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 Zetramorium caespitum, a little ant very like Myrmica, and not
uncommon in South-East England. The female Anergates is at

IV MYRMICIDES—AWNERGATES 161

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 Zetramoriuvm are found in company
with the ¢ and 9 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 Zetramorivm, 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 Zetramorium, 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 Zetramoriwm must also be
destroyed, for no young of the Zetramoriuvm 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 7'etramorium 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 Zomognathus attack the nest
of another little Myrmicid, Leptothorax acervorum, and succeed
by their own pertinacity and the fears of the Leptothorax in

\OD ES AYE M

162 HYMENOPTERA CHAP.

obtaining possession of it; the legitimate owners disappear, leav-
ing the Yomognathus in possession of their larvae and pupae ;
these complete their development only to find themselves the
slaves of ZYomognathus. 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 Zomognathus secure a supply by raids after the
fashion of the Amazon-ant, bringing back to their abode larvae
and pupae of Leptothorax to be developed as slaves. It was for-
merly supposed that the Zomognathus continued its species by
perpetual parthenogenesis of the workers, for neither males nor
females could be found. Adlerz’ has, however, now discovered the
sexual individuals. The male is an ordinary winged ant, and is
so lke that sex of the Leptothorax, 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 Zomognathus 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 ZYomognathus are so
similar to those of the Leptothoraz that Adlerz is quite unable to
distinguish the two.

Strongylognathus testaceus and S. huberi live in association with
Tetramorium caespitum, and are cared for by these latter ants; 1b
is notable that as in the case of the slave-making Polyergus
rufescens the mandibles of the Strongylognathus ave cylindrical
and pointed, and therefore unsuitable for industrial occupations.

1 Bih. Svenska Ak. xxi. 1896, Afd. iv. No. 4.

IV ANTS—MYRMICIDES 163

NS. testaceus is a weak little ant, and lives in small numbers in the
nests with 7. 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 7. 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 7. caespitum in order to carry off worker
pupae, just as Polyergus rufescens does. But the facts that SN.
testaceus 18 a weaker Insect than the Zetramoriwm, and that only
a few of its worker-caste are present in a community where there
are many workers of the Z’etramoriuwm, 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 hubert 1s a much stronger Insect than its
congener, S. festaceus, and Forel has witnessed its attack on
Tetramorium caespitum. Here the raid is made in a similar
manner to that of Polyerqus rufescens on Formica ; the Tetramo-
rium 1s 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. hubert, being similar in form to those of Polyergus
rufescens, are used in a similar manner.

Although 7” 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 Myrmica 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 JL. ruginodis and M. scabrinodis, which sometimes,
at the time of the appearance of the winged individuals, form vast
swarms.

The tiny Monomorium 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

104 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 1sS0-
lated chambers connected by
galleries ; some of the chambers
are used as store-houses or
eranaries, considerable quantities
of corn, grass, and other seeds
being placed in them. A.
structor and A. barbarus have
been observed to do this in
Southern Europe by Lespes,
Mogeridge, and others.

In the deserts about Algeria
and Tunis a harvesting ant,
Aphaenogaster (Messor) aren-
arius, 18 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
iis: ape holes, which are the orifices of

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

iene worker or soldier; D, small the holes are surrounded by

sie pellets of sand projecting some-
what above the general surface, and consequently making the
places conspicuous. The subterranean works occupy an area of
fifty 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 comes to the surface.”

Pogonomyrmex barbatus and other species have been observed
to do harvesting in North America. After the workers of P.
barbatus have taken the seeds into the nest they separate the
husks and carry them out, depositing them on a heap or kitchen-

1 Until recently this genus was generally known as Aéta, but this name is now
applied to the leaf-cutting ants, that were formerly called Oecodoia.
2 Forel, Bull. Soc. Vaudoise, xxx. pp. 29-30, 1894.

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, Pheidole—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 Aphaenogaster ; 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

Fic. 71.—Cremastogaster 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.

ul. The group ATTINI is distinguished by the presence of a
‘arina 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 Oecodoma)
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

166 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 be 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. fervens made an underground route at an
average depth of 18 inches, and at an occasional depth of 6
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 600 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

1V MYRMICIDES—LEAF-CUTTING ANTS 167

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—ramities.
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. Méller 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 confréres 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

168 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 J/yrmica.
Possibly the females may, after losing their wings, again enter
the large communities. Von Ihering states that the workers of
Atta lundi are fertile.
ii. The group PSEUDOMYRMINI includes the genera Psewdo-
myrma and Sima, 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 Pseudo-
myrma 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. Belt
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

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

a Fia. 72.—Sima rufo-nigra and
its associates. A, winged
female ; B, worker, of the
ant; ©, Rhinopsis rufi-
cornis, a fossorial wasp of
the sub-family A mpuli-
cides ; 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 Fis. 73.—Stridulating file
eoande: of Stina leviceps.

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
no 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 bushes,
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. atratus has
been observed to construct its nests
in the dead, suspended branches of
* woody climbers; a number of neatly
Fic. 74. — Oyrptocerus atratus, 4-17: s

a pee ienaeole | meicene eulled: holes: are alll tlabacan.bemsees
pressed first joint of the hind externally ; but, inside, the wood is
foot is shown at a and 6 in rene ¢ : .
event wasikions. freely perforated with intercommuni-
eating 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 Cryptocerus 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 Strumigenys and Epitritus,
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 with one
node at the base, and having also great capacity of movement
between the first and second segments, between which there is
usually a slight constriction. Sting well developed.

This sub-family includes numerous genera and about 400
species. The Ponerides have an elongate hind-body ; the second
segment behind the node is capable 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 Ey

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 shghtly depressed.

We follow Forel in including the Australian bull-dog ants—
Myrmecia—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 known it would appear that the three sexual
forms are not so differentiated as they are in most of the Cam-
ponotides and Myrmicides.

The species of the genus Leptogenys are believed by Emery
aud Forel to possess an
apterous female. Mr.
Perkins has observed that
the Hawanan ZL. falcigera
has workers with differ-
ent kinds of sting, but
no true female. Males of
this species are, however,
abundant. = Wroughton
has recently discovered
that one member of this
genus is of Termito-
phagous habits, but this is not the case with Z. falcigera. Dino-
ponera grandis (Fig. 75) is the largest of the Ponerides, its -

Fic. 75.—Dinoponera grandis, worker. Amazons.

172 HYMENOPTERA CHAP.

workers attaining an inch anda 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, hving 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, /.
punctatissima, 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. ergatandria.! The discovery of this form has led
him to express some doubt as to whether Ponera punctatissima
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 Cardiocondyla and a few other Myrmicides exhibit an
analogous peculiarity.

The genus Myrmecia is confined to the Australian continent
and 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 (Paraponera clavata, eg.) that differ but little
in this point from J/yrmecia, and, moreover, the pupae of Jyr-
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. London, 1893, pp. 365-467.

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 “ bull-
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 TREE Uae yiessT i rarirnice
high, and the colonies are immense in Australia. Female after
numbers, while the Insect is an inch or ‘“*St#2S 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 (Hucharis myr-
meciae) of brilliant colour and considerable size, so that we have
the curious fact of the hordes of this most formidably armed ant,
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 sufiicient observations been made as to
the mode in which the attack is made. Lowne says that JZ. gulosa
itself attacks large beetles of the genus Anoplognathus and buries
them ; and he also adds the very curious statement that JZ nigro-
cincta, When running, is able to take leaps of a foot in length.

The Odontomachi were formerly considered a distinct sub-

—

family, distinguished by the peculiar mandibles (Fig. 77

eek 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
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.

Fic. 77.—<Anochetus ghiliani,
worker. Tangier.

Sub-fam. 5. Dorylides.—Clypeus extremely small, the antennae
inserted very near the front margin of the head. Hind
body usually elongate and subeylindrical, with an imperfect
pedicel formed by the constriction of the back of the first
segment, but occasionally there are two nodes in the workers.
Distinctions between the two sexes, and between the workers
and sexed forms, enormous, the queens truly wingless. The
Semales and workers usually blind, or at any rate destitute of
Jfacetted eyes. (In Ecitonini the antennae are not inserted
quite at the front of the head, and there are two nodes in the
pedicel. )

IV DORYLIDES—WANDERING ANTS A)

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
maintaimed as distinct from Ponerides. The chief character of
the Dorylides is that the males are much less ant-lke 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 (Zciton) from the Myrmicides.'

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, pecuhar 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. (1.) The EcITONINI
consist of the species of the
genus Heiton, the wandering
ants of America, and of Labi-
dus, which there is now good

reason for believing to con- F!6- 78.—Various forms of worker of Eciton
= hamatum. Guatemala.

sist of the males of Eviton.

The female is still uncertain. The Zciton are nomad ants
having no fixed abode, but wandering from place to place in
search of prey, and forming temporary resting-places. The

? For a valuable revision of Dorylus and its allies see Emery, Zool. Jahrb. Syst.
vill. 1895, pp. 685, etc. We, however, doubt the wisdom of extending the sub-
family so as to include Cerapachys, Purasyscia, ete.

176 HYMENOPTERA 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 W. 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
together 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. I was surprised to see
in this lving 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
the centre of the cluster and brought out clinging to it many
ants holding larvae and pupae.”

Turning now to the JLabidus question: many American
species of this genus have long been known, though all of them

o
(=)

Iv DORYLIDES—-WANDERING ANTS Wry

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
be the male of Heiton, 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, who observed the temporary nests of Heiton hamatum,
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 Miller. 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
Lalidus (L. burchellv) was also found on friendly terms with the
Eciton-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-
fore, 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 André under the name of Pseudodicthadia incerta.
He thought this might. prove to be the female of citon-
Labidus ; but his description and figure are imperfect, and do not
greatly support his idea of a connection between Zeiton and
Pseudodicthadia.

il. The group Dory ini 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.

.

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
Dorylus and of the worker
Typhlopone or Anomma. This
remained for many years with-
out confirmation, but in 1880
Trimen announced the dis-
Erie: 79.—Dorylus helvolus. Africa. A, male ; covery in South Africa of an
B, female (Dicthadia) ; C, worker major

(Typhlopone); D, worker minor. (After Enormous apterous female ant,

Emery. ) 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 should 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, ike 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 1s not unlikely
that considerable variety will be found to prevail. The driver
ants of Africa, belonging to the genus Anomma, are in some

B

Iv DORYLIDES—-DRIVER ANTS 179

respects similar to Heiton in habits, 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 4. arcens, 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
he believes all the
supposed species of
ties genus - to. he
merely varieties of A.

burmeistert. The
female of the driver Frc. 80.—Body of male of Dorylus sp. Delagoa Bay.
. . . oe >» ivisi Ss : S : 7
ants is still quite a, Pronouns i Ls c, ees e Re es, Gs
metanotum ; é, propodeum ;_//, first abdominal seg-
unknown. A Dorylus ment; g, 2, points of insertion of anterior and pos-

r : lerior wings.
has been ascertained . ioe

to be the male of Zyphlopone. 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 7'yphlatta Sm. has been recently
identified by Wroughton and Forel as the worker-condition of
which dAenictus 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 Rhogmus. Only
the male is known.

180 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 lterature
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: 2 unknown.

Pseudodicthadia : female only known, possibly that of Heiton.

Cheliomyrmex : workers and soldiers only known.

Aenictus = the male, Typhlatta = worker: unknown.

Rhogmus: male; female unknown. (According to Emery

the worker is very small and like A/aopone.)

Anomma: only worker known; male probably a Dorylus.

Dorylus = male ; Dicthadia = 9: Alaopone and Typhlopone =

workers.

Sub-Fam. 6. Amblyoponides.— Abdomen destitute of distinct
pedicel ; the articulation between the first and second
segments behind the true petiole being broad.

We follow Forel in separating Amblyopone 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 Scolidae. Forel con-
siders the Amblyoponides to approach
closely to certain divisions of the Scolidae
(Thynnides, e.g.). Little is known of these
Insects, though they are widely distri-
buted. Amblyopone is found in Australia
and New Zealand; the allied genus
Stigmatomma has a wide distribution,

Fra. 81.—A mblyopone, worker.
Tasmania. uae : : :
occurring even in Europe. The social

life is beheved to be imperfect, and the habits subterranean and
sedentary. 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 the fact that a few species of ants are

IV INSECTS INHABITING ANTS’-NESTS 181

used by 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 nitidulus, lives
only in the large nests of Formica rufa; these ants tolerate
the little Formicorenus, which so far as.is known does them
neither good nor harm. There are also a considerable number of
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. Not-
withstanding the militant lhves 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.
Coceidae 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 Wasmann, Berlin 1894.

12 HYMENOPTERA CHAP.

of a 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 lve 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 ry
a ; es Fic. 82.—The beetle, A teme/es, soliciting

most curious of all the ant’s- 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 quid
to appear on the hp, and remaining passive while the guest
partakes of the proffered bonne bouche. 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 Hymenoptercus 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.' 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, ete., 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” 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 lve 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. internat. Zool. iii. 1896,
pp. 411-440. ? For explanation of this term see vol. v. p. 524.

CHAP IMR. V.

COLEOPTERA——OR BEETLES

Order V. Coleoptera.

Apparently wingless Insects when at rest, but really with four
wings; the elytra, or anterior pair, shell-like,. reposing
on the back of the body and fitted together accurately
along the middle by a straight suture; the posterior pair
membranous, folded together wnder the elytra. Mouth with
mandibles ; lower lip not divided along the middle. Meta-
morphosis great and very abrupt ; the larva being a grub or
maggot, which changes to a pupa (usually soft) in which the
external structure of the perfect Insect 1s 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 softand 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-
flees. 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 Vertebrata. 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 185

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. 85). 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

Fra. 83.—Under-surface of a beetle,
Harpalus caliginosus ; legs and
antenna of one side, and some parts
of the mouth removed. A, an-
tenna; B, mandible ; C, labrum ;
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,
buccal fissure ; V, plates of ven-
tral segments. 1, Prosternum ;
2, prosternal episternum ; 3, pro-
sternal epimeron ; 4, anterior and
middle coxal cavities ; 5, inflexed
side of pronotum; 6, mesosternum ;
7, mesosternal episternum; 8,
mesosternal epimeron ; 9, meta-
sternum ; 10, posterior division of
metasternum or ante-coxal piece ;
11, metasternal episternum ; 12,
metasternal epimeron; 18, 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-
conte and Horn.)

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
fight. 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 fivein 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

186 COLEORTERA: 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 metasternum'; 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 Verhoeff has inaugurated a careful study of
the comparative anatomy of the terminal segments’; 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

! 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.

* In a series of memoirs in various German periodicals during the last five or six
years (see especially Deutsche ent. Zeit. 1893 and 1894, also subsequent years of
Arch. Naturges.). It should be noticed that in the course of his studies Verhoeff
has modified some of his earlier views.

— —eeeororrrearene enn

NS ae

Mi 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 tegule 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 1s 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 (Je/oé). 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, 1t 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, Pachypus, 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 eases 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 propygidiun.

188 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 ; 1t 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 seginent: it is
Fic. 84.—Larva of a beetle, Family sometimes armed with hooks.

Tey cidae {arama moscha’e). Three pairs of small thoracic legs

1e first spiracle is placed just at the 5

hind margin of the large prothoracic are often present, but are very

ea habia pie 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,
one 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, lke Lepi-
dopterous pupae, and others (Coccinellidae) are intermediate

Vv 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 1ts 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 distinetions) 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 familes; 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 classity

190 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.
Adephaga—(Caraboidea of some authors).—Antennae never lamelli-
form, 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 palpiform.
Abdomen with six (or more) ventral segments visible.

Series 3. Polymorpha.—Antennae frequently with either a club, ae. the
distal joints broader [Clavicorn series. of authors], or the joints from
the third onwards more or Jess 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. Heteromera.—Front and middle tarsi five-joited, hind tarsi four-

jointed. Other characters very variable.

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.

Strepstptera (see p. 298).

bo

Series

or

Series

The first and second series, with much of the third, form the
Pentamera, the fifth and sixth the Tetramera [ or Pseudotetra-
mera'|, The term Isomera was applied by Leconte and Horn
to a combination of series 1, 2, 3, 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
club, the leaves of which are movable, and in repose are
more ov less perfectly coadapted so as to have the appearance
of being but one piece.

This series includes three families, Passalidae, Lucanidae, and
Scarabaeidae; the latter includes an enormous majority of the
species, and in them the structure of the antennae characteristic
of the series is well developed; but in the other two families

1 We consider this term inferior to Tetramera for nomenclatorial purposes.

Ce ee

v LAMELLICORNS IgI

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 grubs’ with
curved 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 © > tvery
peculiar, each one
being more or less
completely sur-
rounded by a chitin-
guses plates,» The
spiracles usually form

s Fic. 85.—Autennae of Lamellicorns. 1, Neleus inter-

a system entirely ruptus,; 2, Lucanus cervus $; 3, Phanaeus splen-

closed, except at the ine. 2; 4, Phileurus didymus 92; 5, Polyphylla
Jullo 6.

moment when the

skin is shed and the tracheal exuviae are detached. Meinert '
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* 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

1 Danske Selsk. Skr. (6), viii. No. 1, 1895.
? Horae Soc. ent. Ross. xiv. 1879, p. 15.

192 COLEORTERA _ 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.—Labrum large, mobile ; mentum deeply
cut out in the middle for the accommodation of the ligula; the
lamellae of the antenna brought together by the curling up of the
antenna. The elytra entirely cover the dorsal surface of the abdo-
men. There are four or five hundred species of this family known ;
they are usually shining-black, unattractive beetles, of large size,

Fic. 86.—View of one
side of meso- and
metathorax of a
Passalid larva from
Borneo showing the
stridulating organs.
a, 6, Portions of the
metathorax ; ¢, coxa
of 2nd leg ; d, striate
or stridulating area
thereon ; e, basal
part of femur of
middle leg ; 7, hairs
with chitinous pro-
cess at base of each ;
g, the diminutive
3rd leg modified
for scratching the
striated area. x 20.

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 species
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

Vv ; LAMELLICORNIA——STAG-BEETLES 193

provided with highly-developed stridulatory structures. No
auditory organ is known, unless the peculiar spiracles be
such.

Fam. 2. Lucanidae (Stag-beetles)—Labrum indistinct, fined ;
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
mandibles bear as many
Fic. 87.—Head and prothorax of forms of the male of projections as the large

a stag-beetle ; Homoeoderus medlyi (Africa). A, forms do, or even more.

Large, B, intermediate, C, small forms. (From a it His Hee Pn
photograph by R. Oberthiir. ) n 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 hable 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 Lucanidae, and instead of
the mandibles of the male being
Fic. 88.—Sinodendron cylindricum. 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 Melolontha vulgaris,
but attains a larger size, and the anal aperture is placed longitu-
dinally instead of transversely ; 1t 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

Vv LAMELLICORNIA—-SCARABAEIDAE 195

usually leave the pygidium uncovered. The number of visible
ventral segments is usually six, or at the sides seven, not five, as in
Lucanidae and Passalidae. This is one of the most important
famihes 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 thé classification of this extensive
family, which, moreover, be it remarked, is increasing by the dis-
covery of about 500 new species every year.

Abdominal spiracles piaced in a line on the connecting membranes, and en-
tirely covered by the wing-cases (Laparosticti). Sub-fam. 1. Copriprs.!

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-fam. 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. DyNasTIpDEs.
The claws of the tarsi equal; the front coxae more prominent, shorter
transversely. Sub-fam. 5. CETONIIDES.

1, The CopribEs form an immense group of about 5000 species ;
they differ somewhat in habits from other Lamellicorns, inasmuch
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. 7Zvox 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 in which the elytra cover the
pygidium, and in which the number of conspicuous ventral segments is reduced to
five or even four. We use the term Coprides as equivalent to the ‘‘ Laparosticti”

of Lacordaire (Gen. Col. iii. 1856) ; it thus includes the ‘‘Conrini” and ‘‘ Glaphy-
rini” of the Catalogus Coleopterorum, vol. iv. Munich, 1869.

196 COLEOPTERA CHAP.

for stridulation 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. No 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 1s 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 valley of the Nile 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 hfe. Many accounts have been given
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 Scwrabaeus detaches a portion of the dung
and forms it into 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.
When 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

v _ LAMELLICORNIA—COPRIDES 197

exhausted, when the Scarabaeus seeks a fresh store of provender
to be 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 Scarabaeus 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
Ob thes ere in! such, a
manner that the grub to
be hatched from the ege
shall find the softest and
most nutritive portions
close to it, while the
coarser and more innu-
tritious parts are arranged
so as 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
beetle 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 egg
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 oviposition, survives, and sees the growth of her
young to the perfect state, and then produces another generation.
No similar case can be pointed out in Insects, except in the Social
kinds; but from these the Coprides observed by Fabre differ pro-
foundly, inasmuch as the number of eggs produced by the mother
is extremely small; Copris hispanus, for instance, producing in
each of its acts of oviposition only one, two, or three eggs.

Fig. 89.—Scarabaeus sacer. Portugal.

198 COLEOPTERA CHAP.

uu. 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. Jelolontha vulgaris, 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! on its
anatomy is one of the classical works of Entomology. This In-
sect 18 SO injurious in some parts of France that money is paid
by the local authorities for its destruction. M. Reiset informs

=

us that under this arrangement 867,175,000 perfect cockchafers,

and 647,000,000 larvae were destroyed in the Seine-inférieure
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 JZ vulgaris oceupies 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.”
In the perfect state the Insect is sometimes injurious from the
large quantity of foliage it destroys. Schiddte* 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.
il. The RuTELIDES number about 1500 species ; there are many

1 Considérations genérales sur Vanatomie comparée des animaux articulés, etc.,
Paris 1828, 4to. xix. and 435 pp., and Atlas of ten (xx.) plates, and 36 pp.

? Raspail, Mém. soc. zool. France, vi. 1893, pp. 202-213.

3 Ann. soc. ent. France, (v.) iv. 1874, p. 39.

i

V LAMELLICORNIA——-DYNASTIDES—CETONIIDES 199

Insects of brilliant metaflic colours amongst them, but very little
is known as to their life-histories. The larvae are very much
hike 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
largest 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 nasicornis
is sometimes found in numbers in spent tan. The growth and
development of the individual is believed to be but slow.

y. The CETONIIDES are renowned for the beauty of their colours
and the elegance of their forms; hence they are a iavourite
group, and about 1600 species have been catalogued. They are
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 flight is peculiar; the elytra do not extend down the

200 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 Rose-chafers. The larvae of C. floricola 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 Huphoria 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
North 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 Zomap-
tera stridulate in a pecuhar manner, by rubbing the edges of the
hind femora over a striate area on the ventral seements.

Series II. Adephaga or Caraboidea.

All the tarsi five-jointed ; antennae filiform, or nearly so; mouth-
parts highly developed, the outer lobe of the maxilla nearly
always divided into a two-jointed palpus ; supports of the
labial palpi developed as joints of the palpi, and in some
cases approximate at their bases. Abdomen with the exposed
segments one more in number at the sides than along the
middle, the number being usually five along the middle, six
at cach 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 Carnivora or
Filicormia. 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

‘

Vv ADEPHAGA—TIGER-BEETLES 201

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
Rhyssodidae; 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 (7 iger-beetles).—Clypeus extending
laterally in front of the insertion of the antennae. Lower lip with
the palpi usually greatly developed, but with the ligula and para-
glossae very much reduced, often scarcely to be detected. Maxillae
with the outer lobe forming a two-jointed palp,’ the inner lobe
elongate, furnished at the tip with a hook-like process, which is
usually articulated by a joint with the 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 MJegacephala, 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 bafile the most practised hand and eye.” He further
says that the species he observed (being of diverse colours)
agreed in colour with the general colours of the “locale they
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

* In Theratides this outer lobe is in a rudimentary state, like a seta.

202 COLEOPTERA CHAP.

in this position waits for its prey. This consists of Insects that
may alight on the spot or run over it.. When 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 be 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 Cicindela, most of which
are so active on the wing, agree
in these respects with the African
species of Manticora, which are
entirely apterous, and are the
Fie. 80. 5 Crema nu erides BNE largest of the Cicindelidae. Pér-

A, larva (after Schisdte) ; B, imago, Inguey found a breeding-ground

eee of IZ. tuberculata 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 Cicindelidae, 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.'

Tiger-beetles display considerable variety of structure, especially
as regards the mouth, which exhibits very remarkable develop-

1 The first portion of a classification of Cicindelidae by Dr. Walther Horn,
Revision der Cicindeliden, Berlin, 1898, has appeared since this was written.

Vv CICINDELIDAE——-TIGER-BEETLES 20

ments 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 innutritious 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-
. - Fic. 91.—Mouth- parts of tiger-beetles. A,
mentary canal that it is Profile of Pogonostoma sp. (Madagascar): a,
difficult to find the orifice of antenna ; 6, labial palp; c, maxillary palp ;
Sra wie : d, palpiform lobe of maxilla ; e, mandible ; /,
communication ; the labium

labrum. B, Section of head of Mamnticora
being much modified to maxillosa (South Africa): ¢, front of upper

: ‘ part of head-capsule ; 6, gula; c, tentorium ;
form the posterior wall. d, eye ; e, labrum ; f, left mandible ; y, max-
For the capture of the prey, a & HEE UIE? palps 4, labial palp; 4,
a : hee support of this palp ; 7, labium.

always living but of various

kinds, a mechanism with great holding power and capable of
rapid action is required. The mouth of the terrestrial Manticore
(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
only the mounds of Termites. The characteristic feature common

204 COLEOPTERA CHAP. ©

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. Raffray
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 (Grownd-beetles)—Clypeus not extending
laterally in front of the antennae. Maxillae with the outer lobe
destitute of an articulated hook at the tip. Antennae covered
(except the basal joints) with a minute pubescence. Hind legs not
very different from the middle pair, formed for running, as usual
in beetles. 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
Carabidae 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.
92, A) are usually elon-

Fic. 92.—Leistus spinibarbis. A, Larva (after é >
Schiodte) ; B, imago. Britain. 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

aN ADEPHAGA—-CARABIDAE ZO

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 Pferostichus and Harpalus, are said to be fond of
ripe strawberries. The most anomalous forms of Carabidae are the
Pseudomorphides, a sub-family almost pecular 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 J/ormolyce 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 Anophthalmus, 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 the
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 7vechus, 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
eavernicolous 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 dépus, 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! A few Carabidae, of the genera Glyptus and Orthogonius,
are believed to live in the nests of Termites. Savage found the
larva of G. sculptilis in the nests of Termes bellicosus; 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.” Mr. Haviland found Rhopalomelus angusticollis in
Termites’ nests in South Africa. Péringuey 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 Helluodes 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. G. H. Horn ;? he arranges Carabidae
in three sub-famihes; we think it necessary to add a fourth for
Mormolyce :

1. Middle coxal cavities enclosed externally by the junction of the meso- and
meta-sternum ; neither epimeron nor episternum attaining the
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 epimera,
but not by those of the episterna. Sub-fam. 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

1 Natural History of aquatic Insects, 1895, p. 376.

2 Tr. Amer. ent. Soc. xv. 1888, p. 18.
3 Op. ctt. v. 1881, p. 91; cf. Sharp, Zr. ent. Soc. London, 1882, p. 61.

Vv 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
Mormolyce 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 lmited to the genus
Amphizoa ; 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
North 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; ae
they dot snot swim, wae FIG. perrar Ue ee pee ene America.

; ; 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 Carabus ;
but at 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 than the tibiae. This family
is limited to the one genus Pelobius (Hygrobia of some authors).
Like Amphizoa, to which it is in several respects analogous, it
has 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. Pelobius may be brietly
described as a Carabid adapted to a considerable extent for
living in and swimming about in water; differing thus from

Fic. 94.—Pelobius turdus. Britain. A, Young larva; B, adult larva; C, imago.
(A and B after Schiddte. )

Amphizoa, which has no special adaptation for swimming. The
larva of Pelobius is remarkable; it breathes by means of branchial
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 Pelobius
tardus is remarkable for its loud stridulation. The sound is pro-
duced by an apparatus described correctly by Charles Darwin ;*

1 Descent of Man, i. 1890, p. 338; The views of Landois and Recker, Arch. f.
Naturgesch. lyii. 1, 1891, p. 101, are erroneous.

Vv 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 file therewith.
The Insects are called squeakers in the Covent Garden market,
where they are sold.

Fam. 8. Haliplidae.— Antennae bare, ten-jointed ; meta-
sternum marked by a transverse line; posterior coxae prolonged
as plates, covering a large
part of the lower surface
of the abdomen; the
slender, but clubbed, hind
femora move between these
plates and the abdomen.
The MHaliplidae 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 1s 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,
B), there are no spiracles,
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

/

Fia. 95.—Cnemidotus caesus. England. A, Imago ;
B, larva, highly magnified, (After Schiddte.)

2m© 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 swimming, not capable of ordinary walking : meta-
sternum without a transverse line across it ; behind closely united
with the extremely large covae. 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-

Fia. 96.—Cybister roeseli (=laterimarginalis De G.) vinalbimetausiar }
Europe. A, Larva (after Schiddte) ; B, ¢ imago. ginal instars are Much
less profoundly

changed, and are capable of existing perfectly well on land, and
of taking prolonged flights through the air; (3) the pupa is, so
far as known, always terrestrial. The larvae and imagos 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.! The hind
pair of legs is the chief means of locomotion. These swimming-
legs (Fig. 97) are deserving of admiration on account of their
mechanical perfection; this, however, is exhibited in various
1 See J. Linn. Sov. Zool. xiii. 1876, p. 161.

V ADEPHAGA—WATER-BEETLES Cileu

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-eases, ‘The air thot Ps, $7: oslomlogies of Cte
enters the space between the B, the movable parts in the striking posi-
ese tt axa) im) Oust Momeni 6 fms
there when the Insect closes

the chink and again dives beneath the water. The enlargement
of the terminal stigmata in Dytisews is exceptional, and in forms
more highly cooehnred in other respects, such as Cybister, these
spiracles remain minute; the presumption being that in this
case respiration is carried on almost entirely by means of the
supply the Insect carries in the space between the elytra and the
base of the abdomen.’ The structure of the front foot of the male
Dytiscus, and of many other water-beetles, is highly remarkable,
the foot being dilated te 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

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. 60
the large stigma on the terminal segment of the abdomen is omitted, though it is
referred to in the text.)

2ilee 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. 213

best 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.!

Series III. Polymorpha.

Antennae frequently either thicker at the tip (clavicorn) or serrate
along their inner edye (serricorn) ; but these characters, as
well as the number of joints in the feet and other 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 Loman 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,” Sez. Trans. R.
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 the pronotum—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,’ and Raffray 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. faviert, is not uncommon
in the Iberian peninsula and South France,
and a’single species was formerly found in
Brazil. The position the family should
Fic. 98.— Paussus cepha- occupy has been much discussed; the only
ie tafe) 4% forms to which they make any real ap-
(After Raffray.) y y i
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 Raffray’s view (he being the most
recent authority on the family)? who concludes that this is
an anomalous group not intimately connected with any other
family of Coleoptera, though having more affinity to Carabidae
than to anything else. The recently discovered genus Proto-
poussus 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 Raffray is erroneous; in a specimen of the
genus Arthropterus the writer has dissected, he finds that there
' Descriptions of larvae that may possibly be those of Paussids have been pub-
lished by Xambeu, Ann. Soe. Linn. Lyon, xxxix. 1892, p. 137, and Erichson, Arch.
Naturgesch. xiii. 1847, p. 275.
2 Arch. Mus. Paris (2), viii. and ix. 1887.

Vv 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
along 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

Fic. 99.—A, Larva of Gyrinus (after Schiddte) ; B,
waters. Of all these, under side of Gyrinus sp. (after Ganglbauer). 1,
ee Some aay ai Prosternum ; 2, anterior coxal cavity ; 3. meso-
Gyrinidae are in their thoracic episternum ; 4, mesoepimeron ; 5, meso-
construction the most sternum ; 6, metathoracic episternum ; 7, middle

af Pee fay: ] coxal cavity ; 8, metasternum ; 9, hind coxa; 10,
adapte or such a ventral segments. [N.B.—The first ventral segment
career. They are able really consists, at each side, of two segments united ;
: : this may be distinctly seen in many Gyrinidae. ]

to dive to escape danger,

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

w i pele 8 )

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

DOr, 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 lfe, 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 Hnhydrus and
the Porrorhynchus of tropical Asia.’ 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 Adephagous
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-
onomists to the maintenance of isolated families. Surely if
there are in nature some families allied and others isolated, it 1s
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.—7arsi jfive-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 Régimbart, Ann. Soc. ent.
France, 1882, 1883, and 1886. For a catalogue, Séverin, Ann. Soc. ent. Belgique,
XXXxili. 1889.

v POLYMORPHA——HYDROPHILIDAE PAM AE

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
larger (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 often 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 five 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 lttle mast, which is supposed by
some to be for the purpose of securing air for the eggs. Helo-
chares 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 Dumeril
Fic. 100. — Sercheus emar- states that in Hydrous caraboides the

ginatus ¢. Britain. A, J

Upper surface of beetle; alimentary canal undergoes a_ great

Be er ae Eee change at the period of metamorphosis,

ruptured and some of the becoming very elongate in the adult,

eg 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-lke 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 Serosus the
terminal stigmata are obsolete, and the sides of the body bear
long branchial filaments. Cussace says that in Spercheus (Fig.
101) there are seven pairs of abdominal spiracles, and that the
larva breathes by presenting these to the air;+ but Schiddte
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 Hydrobius. No doubt Cussae was wrong in
supposing the peculiar lateral abdominal processes to be stig-

1 Ann. Soc. ent. France, xxi. 1852, p. 619.

V POLYMORPHA—HYDROPHILIDAE——-PLATYPSYLLIDAE 219

matiferous. In Serosus there are patches of aériferous, minute
pubescence on the body. The pupae of Hydrophilides repose on
the dorsal surface, which is protected by
splous 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 Sphaeridiides 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- Fre. 101.—Larva of Spercheus
ing. In Britain we have nearly ninety Sen ee aE ae
species. Some taxonomists 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
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 Riley.’ Little is known as to the
food and life-history. Horn states that the eggs are placed on

‘ Horn, Tr. Amer. ent. Soc. xv. 1888, p. 23 ; Riley, Insect Life, i. 1889, p. 300.
I J y I

220

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

§ Sy)

Sot Hs

Fic. 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
mingaudi ; others have supposed that it eats scales of epithelium

Fic. 103.—Leptinus testaceus.
Britain.

or hairs of the beaver.

Fam. 14. Leptinidae. — Antennae
rather long, eleven-jointed, without club, but
a little thicker at the extremity. Eyes absent
or imperfect. Tarsi five-jointed. LElytra
quite covering abdomen. Mentum with the
posterior angles spinously 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
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

a

en a A

Vv POLYMORPHA—_LEPTINIDAE——SILPHIDAE 2

i)
al

in curious places, including the nests of mice and buimble-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 alhed 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.—Zhe 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 cozae are conical and contiguous :
prothoracie epimera and episterna not distinct. Visible abdomi-
nal segments usually five, but sometimes only four, or as many as
seven. Tarsi frequently five-jointed, but often with one joint less.
Elytra usually covering the body and free at the tips, but oceasion-
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 (ecrophorus), so well known from their habit
of making excavations under the corpses of small Vertebrates,
so as to bury them. Besides these and Si/pha, 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.

* Insect Life, i. 1889, pp. 200 and 306.

COLEOPTERA CHAP.

NO
NO
NO

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 S.
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 Si/pha larvae
live, like the Necro-
phorus, on decomposing
animal matter, but run

=

/
H
|
Lo |
o|
aN) i

ow

Fra. 104.—A, Larva of Silpha obscura. Europe. about to seek it ; hence
Gatton eanede) B, Ptomaphila lacrymosa, many specimens of some
mie of these large larvae may

sometimes be found amongst the bones of a very small dead bird.
We have found the larva and imago of S. thoracica in birds’ nests
containing dead nestlings. S. atrata and S. laevigata make war
on snails. S. /apponica enters the houses in Lapland and ravages
the stores of animal provisions. S. 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 truftle-beetle.

The number of species of Silphidae known must be at present
nearer 900 than 800. Of these an unusually large proportion be-
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 in con-
siderable force in New Zealand; and is probably well represented

v SCYDMAENIDAE—_GNOSTIDAE——PSELAPHIDAE

i)
i)

Los)

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 Dr. Horn 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.—JMinute Insects allied to Silphidae,
but with the hind coxae separated, and the facets of the eyes coarser ;
the tarsi are five-jointed ; the number of visible abdominal segments is
siz. 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. Jastigus
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
Staphylnidae rather than to Seydmaenidae. The South European
Leptomastax is remarkable on account of the slender, long, sickle-
shaped mandibles. The Oriental genus C/idicus 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. — Winute Insects with three-jointed
antennae, five-jointed tarsi, and three apparent ventral segments, the
jirst of which, however, is elongate, and consists of three united
plates. Hlytra 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.”

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. Tarsi with
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.
2 Westwood, Zr. ent. Soc. London (N.S.) iii. 1855, p. 90; Wasmann, Kit.
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 on 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 Chenniwm bituber-
culatum—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 Raffray,| who has himself discovered
and described a large part of the known species.

Fam. 19. Staphylinidae.—H/ytra very short, leaving always
some of the abdominal segments exposed, and covering usually only
two of the segments. Abdomen usually elongate, with ten dorsal, and
seven or eight ventral segments ; of the latter six or seven are usually
exposed ; the dorsal plates as hard as the ventral, except sometimes
in the case of the first two segments; the segments very mobile, so
that the abdomen can be curled upwards. The number of tarsal
joints very variable, often five, but frequently as few as three, and
not always 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.” At present

1 Rev. ent. franc. ix. 1890.
Die Kifer von Mittelewropa : I. Familienreihe, Staphylinoidea. Vienna, 1895
and 1899.

Vv POLYMORPHA—STAPIIYLINIDAE 225

about 9000 species are known, some of which are minute, while
scarcely any attain a size of more than an inch in length, our
common British black cock-tail, or “devil’s coach-horse beetle,”
Ocypus olens, 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 1t overcomes this difficulty
by shght 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
oo pegments oe Fic. 105.—Staphylinidae. A, Larva of Phi-
Insect may be is at present lonthus nitidus. Britain. (After Schiddte.)
quite mysterious. The habits aa pea caer na vegas
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, 7.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. VI Q

i)
to
OV

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 Lomechusa 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 Lomechusa, as described by
Wasmann (to whom we are indebted for most of our knowledge
of this subject), 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 als» do 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 Atemeles 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 recelving any direct support from them. The most

1 Vergleichende Studien iiber Ameisengdste, Nijhoff, 1890; and Tijdschr. ent.
Xxx. 1890, pp. 93, etc. ; Biol. Centralbl. xv. 1895, p. 632.

(esl werd YL NTDAE——SPHABRIIDAE-——TRICHOPTERYGIDAE 227

remarkable Staphylinidae yet discovered are some viviparous
species, forming the genera Corotoca and Spirachtha, that have
very swollen abdomens, and live in the nests of Termites in
Brazil:* very little is, however, known about them. <A very
large 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
Veileius has the power of making itself disagreeable to the
hornets by some odour, and they do not seriously attack it. The
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 labrum. The pupae of some are
obtected, z.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 1s 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 with only
three visible ventral segments. This family
includes only three or four species of In-
secis about 4, ofan inch long. They are Fie. 106.—Prichopteryx faset-
7 cularis. Britain. A, Out-
very convex, and may be found walking line of perfect Insect; B,
onmud. SS. acaroides occurs in our fens. ae = ea gs iS
Mr. Matthews considers that they are above; E, pupa; F, wing;
most nearly allied to Hydrophilidae.’ ee ruelob imag.
Fam. 21. Trichopterygidae.—Lztremely minute: antennae

1 Schiddte, Ann. Sci. Nat. Zool. (4) v. 1857, p. 169.
* Biol. Centr. Amer. Col. ii. pt. i. 1888, p. 156.

i)
i)
ice)

COLEOPTERA CHAP.

clavicorn (basal and apical joints thicker than middle joints) ; tarsi
three-jointed ; elytra sometimes covering abdomen, in other cases
leaving a variable number of seyments exposed ; wings fringed.
This family comprises the smallest Insects ; Vanosella fungi being
only =), of an inch long, while the largest Trichopterygid is
only ;4, 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.’

Fam. 22. Hydroscaphidae.—Lxtremely minute aquatic Insects,
with elongate 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.—JIinute beetles. Tarsi four-jointed,
but appearing only three-jointed, owing to the hind joint being
concealed by the emarginate (or notched) second joint. Sia free
ventral segments. Masxillae with only one lobe. Antennae 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
Orthoperus brunnipes was found
abundantly by Perris in thatch in
France. Mr. Matthews proposes
to separate the genus Aphanoceph-

| alus as a distinct family, Pseudocory-

Fic. 107.—A, Larva of Orthoperus lophidae.* In Corylophidae the
brunnipes (after Perris); B, O. ato- : . : ; oui

marius, perfect Insect. Britain, Wings are fringed with long hairs,

as is the case in so many small

Insects: the species of Aphanocephalus are rather larger Insects,
and the wings are not fringed; the tarsi are only three-jointed.

* Monograph, Trichopterygia illustrata, by A. Matthews, London, 1872.
* For further information refer to Matthews, An Essay on Hydroscapha, London,
1876, 20 pp. 1 pl. 3 Ann. Nat. Hist. (5) xix. 1887, p. 115.

v POLYMORPRA—SCAPHIDIIDAE—SYNTELIIDAE 229

Fam. 24. Scaphidiidae.— Front coxae small, conical; prothorax
very closely applied to the after-body ; hind coxae transverse, widely
separated: abdomen with six or seven visible ventral plates ;
antennae at the extremity with about five joints that become
gradually broader. Tarsi five-jointed. This family consists of a
few beetles that 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
Seaphisoma agaricinum by Perris; itis mega eae audi
like the larva of Staphylinidae, there are cinum. Britain. A
nine abdominal segments in addition to a aa ane eurishs B
very short, broad pseudopod, and very short
cercl. This larva feeds on agaries ; it goes through its development
in about three weeks; unlike the adult it is not very active.

Fam. 25. Synteliidae.— Antennae clavicorn, with very large
club: labium, with hypoglottis and the parts beyond it, exposed.
Front coxae transverse. Abdomen with five visible ventral segments,
and eight or nine dorsal, the basal four 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 * allied to Silphidae. The
perfect Insects live on the sap running from
trees: but nothing else 1s known of their
natural history. Like so many others of
the very small families of aberrant Cole-
optera, it has a very wide distribution ;
Fic. 109.—Syntelia west. Syntelia being found in Eastern Asia and

ee dee Mexico, while the sub-family Sphaeritides

occurs, as a single species, in Europe and
North America. The earlier instars are unknown.

' Larves de Coléoptéres, 1878, p. 11, pl. i. 2 Biol. Centr. Amer. Col.ii. pt. i. p. 438.

230 COLEOPTERA CHAP.

Fam. 26. Histeridae.—Very compact beetles, with very hard
integument, short, bent antennae, with a very compact club: no
hypoglottis. Elytra closely applied to body, but straight behind,
leaving two segments exposed. Abdomen with five visible ventral
segments ; with seven dorsal segments, all hard. Front coaxae
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 ;' in Britain
we have about 40. The larvae
are without ocelli or labrum,
Fic, 110.—Platysoma depressum. Europe. but have well-developed mandi-

Tee te eS eae ane bles, the second and _ third

thoracic segments being short,
the ninth segment of the abdomen terminal, with two distinctly
jointed cerci.” 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 (7rypanaeus) ; 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

1 The family was monographed by the Abbé de Marseul in Ann. Soc. ent. France,
1853-1862, but great additions have been made since then.
* For characters of larvae of various genera, see Perris, Larves, etc. p. 24.

Vv. HistERIDAR——PHATACRIDAE—-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 lve in company with Termites.

Fam. 27. Phalacridae.—-Lody 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
Fic. 111. — Olibrus bicolor. pass the Insects over as having only four-

Europe. A, Larva (after - . : A

Heeger) ; B, perfect Insect. jointed tarsi; their structure, so far as

the front two pairs are concerned, being
very nearly that of many Phytophaga. The laryae 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’ 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 five visible plates. These Insects are numerous,
about 1600 species being at present known; in many of them
the elytra nearly or quite 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
earcases. We have about 90 species in Britain ; several forms of

1 SB. Ak. Wien. xxiv. 1857, p. 330.

Zee COLEOPTERA CHAP.

the genera Meligethes and Epuraea are among the most abundant
of our beetles. Most of what is known as to the larvae is
due to Perris; several have been found living in flowers; that
of Pria haunts the flower of Solanum dulcamara at the junction
of the stamens with the corolla; the larva of JZeligethes aeneus
sometimes occasions much loss by prevent-
ing the formation of seed in cultivated
Cruciferae, such as Rape. 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 Soronia lives in
Fic. 112.—Pria dulcamarae, 'eTMenting sap, and has four hooks curv-
Britain. A, Larva (after ing upwards at the extremity of the body.
Perris) ; B, perfect Insect. : :
The curious genus Cybocephalus 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 hizophagus has given rise to
much discussion; although now usually placed in Nitidulidae,
we think it undoubtedly belongs to Cucujidae.

Fam. 29. Trogositidae.— Differs from Nitidulidae in the struc-
ture of the tarsi ; these appear to be four-
jointed, with the third joint similar in
size and form to the preceding; they
are, however, really five-jointed, an ex-
tremely short basal joint being present.
Hind coxae contiguous. The club of each
antenna is bilaterally asymmetric, and

the sensitive surface is confined to certain
parts of the joints. There are some 400
or 000 species of Trogositidae, but nearly
all of them are exotic. The larvae (Fig.
113, A), are predaceous, destroying other Fic. 113.—Temnochila coerulea.
? ? ) J 5

. wes Europe. A, Larva (after
larvae in large numbers, and it is pro- _perris) ; B, perfect Insect.
bable that the imagos do the same.
The larva of Tenebroides (better known as Trogosita) mauritanica
is found in corn and flour, and is said to have sometimes been very

eee

V POLYMORPHA—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. Nemosomides,
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.!
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 ; five 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; New Zealand has produced
the greatest variety of forms; the forests
of Teneriffe are rich in the genus Zar-
phius. 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
Fic. 114.—Bitoma crenata, underground and is quite blind, the
ae roo optic lobes being absent. Some Coly-
diidae are more active, and enter the
burrows of wood-boring Insects to destroy the larvae (Colydium).
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.?

' Catalogue of Trogositidae, by Leveillé, in Ann. Soc. ent. France, 1888, p. 429.
* For classification, see Sharp, Biol. Centr. Amer. Col. ii. pt. i. 1894, p. 443.

234 COLEOPTERA CHAP.

Fam. 31. Rhysodidae.— 7Zarsi fowr-jointed ; mouth-parts
covered by the large mentum ; front tibiae notched on the inner
edge. This family consists only of a few species, but is found
nearly 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 Adephaga,’ 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
familes of the Coleoptera Polymorpha, but also with most of the
ereat series; Adephaga, Rhynchophora, Phytophaga, and Hetero-
mera, being each closely approached.

Fam. 32. Cucujidae.—TZwisi five- or four-jointed, the first
joint often short: antennae sometimes clubbed, but more often
quite thin at the tip; front and middle coxae deeply embedded,
globular, but with an angular prolongation externally ; abdomen
with five visible ventral segments, all movable. This family and
the Cryptophagidae are amongst the
most difficult families to define ;
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 flat.
Many of the larvae are also very
Fie. 115.—Brontes planatus, Britain. flat, but Perris says there is great

A, Larva; B, pupa; C, perfect 4. : : Wag ; =
Insect. (A and B after Perris.) | Clversity in their structure: -they
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, Aaf. Mittelewropas, i. p. 530, as well as Leconte and Horn
Classification, etc., p. 130.

V POLYMOR PHA——CRYPTOPHAGIDAE—-HELOTIDAE 2: 35

Fam. 33. Cryptophagidae.—/ront and middle coxae very
small and deeply embedded; antennae with enlarged terminal joints;
tarsi five-jointed, the posterior sometimes in the male only four-
jointed ; abdomen with five visible ventral segments, capable of
movement, 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 (Antherophagus, Telmatophi-
/us) are found chiefly on growing plants,
especially in flowers. Although the
unago of Antherophagus lives in flowers,
yet the larva has only been found in ‘
the nests of bumble-bees; there is Fic. 116.—Cryptophagus denta-
reason for believing that the imago a eee oe ides
makes use of the bee to transport
it from the flowers it haunts to the nests in which it is to
breed;+ 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 Perris found the one we figure.

Fam. 34. Helotidae.—J/ront 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 Perris, Larves, etc., p. 75.

236 COLEOPTERA CHAP.

There is but one genus, 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.’
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
region (including Japan) with one species in Eastern Africa.

Fam. 35. Thorictidae.—7arsi jive-jointed, none of the joints
broad ; front coxae small, rather prominent, but not at all trans-
verse; five visible ventral plates, all mobile; metasternum very
short ; antennae short, with a solid club. This httle family, con-
sisting of the genus Zhorictus, 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
pronotum, or on the under surface of the body on each side of the
breast ; Wasmann thinks that when the beetles are riding about,
as above described, the ants have then
an opportunity of getting at the patches
on the under surface.

Fam. 36. Erotylidae.—7arsi jive-
jointed, but with the fourth usually very
small, the first three more or less broad
and pubescent beneath. Antennae strongly
clubbed. Front and middle coxal aceta-

h bula round, without angular prolongation
Fic. 117.—Tritoma Bipustulata. externally ; five visible ventral segments.
Erotylidae. Britain. A, Larva This is now a large and important family
(after Perris) ; B, perfect Insect. = ; rage os
of about 1800 species, but it is chiefly

exotic and tropical, its members haunting the fungoid growths

1 Ritsema, Catalogue of Helota, Notes Leyden Mus. xiii. 1891, p. 223, and xv.
1893, p. 160.

V EROTY LIDAE——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 Languriides (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 Langurides.
Fam. 37. Mycetophagidae.—Zarsi four-jointed, slender, the
front feet of the male only three-jointed ; coxae oval, not deeply
embedded ; abdomen with five ventral seq-
ments, all movable. A small family, of in-
terest chiefly because of the anomaly in the
feet of the two sexes, for which it 1s 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 4, Menta ae:
about a dozen species in Britain, and ciatus. Mycetophagidae.
scarcely 100 are described from all the Sas Sateen
world. The Diphyllides, placed by Leconte
and Horn in this family, seem to go better in Cryptophagidae.
Fam. 38. Coccinellidae (Lady-birds).—Tuarsi apparently three-
jointed ; the first two joints pubescent beneath ; the third joint
consisting really of two joints, the small true third joint being
inserted near the base of the second joint, the upper surface of which
is grooved to receive it. Head much concealed by 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—hbears, 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,

AEN COLEOPTERA CHAP.

and Acari that are so injurious to cultivated plants. They
also eat various other soft-bodied Insects that attack plants. As
they are excessively voracious, and are themselves singularly
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-family
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.’

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 concomitantly change a good deal in
colour and form; the larval life does not usually exceed four or
five weeks; at the end of which time the larva suspends itself
by the posterior extremity, which is glued by a secretion to some
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 extremity of the body, but retain it as a covering for the
pupa. The larvae that feed on plants are much less active than
the predaceous forms. We are well supphed 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 Zool. Anz. xviii. 1895, p. 244.

V COCCINELLIDAE—-EN DOMYCHIDAE——-MYCETAEIDAE . 239

Pseudotrimera. But they are generally placed in the Clavicorn
series, near Endomychidae. Verhoeff has recently made con-
siderable morphological studies on the male genital organs of
Coleoptera, and as the result, he concludes that Coccinellidae
differ radically from all other Coleoptera as regards ‘these
structures, and he therefore treats them as a distinct series or
sub-order, termed Siphonophora. The genus Lithophilus has been
considered doubtfully a member of Coccinellidae, as the tarsi
possess only in a slight degree the shape characteristic of the
family: 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
treats as another separate group, Coccidulini.

Fam. 39. Endomychidae.!\— Varsi apparently three-jointed, the
Jirst two joints broad, the terminal joint elongate ; at the base of
the terminal joint there is, however, a very small joint, so that the
tarsi are pseudotetramerous ; antennae rather large, with a large
club ; labium not at all retracted behind the mentum ; front and
middle 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. We 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.—7Zursi fowr-
jointed, the first two joints not very different
from the third, usually slender; abdomen
with five visible ventral segments, which are
movable; front and middle coxae globular.

The little Insects composing this family are F eee eer res
by many placed as a division of Endomy- (after Blisson) ; B, per-
chidae, and Verhoeff is of opinion that the — '¢t I™sect

group is an altogether artificial one; but we think, with Duval,
it makes matters simpler to separate them. There are only

? Gerstaecker, Monographie der Endomychiden, Leipzig, 858, 1433 pp. Since this
work was published, the species known have been multiplied two or three times.

240 COME ORME RA CHAP.

some forty or fifty species, found chiefly in Europe and North
America. We have three in Britain; one of these, Mycetaea
hirta 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.—TZarsi three-jointed ; anterior coral
cavities round, not prolonged externally ; abdomen with five visible
and mobile ventral segments. 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
Perris to have the mandibles replaced by fleshy appendages. The
pupa of Latridius is remarkable, on account of the numerous long
hairs with heads instead of points; the larva of Corticaria is
very like that of Latridius, but some of the hairs are replaced by
obconical projections. The sub-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 J/onotoma
live in ants’ nests, but probably have
no relations with the ants. <A few
species of Latridiudes proper also main-
tain a similar life; Coluocera formicaria
is said to be fond of the stores laid up
by Aphaenogaster 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 Fic. 120.—Latridius minutus.
have about 40 species in Britain. The peeeae Pee
North American genus Stephostethus has
the prosternum constructed behind the coxae, somewhat in the
same manner as it 1s in the Rhynchophorous series of Coleoptera.

Fam. 42. Adimeridae.—T7warsi appearing only tio-jointed,
a broad basal joint and an elongate claw-bearing joint, but between
the two there are two very small joints. This family consists
only of the American genus Adimerus; nothing is known of

Vv 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— 7Z'a7rsi
Jive-jointed ; antennae usually short,
with the club frequently very large
in proportion, and with the under
side of the thorax bearing a hollow
Jor its reception. Front coxae rather
long, oblique: hind coxa formed to
receive the femur when in repose.
A family of 500 or 400 species of
small or moderate-sized beetles; the F's. 121.—Adimerus setosus. Adi-
: meridae. A, the Insect; B, one
surface, usually covered with fine foot more enlarged. Mexico. From
hair, forming a pattern, or with LONE AE EGS CH gett NPL
scales. Byturus, 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 im
Fic. 122.—Tiresias serra. furniture. The young larva of this species
Larva. New Forest. : .
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 Yiresias serra, which lives amongst cobwebs in
VOL. VI R

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 (P1//-beetles).—Oval orround,convex beetles ;
tarsi five-jointed, front coxae not exserted, transverse ; hind coxa
shielding the retracted femur. The whole 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
inanunate 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
no 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 mentum, abruptly clubbed antennae,
and the head cannot be retracted and concealed. The genus
Nosodendron seems 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-
podeal 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

ee

Vv 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—J/inute 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, clubbed ; tarsi four-
jointed ; 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
JSorwards; antennae short, the terminal seven joints broad and short,
Sorming a sort of broad serrate club ; legs armed with stout spines ;
tarsi 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 shghtly 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,
usually elongate, behind forming a process received into a definite
cavity on the mesosternum ; head retractile, the mouth protected
by the prosternum. Tarsi five-jointed, terminal joint long.

244 COLEOPTERA CHAP.

Although the characters of these Insects are not very different
from those of Byrrhidae, of Dascilidae, and even of certain
Elateridae, there is practically but little difficulty im 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 pecuhar, 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
acuminatus 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 tracheal tube; the body
is terminated by a process from which there can be protruded
bunches of filamentous branchiae. The larvae of Jlacronychus
quadrituberculatus 1s somewhat sunilar, 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 Psephenus
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, without mentioning any names, a detailed account
of Brazilian Parnid larvae, that may perhaps be allied to
Psephenus.

Fam. 49. Derodontidae.—7w7'si jfive-jointed, slender, fourth
joint rather small; front coxae prominent and transversely pro-

} Stettin. ent. Zeit. xlii. 1881, pp. 104-112.

Vv POLYMORPHA—DERODONTIDAE—CIOIDAE 245

longed ; middle coxae small; abdomen with five visible segments,
all mobile, the first not elongated. One of the smallest and least
known of the familes of Coleoptera; it
consists of four or five species of small
Insects of the genera Derodontus and
Peltasticta, found in North America,
Europe, and Japan. The distinction of the
family from Cleridae is by no means cer-
tain; our European Laricobius apparently
possessing characters but 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-
bedded ; abdomen with five ventral seg- ee eee
v pr db merica,
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
hooks curved upwards; that of Cis
mellier (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
dsritain. The Hawaiian Islands have
. =e a remarkably rich and varied fauna of
Fic. 124.—Cis melliet. Martin- Cioidae.
vipa; @ mwas De towimi Fam. 51. Sphindidae—This family
portion of body of larva. of half a dozen species of rare and
coal) small Insects, differs from 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, eg. Reticularia hortensis, that are at first pulpy and

246 COLEOPTERA CHAP.

afterwards become powder. The larvae of both of our British
genera, Sphindus and <Aspidiphorus, have been described by
Perris, 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).— 7 arsi
Jive-jointed, but the first joint very short and imperfectly separated
Jrom 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 differmg very much in appear-
Fic. 125.—Apate capucina, ®2Ce, they have similar habits and larvae.

Europe. A, Larva (after The typical Bostrichides are remarkable

cerris) ; By pertect In- for their variety of sculpture and for the

sect. J i

shapes of the posterior part of the body ;
this part is more or less conspicuously truncate, and furnished
with small prominences. Dinapate wrightii, 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. capucina. Some of the forms (Phonapate) 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.—7a7'si five-jointed, first joint not reduced
in size, often longer than second; front and middle coxae small,
not transversely extended, the former slightly prominent; five
visible ventral segments ; prosternum very short. Here are in-
cluded two sub-families, Ptinides and Anobiides; they are
considered as distinct families by many authors, but in the

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Vv POLYMORPHA—PTINIDAE 247

present imperfect state of knowledge’ 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; Anobiwm (Sito-
drepa) 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

Fic. 126.—“ Biscuit-weevil.”
Anobium paniceum.

Fig. 127. — Early
stages of Anobiwm
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 lke a little hard seed. Anobiuwm
striatum is a common Insect in houses, and makes little round
holes in furniture, which is then said to be “worm-eaten.” <A.
(Xestobium) 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
is standing.
iia) 19) sueere ones eenge The very anomalous genus Letrephes
West Australia. (After (Fig. 128) is found in ants’ nests in
Westwood Australia. Westwood placed it in Pti-
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.—WSeven (or even eight) visible ven-
tral segments, the basal one not co-adapted in form with the coxae ;
tarsi five-jointed. LIntegument softer than usual, the parts of the
body not accurately co-adapted. 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 1t gives is usually conspicuous, frequently much more
so than that of its mate; in other cases the males are the most
brilhant. 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-flies

lg gree ge

ire,

Vv MALACODERMIDAE—FIRE-FLIES—-GLOW-WORMS 249

(Luciola) 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 wtaliea in the Val 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... In our British glow-
worm the female is entirely apter-
ous—hence the name glow-worm Fic. 129.

Phengodes hieronymi. Cor-
; doba, South America. (After Haase.)
—but the male has elytra and A, Male; B, female. 7, Z, Positions

ample wings, and frequently flies of luminous spots ; ls, spiracles. About

. : : x 3.
at night into lighted apartments.

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 hemipterus 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

? Bull. ent. ital. 1886, p- 406, and Ent. Zeit. Stettin, xliii. 1887, pp. 201-206.
Emery does not mention the name of the species, but we 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 Zampyris has given rise
to many contradictory statements; the lhght 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 LZ. noctiluca); the luminescence is, however, most
marked in the female imago, in which it is concentrated near
the extremity of the abdomen; here there are two strata of
cells, and many fine capillary tracheae are scattered through the
luminous substance. Wielowiejski concludes that the hght-
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."
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 Lampyris. 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” 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 Zettschr. wiss. Zool. xxxvii. 1882, p. 354; also Emery, op. cit. xl. 1884,

p- 338. For another theory as to the luminescence, see p. 259.
2 Bull. Soc. Zool. France, xii. 1887, p. 187, postea.

Vv POLYMORPHA—GLOW-WORMS PAS AI

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 *
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) asa perfect Insect ;
he is, indeed, corroborated by Riley.” 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
shghtly 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
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. Willey reeently found in New 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 Ceylonese 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. Zeitschr. xxxii. 1888, pp. 145-167.
2 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 beheved 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 lke perfect Insects as
do the imagos.

The other divisions of Malacodermidae—lLycides, 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 maybe 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 distinctly
co-adapted with the coxvae. ‘These
Insects are extremely numerous,
but have been very little studied.
In many works they are classified
with Malacodermidae, but were
correctly separated by Leconte
and Horn, and this view is also
taken by Dr. Verhoeff, 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
Fic. 130.—Malachius aeneus. Britain. the abdomen and the _ hind

A, Larva (after Perris); B, female 7 oa pA
imago. coxae. The characters are singu-
larly parallel with those of
Silphidae ; but in Melyridae the antennae are filiform or serrate,

Vv POLYMORPHA—MELYRIDAE——-CLERIDAE 2

1
ie)

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. Zygia—they are much firmer. Thus these
Insects establish a transition from the Malacodermidae to ordi-
nary Coleoptera. Although the imagos 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. 150, larva of Malachius aeneus). These are said by Perris
to bear a great resemblance to those of the genus Telephorus,
belonging to the Malacodermidae.

Fam. 56. Cleridae.— TZ arsi jive-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 coaue
prominent, usually contiguous, rather large, but their cavities not
prolonged externally ; labial palpi usually with large hatchet-
shaped terminal joint ; ventral segments five or sia, 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 lttle 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 Z/lus elon-
gatus (Fig. 131) enters the burrows of Pfilinus pectinicornis im
search of the larva. The members of the group Corynetides
frequent animal matter, carcases, bones, ete., 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 Perris has also shown
that the larvae of Necrobia rujficollis really use the puparia formed

1 Larves des Coléoptéres, 1878, p. 208.

bem COLEOPTERA CHAP.

by Diptera. Some of the species of Neerobia have been spread
by commercial intercourse, and V. rujfipes appears to be now one
of the most cosmopolitan of Insects. The beautifully coloured
Corynetes coeruleus is often found in our houses, and is useful, as
it destroys the death-watches (Anobiwm) that are sometimes very
injurious. Zrichodes apiarius, a very lively-coloured red and
blue beetle, destroys the larvae of the honey-bee, and Lampert
has reared Tvrichodes alvearius
from the nests of Chalicodoma
muraria, 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 Z7ri-

chodes ammios recorded — by
Fro 0h gas Meee HOnetee Mayet)! thas Tueech aus

C, termination of the body, more mag- Immature form, destroys Acri-
nified.

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.— Llongate beetles, with soft integu-
ments, front and middle corae exserted, longitudinal in position ;

tarsi slender, five-jointed ; antennae short, serrate, but rather broad.
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 Lymeaylon 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, Soc. ent. France, 1894, p. 7.

Vv 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—Z.
navale and Hylecoetus dermestoides—are now very rarely met
with.

Fam. 58. Dascillidae.—Small or moderate-sized beetles, with
rather flimsy integuments, antennae either serrate, filiform, or
even made flabellate by long appen-
dages ; front coxae elongate, greatly
exserted ; abdomen with five mobile
ventral segments ; tarsi five-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 Parnidae
and Byrrhidae, that are placed in
Clavicornia, than it isto any of the py, 139, Hydrocyphon deflewicollis.
ordinary families of Serricornia. It Britain. A, Larva (after Tournier) ;
is probable that careful study will Bata
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,
and fragile structure. The few larvae known are of three or
four kinds. That of Daseillus 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
Cyphonides 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 Hydrocyphon (Fig. 132, A)

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
Tournier, when alarmed, it enters the water and goes beneath it
for shelter. The third form of larva belongs to the genus Fuci-
netus, it lives on fungoid matter on wood, and has ordinary
antennae of only four joints It is very doubtful whether
Hucinetus is related to other Dascyllidae ; some authorities indeed
place it in Silphidae.

Fam. 59. Rhipiceridae.—T7a7si jfive-jointed, furnished with
a robust onychium (a straight chitinous process bearing hairs)
between the claws ; antennae of
the male bearing long processes,
and sometimes consisting of a
large number of joints. Man-
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 dejeant (Fig. 1335, A) is
described by Schiédte as hard,
cylindrical in form, and peculi-

Fic. 133.—A, Larva of Callirhipis dejeani arly truncate behind, so that
(after Schiddte) ; B, Rhipicera mystacina there appear to be only eloht
male, Australia ; C, under side of its hind ° : 4
Panis abdominal segments, the 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 pectinate, rarely
jiliform. Front coxae small, spherical. Thorax usually with

1 Perris, Ann. Soc. ent. France (2) ix. 1851, p. 48.

v CLICK-BEETLES 25

hind angles more or less prolonged backwards ; with a prosternal
process that can be received in, and usually can move in, a
mesosternal cavity. Hind coxa with a plate, above which the femur
can be received. Visible ventral segments usually five, only the
terminal one being mobile. Tarsi five-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 going off
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! our British Lacon
murinus is provided near the ses ety ale ee pier
tip of the upper side of the ab- — imago.

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 1s difficult to look on this clicking power as of very great value
to the Elateridae, yet their organisation 1s profoundly modified so
as to permit its accomplishment. The junction of the prothorax
with the after-body involves a large number of pieces which are
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

A
2

&

a

1 Arch. Naturgesch. x\viii. 1, 1882, p. 371.
WGI. WAL

TR

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-
sternal 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. V. 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 New World,
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 Pyrophorus, 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. noctilucus is one of the most
abundant and largest of the Pyrophorus, and possesses on each side
of the thorax a round polished space from which lght 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
flying, this central luminous body can be alternately displayed
and concealed by means of shght 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 hfe 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,' who
comes, however, to conclusions as to the physiology of the
luminous processes different from those that have been reached
by Whielowiejski and others in their investigations on Glow-
worms. He considers that the ight is produced by the reactions
of two special substances, luciferase and luciferine. Luciferase
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 “Tes Elatérides lumineux,” Bull. Soc. Zool. France, xi. 1886; also Legons
de Physiologie générale, Paris, 1898, and C.R. Ac. Seti. exxiii. 1896, p. 653.

26

O

COLEOPTERA CHAP.

Schiodte.

Eucnemides,

Leconte and Horn also adopt this view, except that
they treat Throscides as a distinct family.
Throscides, and Cebrionides are all considered dis-

By most authors

tinct families, but at present it is almost impossible to separate

them on satisfactory lines.
and Horn exhibits the characters

imago is concerned :—

Posterior coxae laminate ;

Posterior coxae not laminate ;

Fic.

Labrum concealed ;

The following table from Leconte
of the divisions so far as the

trochanters small.
antennae somewhat distant from
insertion narrowing the front

eyes, their
BESET TIS.

the

Labrum visible, meer antennae arising near ane eyes under the frontal

mar, gin

ELATERIDES.

Tanai transverse, connate w ih the from:

Ventral segments six ; claws simple ;

tibial spurs well developed.

CEBRIONIDES.

Ventral segments five ; claws serrate ; tibial spurs moderate.
PEROTHOPIDES.
trochanters of middle and posterior legs very
long CEROPHYTIDES.
Throseides are

135.

Larva of Fornax
n. sp. Hawaii. A, Upper
side; B, under side: ss,
position of spiracles; C,
head more enlarged; D,
under side of terminal seg-
ment ; @, anus.

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 7hroscus 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-lke, man-

dibles. The larvae of Eucnemides (Fig.
135) are very little known, but are
highly remarkable, inasmuch as it is

very difficult to find any mouth-opening
in some of them, and they have no legs.
The other divisions possess very few species
compared with Elaterides. In Britain
we have about sixty species of Elate-

rides, four of Throscides and three of Eucnemides; Cerophytum

was probably a native many years

ago. Neither Perothopides

Vv POLYMORPHA—BUPRESTIDAE 261

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 the second, the others mobile. Tarsi
Jive-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
briluant 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

Fia. 186.—A, Larva of Euchroma
metal. The dorsal plates of the abdo- goliath (after Schiddte) ; B, imago
men are usually soft and colourless in of Melanophila decostigma.

: : Europe.

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 per cent 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

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 Hthon dwell in
galls on various plants. Buprestid larvae are of very remarkable
shape, the small head being almost entirely withdrawn into the
very broad thorax, while the abdomen is slender.t 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, those 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 five.
Several of the families of the series are of doubtful validity ;
indeed beyond that of Tenebrionidae 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
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
lees; 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 Tt 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 Westwood (Mod. Classif. i. 1839, p. 229), and figured as probably
Buprestis attenwata, is doubtless a Passalid.

v HETEROMERA 263

Fam. 62. Tenebrionidae.— Front coxae short, not projecting
from the cavities, enclosed behind. Feet destitute of lobed joints.
Claws 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
used by the great majority of the
terrestrial forms. Many of the Fis. 187.—Tenebrio molitor. Burope, ete. A,

. {es Larva (meal- worm); B, pupa (after
wood - feeding Tenebrionidae, Schiddte); C, imago.
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.—dAnn. New York Ac. v. 1890, p. 416.

In Eleodes, 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 1s 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—
we. the larva of Tenebrio molitor—ais 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 closed, tips of
the front coxae free, claws smooth, penultimate joint of the tarsi
broader, pubescent 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, Lagria 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.

Vv HETEROMERA 265

Fam. 65. Othniidae.—Onlyabout ten species are knownof 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 Othnius have been found in Japan and Borneo, as
well as in North America. Nothing is known as to their
metamorphoses.

Fam. 66. Asgialitidae.— All the coxae very widely separated ;
no co-adaptation between the sides of the abdomen and the edges of
the wing-cases ; five ventral segments and tip of a siath 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 ina 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 inflexed
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 Rhinosimus have the head prolonged in
front of the antennae so as to form a beak. The larva of Pytho
depressus 1s 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. Pyrochroa rubens is
common in some parts of England; the larva is found under
the bark of tree-stuinps; 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—AHead with an abrupt narrow neck ;
prothorax narrower than the elytra. Middle and hind coxae placed
in definite acetabula. Claws simple. 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 without a narrow neck. | Penultimate tarsal
joint broad ; claws 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. | Nacerdes melanura is common on our

Vv TIETEROMERA—MORDELLIDAE ~ ZO7

coasts, where its larva lives in timber cast up by the sea, or
brought down by floods, 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 Ascelera
caerulea there are six dorsal and
three ventral pairs of these re-
markable pseudopods. We have
six species of Oedemeridae in
Britain, including <Asclera as Py tele cn dealion. me eaten
well as Nuacerdes. B, pupa (after Schiddte) ; C, imago.
Fam. 74. Mordellidae (incl. 9 “™""s*
Rhipiphoridae)— Head peculiarly formed, vertex lobed or vidged
behind, so that in extension it reposes on the front edge of the pro-
notum ; capable of great inflection and then covering the prosternum ;
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 Rhipiphoridae
cannot at present be satisfactorily distinguished from Mordeilidae.
Leconte and Horn separate the two by the fact that the sides
of the prothorax form a sharp edge in Mordellidae, but not in
Rhipiphoridae. 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. 139, 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 Rhipiphorids, of the groups Myoditini and Rhipidini,
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
Anaspini swarm on the flowers of bushes and Umbelliferae. The

268 COLEOPTERA CHAP.

life-histories appear to be singularly varied; but 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 JMordellistena floridensis.
Coquillett has found the larvae of JZ pustulata 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
Mordellistena, and fed voraciously on Dipterous larvae in the
stems of a plant. The little that is known as to the meta-

Fia. 189. — Mordelli-
stena floridensis.
America. (After
Riley.) A, Larva;
B, pupa; C, imago ;
D, outline of de-
tached head of im-
ago of MW. pumila,
to show the neck.

morphoses of Mordella and Anaspis shows that they live in old
wood, but does not make clear the nature of their food.
Although it has been ascertained that the Rhipiphorides
exhibit instances of remarkable metamorphosis, their life-
histories are still very imperfectly known. Dr. Chapman has
ascertained some particulars as to MJetoecus paradoxus, which has
long been known to prey in the larval state on the larvae of the
common social wasps." 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 wasps’ 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. Hist. (4) vi. 1870, p. 314; and Ent. Mag. xxvii. 1891, p. 18.

V HETEROMERA—CANTHARIDAE 269

access is gained to the cells the little J/etoecus 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 Jetoecus, in the cell of the wasp. The
wasps, though they investigate the cells, do not apparently entertain
any objection to the JJetoecus, though there may be sometimes as
many as twenty or thirty of the destroyers in a single nest. <A
few hours after the J/etoecus has become a winged Insect and has
escaped from the cells, it appears however, from the observations
of Erné? 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. Lmenadia
flabellata, 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.2, An old record to the
effect that a second species of Hmenadia, E. bimaculata, lives in
the stalks of Kryngiuwm campestre, on the pith, is now thought to
be erroneous. Fabre has found the larvae and pupae of another
Rhipiphorid in the cells of a bee, Halictus sexcinctus.

The most remarkable of the Rhipiphorids, from the point of
view of its habits, is certainly Symbius 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 coming from India; the male has
been met with in several European countries, but the female is
excessively rare.

Fam. 75. Cantharidae or Meloidae (A/ister-beetles, Oil-beeties).
—Head with an abrupt neck ; elytra and sides of the abdomen with-
out any coadaptation ; each claw of the feet with a long appendage
closely 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. Schweiz. ent. Ges. iv. 1876, p. 556. ” Ann. Soc. ent. France, lx. 1891, p. 447.

270 COLEOPTERA CHAP.

and appled 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 pre-
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 1851,' and the subject
has since been greatly advanced by Fabre, Riley, and others.
As an example of these peculiar histories, we may cite Riley’s
account? of Hpicauta vittata (Fig. 140), a blister-beetle living
at the expense of North American locusts of the genus Calop-
tenus. The locust lays its eggs underground, in masses sur-
rounded by an irregular capsule, and the Hpicauta 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 of 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
this the triungulin at once eats its way, and commences to devour
an egg. Should two or more triungulins enter the same ege-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. Linn. Soc. xx. 1851, p. 297 ; and xxi. 1853, p. 167.
2 Rep. U.S. ent. Commission, i. 1878, p. 297.

V HETEROMERA——-CANTHARIDAE 2

NI
H

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, but is not accompanied by
any very great change of form, though 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

Fic. 140.—Hypermetamorphosis of Hpicauta vittata. North America. (After Riley.)
A, Young larva or triungulin ; B, Caraboid instar 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
hes 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
reduced size and greater whiteness. Itis 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). Itis 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 (/oc. cit.) this series of instars, Riley
changed his views as to their nomenclature.’ The following
summary of the metamorphosis, to which we have added the two
nomenclatures of Riley—the original one, when different from the
amended one, being given in square brackets—may therefore be
useful, viz—Ege ; 1, triungulin-larva—moult; 2, Caraboid larva
[second larva, Caraboid stage]—moult; 3, Scarabaeoid larva
[second larva, Scarabaeoid stage|—moult; 4, Scarabaeoid larva
[second larva, ultimate stage] (large amount of food and much
erowth)—moult ; 5, coarctate larva [pseudo-pupa, or semipupa] ;
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
humeralis, 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. V. p. 159 of this work.

is CANTHARIDAE 273

to any hairy object that may come near them, and thus a certain
number of them get on to the bodies of the Anthophora 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 ghdes 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 ving 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.
VOL. VI T

274 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 1s 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
melittae,’ that was believed by Kirby to be a sort of bee-louse.
Some species of the genus J/eloe are still common in Britain, and
the Insects may be 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 Anthophora, and its life-history seems on 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 Anthophora nest. It
would be supposed that it would be a much better course for these
bee-frequenting triungulins to act like those of Hpicauta, 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 uninstructed, inexperienced atom, is such that we
should @ priori have denounced it as an impossible means of
existence, were it not shown that it 1s constantly successful. It
is no wonder that the female JJ/eloe 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 J/e/oe 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
Meloe as a living Insect is.

The histories of several other Cantharids have been more or
less completely discovered. Fabre has found the larva of
Cerocoma schaefferi attacking the stores of provisions laid up by
a fossorial wasp of the genus TYachytes, 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.t

Some half-dozen species of the genus Cephaloon 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 Cantharide.

Fam. 76. Trictenotomidae.— Large Heteromera, with powerful
Sree projecting mandibles ; the antennae long, but with the terminal
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 than
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 Insectes Vésicants, Paris 1890, 554 pp. Parts of this work were pre-
viously published in J. de ? Anat. Phys., xxi. xxii. xxiii. 1886 and 1887.

iS)
N
O”’

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 wpper surface so as to allow of the 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 completely) occluded in the
mouth.

This great series of beetles includes something lke 35,000
species. It approaches, like all the other series, the Polymorpha,
especially the family Erotylidae placed therein, but 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 Rhynchophora, 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 stein-feeders. The number of
exceptions to this rule is but small, though certain Cerambycidae
and certain Chrysomelidae live on roots.

Fam. 77. Bruchidae.—Prosternum eatremely short ; in front
perpendicular ; behind the coxae, forming merely a transverse
lamina with pointed extremity. Hind femora more or less

Vv PHY TOPHAGA——BRUCHIDAE 207

thickened. This comparatively small family includes about
700 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
Lruchus pisi and B. fabae have, when first hatched, three pairs
of legs which are subsequently lost. They also have peculiar

S pas
SR WC
Pt 0 Fic. 141. — Bruchus
ck pist or pea-weevil.
=x A, Young larva ;
B, prothoracic spin-
ous process; C,
post-embryonic
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. oth 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 lfe-histories of other Bruchidae it 1s probable that
still greater variety prevails. Heeger has found that bruchus
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 comes to another pod into which it
bites its way.

The family has, until recently, been placed in the Rhyncho-

COLEOPTERA CHAP.

1)
Sy
ee)

phorous series, with which it has, however, no direct connection.
On the other hand, it is so closely connected with Chrysomehdae
that it is not possible to indicate good characters to distinguish
the two at present. The Australian genus Carpophaqus, and the
large South American species of Caryoborus 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 Uvodon appears to belong to Anthribidae, not to
Bruchidae. The family Bruchidae is called Mylabridae by some.
Fam. 78. Chrysomelidae.-—<Antennae moderately long ; eyes
moderately large, usually not at all surrounding the insertion of the
antennae; upper surface usually bare, frequently brightly coloured
and 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
Chrysomelidae from Cerambycidae has
yet been discovered, 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. Thepotatobeetle,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
Me 149. Dorjhone ae turnip flea, a tiny hopping beetle, is
lineata, the potato beetle, among the smallest forms of the family,
a Aenean ete and is a member of another very exten-
joint ; 4, true fourth joint; sive subdivision of Chrysomelidae, viz.
Oe eee en 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. :—

Vv PHY TOPHAGA—-CHRYSOMELIDAE 279

Prothorax much narrower at the base than the elytra, and usually with-
out side-margins (raised edges). Sub-fam. 1. Europa; with three divisions,
Sagrides, Donaciides, Criocerides.

The basal ventral plates of the abdominal segments are somewhat shorter
in the middle than at the sides, the fourth one being often invisible in the
middle, while the fifth is very large. Sub-fam. 2. CamprosomEs ; 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. Cycnica ; with four divi-
sions, Lamprosomides, 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
part 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
of 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,’
in S1X groups, Viz. :

1. Elongate larvae, living under water, and there undergoing their meta-

morphosis. (Donaciides.)

Larvae mining in leaves, and undergoing their metamorphosis in the

leaf. (Hispides and some Halticides.)

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.)

5. Peculiar larvae of short form, spiny, and protecting their bodies by
excrementitious matter attached by a special apparatus, the excrement
itself being moditied so as to be suitable for retention. (Cassidides.)

6. Elongate, pallid, larvae with curved abdomen ; living in shell-like cases,
and undergoing metamorphosis therein. (Most of the Camptosomes,
the habits of which are known.)

bo

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 lives in
the earth feeding on roots after the manner of a Rhizotrogus
larva, which it resembles. The larva of Sagra splendida lives

1 Genera des Coléoptéres (Suites a Buffon), 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.

1. Europa. The beetles of the genus Donacia are of special
interest. They form, with the genus Haemonia, a peculiar group,
well 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 aérial 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 on 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 1t 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 ' 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 #. curtist at Cley on
the Norfolk coast on submerged Potamogeton 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 Berlin. ent. Zeit.:1887, p. 225, and 1889, p. 299.

Vv PHYTOPHAGA——_CHRYSOMELIDAE 281

body is pecuharly shaped conformity with the curious habit
of using the excrement as covering. The larva is less elon-
gate than usual, anu has tue 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 Lema 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 hlies—C. merdigera,
é.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
egos 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-
lating organ. Two contiguous
areas at the base of the last C
dorsal segment, where they can
be rubbed by. the tips of the Fic. 143.—Crioceris asparagi. A, Eggs

. ¥ 3 in position on stem of asparagus ; B,
elytra, are slightly elevated and oie egg much enlarged; ©, young
bear very close and fine straight larva. Cambridge.
lines.

ii. The CAMpPTosoMEs, as we have already noticed, are distin-
euished 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-

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 Rhynchophora. 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 Cerambycidee.

i. The division Cycuica includes the great majority of Chryo-
melidae; we hav@ 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 egos 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. Hispa 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 Perris,’ 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. Liége, x. 1855, p. 260.

V PHY TOPHAGA——-CHRYSOMELIDAE 2

[oe]
Ww

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, always
makes use of two leaves, and of two opposed leaves. A know-
ledge of the habits of some of the larger of the exotic Hispides
would be of much interest.

Fic. 144.—Pupa of
Cassidid beetle
(? Aspidomorpha
S5]05)) Soe eee V aKUL
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
palmarum, the larva of which has, in place of the usual fork, a
more complex appendage on the back 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 Aspidomorpha—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 skins 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 Porphyraspis. JP. tristis 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
are 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,
Fro, 145.—Nest of intes- and make an even surface; but the outside

eae rei fibres stand out in a somewhat bushy
of Porphyraspis tristis fashion. The construction is much lke that
ee of a tiny bird’s nest. Sefior Lacerda informed
the writer that the larva makes a nest as soon as it 1s hatched.
Another Porphyraspis—P. palmarum—has been recorded as

Vv PHY TOPHAGA——CERAMBYCIDAE 28

forming similar nests on a species of Thrinaxz in St. Domingo.
Candéze says! that when it has 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 frequently ren-
dered dull by a very minute hairiness, which often forms a pattern ;
antennae usually long, and their insertion much embraced by the
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, whitish 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-
bycidae and those of Buprestidae, and when the thorax of a
Longicorn larva is unusualiy broad, e.g. Asiynomus, this similarity
is very pronounced.

Fic. 146.—Saperda popul-
ned. Britain.

1 Mem. Soc. Liéye, xvi. 1861, p. 387.

286 COLEOPTERA CHAP.

The modes of iife of Cerainbycid larvae exhibit considerable
variety, and much perfection of instinct is aisplayed by the larvae,
as well as by the mother beetles. ‘he larvae of Saperda populnea
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 aftlux of
sap to the spot where they are situated. Haphidion villoswm is
called the oak-pruner in North America. The parent beetle lays
an egg near the axilla of a leaf-stalk or small stem, and the
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 pabulum
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
erowth 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 lite of a Longicorn beetle extended over at least forty-five
years.’ 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. O. Waterhouse had for some years a Longicorn larva under
observation, feeding in this way in the wood of a boot-tree >2 ihe
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. 1890, p. 689.
2 Not a growing tree, but the instrument used for stretching boots.

.
|

Vv 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' has noticed that species of the genus Odontocera
resemble various Hymenoptera, one species being called 0.
braconoides; he also observed that these Hymenoptera - like
Longicorns, instead of withdrawing their underwings under the
elytra as beetles generally do, vibrate them rapidly ike Hymen-
optera. A large number of Longicorns stridulate loudly by rubbing
a ridge inside the pronotum on a highly specialised, striate surface
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
Hawanan genus Plagithmysus produce sound in both these
manners, the thoracic stridulating organ being beautifully de-
veloped, while in some species the margin of the elytra and
base of the femora are also well adapted for the purpose of sound-
production, and in a few species of the genus there are also
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-fam. 1. PRIONIDEs.

2. Front coxaé not greatly extended transversely, thorax not. margined ;
last joint of maxillary palpus not pointed, usually broader (more or. less)
than the preceding joint. Sub-fam. 2. CERAMBYCIDES.

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
on the inner side. Sub-fam. 3. LamipeEs.

The Prionides are on the average considerably larger in size
than the members of the other divisions, and they include some ot
the largest of Insects. The Amazonian 7%tanus giganteus and the
Fijian Macrotoma heros are amongst the most gigantic. Some
of 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. Zeitschr. xli. 1896, 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.‘ 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 Lamuides are the most highly speciahsed 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
inflexed, 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 beak, called
rostrum. Tarsi four-jointed, usually at least the third
joint broad and densely pubescent beneath.

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 1n
which the beak is indistinct, and others in which the tarsi are

1 Sharp, Ann. Soc. ent. Belgique, xxviii. 1884, CR. p. evii.

‘cases it 1s also used to push the

Vv 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
be 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

ege far into the hole in which it
has previously been placed by
the ovipositor; but there are
many forms in which it 1s fairly
certain that it is not so used.
What purpose it serves in the
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
Maat I HO OEE ao OG ie ee Aga ee
this interpretation ) and there of pronotum, head, and rostrum.
seems to be no reason whatever
for considering the Rhynchophora 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 Rhynchophora can be 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, but dis-
tinct and flexible. Antennae often long, not elbowed, the first joint
not very lony. Third joint of tarsus small, usually much concealed
by being embraced by the second joint. Pygidium exposed ; pro-
pygidium deeply grooved in the middle. 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

f Insect makes burrows in dead branches
Bia. 148.— Platyrhanus lati- of hawthorn. The larvae of the genus
rostris, Anthribidae. Britain. =

A, the perfect Insect; B, Brachytarsus have been ascertained te

tarsus and tip of tibia. prey on Coccidae.

Fam. 81. Curculionidae (/Vcevils)—The beak of very variable
length and thickness; the palpi small, nearly always concealed
within the mouth, short, and rigid. Labrum absent. Antennae
of the majority elbowed, ie. with the basal joint longer, and so
formed that when it is laterally extended the other joints can be
placed in a forward direction, This enormous family includes

V RHYNCHOPHORA—CURCULIONIDAE ZOT

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 Amycterides, eg.) 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. No 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 Rhynchites (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

Z2OQ2 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 $, 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 §-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 funnel-

}

Fic, 149.—The leaf-rolling of Rhynchites betulae. Britain. A, Female beetle, magnified ;
B, the beetle forming the first incision on a leaf; C, the completed roll. (Band C
after Debey.)

twister now 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

’

i
ij

Vv RHY NCHOPHORA—CURCULIONIDAE 293

to the roll, and with the other legs drags to it the portion of the
leat on the other side of the midrib so as to wrap this part (7.
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.?

Fam. 82. Scolytidae.— Rostrum extremely short, broad ; tibiae
frequently denticulate externally ; antennae short, with a broad
elub. This family is not at all sharply distinguished from certain
groups of Curculionidae (from Cossonides 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 more extensive account of Rhynchites betulae and others refer to
Wasmann. Der Trichterwickler, Miinster, 1884, and Debey, Beitrage zur Lebens-
und Entwickelungs-geschichte . .. der Attelabiden, Bonn, 1846. The first in-
cludes an extensive philosophical discussion ; the second is a valuable collection
of observations.

V hkKHYNCHOPHORA——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 jot 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,* 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.”

Fam. 83. Brenthidae.— orm elongate; rostrum straight,
directly continuing the long axis of the body, often so thick as to
form an elongate head ; antennae not elbowed. The Brenthidae
form a family of about 800 species, remarkable for the excessive

1 Bull. U.S. Dep. Agric. ent. New series, No. 7, 1897.
? Perris, Ann. Sci. Nat. (2) xiv. 1840, p. 89, pl. iii.

2g6 COLEOPTERA BY

leneth and slenderness of some of its forms, and for the
extreme difference in the sexes that frequently exists. It is
well represented in the tropics
only, and very little is known
as to the natural history and
development. These beetles are
stated to be wood-feeders, and
no doubt this 1s correct in the
case of the majority of the
species; but Mr. Lewis observed
in Japan that Zemioses celtis and
Cyphagogus segnipes are pre-
daceous, and enter the burrows
Fie. 150. — Lupsalis minuta. North ob wood-boring Insects to search
America. A, Larva; B, pupa; ¢, for larvae as prey: they are
ey Bhp Sebel OT nas (CNAEE very much modified in 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 1s known as to the larval history of these carnivorous
forms. Indeed an uncertainty, almost complete, prevails as to
the early stages of this family. Riley has given a sketch of a
larva which he had no doubt was that of Hupsalis minuta, the
North American representative of the family; if he is correct
the larva differs from those of Curculionidae by its elongate form,

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 an 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

Vv BRENT HIDAE—AGLYCYDERIDAE ZO 7,

the body, although this itself is so drawn out as to be quite
exceptional in the Insect world:' 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.” The males of some species
fight; they do not, however, wound their opponent, but merely
frighten him away. In Zupsalis 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 fulerum 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
barbicornis, 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. — 7Z'arsi 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. West-
wood considered the tarsi to be really four-jointed, but it is not

* 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 Brenthus anchorago—ate five times as long as others.

2908 COLEOPTERA CHAP.

at all clear that the minute knot he considered the third joint is
more than the articulation
of the elongate terminal
joimt. 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 New
Caledonia. The former is
believed to live in the stems
of Huphorbia canariensis ;
Fic. 151.—Aglycyderes setifer. Canary Islands. a New Zealand species has

A, Imago 7 B, tarsus according to Westwood Sls Pop fn eemeegan

C, according to nature; D, maxilla; E,

labium. with the tree-fern Cyathea

dealbata.

Fam. 85. Proterhinidae.— 7rsi three-jointed, the second joint

lobed ; head of the male scarcely prolonged, but that of the female

RNS BE
SREERET AAT Mz

2 2eSSSS

Jorming a definite rostrum,; maxillae and ligula entirely covered
by the mentum. 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 the Fie. 152.—Proterhinus lecontei. Ha-
Hawaiian’ Islands, where these Imeyiiaie sere ee pee
Z C, front foot, more magnified.
sects live on dead wood in the
native forests. The genus is numerous in species and individuals.
Strepsiptera (or Rhipiptera, Stylopidae).— Male 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, without 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

V SLREPSIPTERA 299

other Insécts, 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 lke that of MJeloe; 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

+. 153.—Sexes of Strep-
siptera. A, Male of
Stylops dalii (after Cur-
tis); B, female of Yenos
rossvi (after von Siebold).

Strepsipteron is found only in the imago of the host. 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.)
When 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.

300 '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 Git being a
species of the genus Sty/ops 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 very
different from Stylops 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-
Fic. 154.—Young larva ever, that they vary much according to the
of Stylops on a bee's- : : : 7 »
hair. Greatly magni- Species, and also according to the extent of
fied. (After Newport.) the stylopisation. Usually only one Stylops
is developed in a bee; but two, three, and
even four have been observed:’ and in the case of the wasp,
Polistes, Hubbard has observed that a single individual may
bear eight or ten individuals of its Strepsipteron (enos,
n. sp. 2).
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

+ 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
stated that female bees produce more especially female Stylops, and male bees
male Stylops. If any correlation as to this latter point exist, it is far from general.

Vv 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 Sty/ops 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

1843.1 Before that time the

mature female was supposed to be

a larva, and the triungulins found

in it to be parasites. Although a B

the erroneous character of these pie, 155.—Portion of early stages of

views has been made clear, the Xenos rossti. (After von Siebold.)
A, Small male larva ; B, small female

problems that have been sug- — jarva: ¢, Pieces trvace D,

gested present ereat difticulties. full-grown female larva; E, the so-

called ‘‘ cephalothorax ” and adjacent
Apparently the change from the — segmentof adult female. (The newly-

triungulin condition (Fig. 154) to pees ae a 4S Sg ie a
as : Ce of Stylops shown in Fig. 154.

the parasitic larvae (Fig. 155, A, B)

is extremely great and abrupt, and it appears also that during

t=)

Von Siebold, Arch. Naturges. ix. 1843, pp. 137-161. Nassonoff’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.

302 SME RSL TERA 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. Von
Siebold apparently entertained no doubt as to the part of the
female that is extruded being the anterior extremity ; indeed he
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' 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 ganghon, 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?

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, Andrena, are concealed under ground ; and
in the case of the wasps, Polistes, 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. Meddel. v. 1896. p. 148, and Ov. Danske Selsk. 1896, p. 67.
2 Horae Soc. ent. Ross. xiv. 1879, p. 14.

Vv SL OPS: 303

metamorphosis. They have been observed in several species of
Homoptera; and the writer has a large Pentatomid bug of
the genus Callidea, which bears a female
Strepslpteron apparently of large size. This
bug’ 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
Nassonoff think parthenogenesis occurs in
Strepsiptera, but there appear to be no facts tee Stee one
to warrant this supposition. Von Siebold — raeus) with a Strep-
:, 1 . sipteron (Xenos 9) in
speaks of the phenomena of Strepsipterous — josition, one of the
reproduction as paedogenesis, or pseudo- dorsal plates of the
paedogenesis, but we must agree with Re ir
Meinert that they cannot be so classed. tion of part of the
. : : parasite ; 6, line in-

The males of Strepsiptera live for only a dicating the position
very short time, and are very difficult of ces EEE
observation. According to Hubbard the ee
males of Yenos 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 Yenos, 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.

' Named by Mr. Distant Callidea bare ; according to the Brussels catalogue of
Hemiptera, Chrysocoris grandis var. baro.

CHAPTER Vi

LEPIDOPTERA—OR BUTTERFLIES AND MOTHS

Order VI. Lepidoptera.

Wings four; body and wings covered with scales usually varie-
gate in colour, and on the body frequently more or less like
hair: nervures moderate in number, at the periphery of
one wing not exceeding fifteen, but little irregular; cross-
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 body; the larva
with large or moderate head and strong mandibles. Pupa
with the appendages usually adpressed and cemented to the
body so that it presents a more 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 1s
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. Vemognatha.

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. Asa 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,

Fie, 157.—Metamorphosis of a Lepidopteron (Rhegmatophila alpina, Notodontidae).
(After Poujade, Ann. Soc. ent. 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 7; F, male moth in repose ; G, female moth in repose.

Aphidae) and a very small number (Tinea, ete.) 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 ba

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 be
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 (Hphestia
kuehniella, 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: Réaumur 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

rete

VI EXTERNAL STRUETURE 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 Lepidopteron 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-

Fic. 158.—External structure of a female butterfly, Anosia plexippus. (After Scudder.)
a, Base of antenna ; 6, pronotum ; 47, seutum of mesothorax ; ¢, clypeus ; ca, coxa ;
d, scutellum ; d1, scutellum of metathorax ; ¢, post-scutellum (=base of phragma) ;
em, epimeron ; ep, episternum ; 7, scutum of metathorax ; m, basal part of pro-
boscis (=maxilla); 0, eye; p,; labial palp; 7, mesosternum; s, prothoracic
spiracle ; 7, tegula ; ¢r, trochanter ; 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, Castnia) 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 Bombyces and Psychidae; 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 labrum is a very short,

1 Kellogg, Kansas Quarterly, ii. 1893, p. 51, plate II.

oD?

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

Fia. 159.—Mouth of Lepidoptera. Tiger-moth, Arctia caja. A, Seen from front ; B,
from front and below. a, Clypeus; 4, labrum; ¢, epipharynx; d, mandibular
area; d@’, prominence beneath mandibular area; e, one side of haustellum or pro-
boscis ; 7, maxillary palp; g, labial palp.

is not the case’ The mandibles are usually indistinguish-
able, though they, or some prominence possibly connected with
them,” may frequently be detected in the neighbourhood of
the pilifers; they are, according to Walter, largest and most
perfectly developed in Eriocephala, a genus that was not dis-
tinguished by him from Micropteryx and was therefore termed
“niedere Micropteryginen,’ 12¢. lower Micropteryges. The
opinion entertained by Walter that Jicropteryx proper (his
“hohere Micropteryginen”’) also possesses rudimentary mandibles
is considered by Dr. Chapman, no doubt with reason, to be
erroneous.” 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. Zeitschr. Naturw. xviii. 1885, p. 751.

2 The writer is not quite convinced that the supposed mandibles of these Macro-

lepidoptera are really entitled to be considered as such.
3 Tr. ent. Soc. London, 1893, p. 263.

;
q

V1 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 populi, 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 MAxILLAk 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 Hepialus 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 Amer. Natural. xxix. 1895, p. 637. It shouid be recollected that many
Lepidoptera do not possess any proboscis.

5 ILO) 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.'

Lasium.—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. Zeitschr. Naturw. xviii. 1885, p. 168.

Ng mee i ee eh ra tet i eer eee = «>

VI EXTERNAL STRUCTURE 311

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 Miller’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.’ The canal
formed by each maxilla opens into a cavity inside the front part
of the head. This cavity, according to Burgess,’ 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* 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-scutum, and a

1 Amer. Natural. 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. Mier. Sci. 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.
Zeitschr. Naturw. xv. 1882, p. 151.

3 See Cholodkovsky, Zool. Anz. ix. p. 615; Haase, t.c. p. 711; also Riley,
P. ent. Soc. Washington, ii. 1892, p. 310.

ale : LEPIDOPTERA CHAP.

smaller part, the meso-scutellum behind. In front of the seutum
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 1s 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
mesothorax 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 mesothorax, 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 mesothorax
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 MrrarHorax 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-

¥
F.
‘
#

VI EXTERNAL STRUCTURE a

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 1t 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 mesoscutellum 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! a
structure behind the anterior wing, and thinks it may be an
imperfect spiracle, and we have found a similar stigma in
Saturnia pavona. At the back of the thorax there is on each
side in some Lepidoptera (Noctuidae, Arctia, ete.), 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 Peytoureau,” 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,’ 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. Sci. Paris, exviii. 1894, p. 360; and his Thesis, Bordeaux, 1895.
3 O.R. Ac. Set. Paris, exviii. 1894, p. 542.

314 IE PID © PARE RA 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
“uneus”; 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, /.c.). On
each side of the ninth abdominal segment there is a process called
the “ valve,’ the internal wall of which bears some hook-hke 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 _ be
covered externally with
scales like the rest of
the abdomen. Peytou-
reau considers that the
uncus is really the dorsal
plate ofa tenth segment,

and that the scaphium

Fic. 160.—Acherontia atropos. The termination of 6 jg the tenth ventral
body, one side removed. JX, Ninth dorsal plate ; = ee
TX’, ninth ventral; s, lobe, saccus, of ninth plate. Thus, according
ventral plate; Y, tenth dorsal plate, or uncus; to this view. the ninth
sc, scaphium, or tenth ventral plate ; @, position of ; : :
anus ; 4, chitinised band of scaphium ; JV, valve or segment is extensive and

nye oa ae aeetege i Pee leonipl ae Heme

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.

Lrcs.—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,

Ce

P= ee eee ee ee

vI WINGS Bis

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 J/orpho, 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 aérial 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 sinall 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 Castnia (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 (7). 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-
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 jugum 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
humerous 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 nervures
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 WING-NERVURES a7

another, and it is probable that they function almost as four
separate wings instead of as two pairs.

WING-NERVURES.—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, IT. B).
The maximum number of closed cells is six; this is found in
some species of Micropteryz, 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, 1b). This important condition
has not yet been adequately discussed.

Turning to the mode of designation of the nervures,! 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
prefatory 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. Washington,
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

ro-

results. See Spuler, Zeitschr. wiss. Zool. liii. 1892, p. 597.

318 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 161 will enable us to
explain the methods actually in vogue ; I. representing the system,
dating from the time of MHerrich-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

Mio m2

Fic. 161.—Wing-nervuration of Lepidoptera. I, Diagram of moths’ wings (after Hamp-
son) ; II, of a butterfly’s wings (Morpho menelaus 6, after Staudinger and Schatz).
A, front, B, hind wing. I.—c, costal ; sc, subcostal ; m, median ; la, 10, 1c, in-
ternal nervures; /f, 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. IJ.—C, costal ; SC, subcostal ; M, median ;
SM and SN, submedian nervures ; 1A, inner-margin nervure ; UR, lower radial ;
OR, upper radial ; SC! to SC®, divisions of subcostal ; M! to M®, divisions of median
nervure ; 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

eh ND MOD SP. om

eS te Te

v 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,b, e 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. eae Pine Ge SNe

The normal number of nervules B, hind wings. 1a, 1, le, 1d,
is, Gul the. front wing, 11+ 1 or 2 Inner marginal nervures ; 2, lower

branch of median ; 8, subcostal
inner marginal, and on the hind of hind wing; 12, subcostal of
ere Seon, aa: é front wing; ¢, “stalk” of 8 and
pee OL oe olier marcinal. Im- 9 ¢ fenalum: @ retmmcnlam:
the aberrant moths of the genus Ԥ, shoulder; g, articulation of
- 5 a wing.
Castnia the nervuration is unusually -
complex and irregular (Fig. 162), and an analogous condition
occurs in our common Goat-moth (Cossus ligniperda). In
Hepialus 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.’
Internal Anatomy.’—The alimentary canal extends as a long,

1 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, ete.

? The internal anatomy of Lepidoptera has not been extensively studied. For
information refer to Dufour, C.R. Ac. Paris, xxxiv. 1852, p. 748; Seudder, Putt.
New England, i. 1889, p. 47 ; Minot and Burgess, Fourth Rep. U. S. Entom. Comm.
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 Mal-
pighian 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

ex3 cx? LZ
Jipatie

Fic. 163.-—Internal anatomy of Lepidoptera. Section of the body of a female buttertly,
Anosia plexippus. (Atter Scudder.) The portion to the left of the vertical line*
more magnified. I. II. III. thoracic segments; 1-9, abdominal segments; 4a,
antenna; @, anus; ac, aortal chamber ; ag’, etc., abdominal ganglia; agl', agi’,
accessory glands ; ao, aorta ; br, brain; ¢, colon; cp, copulatory pouch ; cz), ca,
cx, coxae ; fr, food-reservoir ; g!, suboesophageal ganglion ; h, dorsal vessel ; @,
intestine ; Jim, area filled by wing muscles ; /.or, ovary, or egg-tubes of left side ;
mv, Malpighian tube (the two others of the right side cut away, except small por-
tions) ; mx, maxilla; 0, oviduct; 00, its orifice ; oe, oesophagus ; ov.c, end of left
ovary ; p, labial palp ; ph, pharynx ; 7.0v, terminal parts of right ovarian tubes,
turned to one side, after the tubes have been cut away ; sd, salivary duct; sgl,
salivary gland ; sp, spermatheca ; sf, stomach; ¢g, thoracic ganglia ; v, 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 Beh

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 ege-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,’ 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, the spermatheca in Pronuba contains some
curious radiate bodies, and Godman and Salvin 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—hbeing 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. V.) the egg is a

1 Tr. Linn. Soc. London (2), v. 1890, p- 143.
2 P. ent. Soc. Washington, ii. 1892, p. 305.
VOLE ve

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, Vol. V.). The egg-shell or chorion is, aceord-
ing to Korschelt ' 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 egos deposited by an individual. The number in the case of
Aporia crataegi is beheved to be from 60 to 100, and in some
Hepialus 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 neustria 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
Vanessae. Some moths cover the eggs with
fur from their own body, which, in the case
of certain of the Eggers (Lasiocampidae),
ie ee a ee seems to have a special supply for the pur-
cardamines, magni- pose. The period that intervenes between
pet atin ny eae Ceposition and hatching of the eggs varies
from a few days to many months. There
seems to be, asa 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
worm-like creature—in old English it was sometimes called

1 Acta Ac. German. li. 1887, p. 238.

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 lke 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 ee 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 give rise to the legs of the imago. The number
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,’ 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, yp. 384-404, Pl. 7. 7 isis, 18455. sede

324 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. The mouth-parts are very different from those of the

Fic. 165.—A, Section of male caterpillar of Anosia plexippus—muscular and tracheal
systems and fat-body not shown: I, pro-, II, meso-, III, meta-thorax ; 1-10,
abdominal segments ; 6, supra-oesophageal ganglion ; c, rectum ; d.v, dorsal vessel ;
g>-g°, ganglia of ventral chain ; h, head ; 7, intestine ; m.v, Malpighian tube ; 2.c,
nerve-cord of ventral chain ; oe, oesophagus ; s, spinneret ; s.o.g, infra-~oesophageal
ganglion ; st, stomach ; s.v, silk-vessel; ¢, testis. B, One of the jointed prothoracic
legs. ©, An abdominal foot with its hooks. (After Scudder and Burgess, mag-
nification about 7.)

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. 166, 7) and sometimes forms a
conspicuous spine projecting downwards. The eyes are ex-
tremely imperfect organs, consisting merely of six, in some cases

' For anatomy of caterpillars refer to Lyonnet’s famous work, 7raité anatomique
de la chenille qui ronge le bois de saule, La Haye, 1762.

VI LARVA ae5

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 a
structures, apparently very imperfect a
functionally, the caterpillar’s sight
being of the poorest character.’ 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 ; Pe ey ae eae
there are no true stigmata on the a caterpillar, with the jaws
second and third thoracic segments, Leela Gia
though traces of their rudiments or ocelli ; e, maxilla ; 7, lingua; g,
vestiges are sometimes visible. spinors te cain pall,

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” 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—(Lymantriidae
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.’

The SILK-GLANDS of Lepidoptera are of great interest from the
physiological poimt 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, Blane, Téte du Bombyx mori... 1891, pp. 163, ete. ; and Landois in
Zeitschr. wiss. Zool. xvi. 1866, p. 27.

2 Entwickelungsgeschichte der Schinetterlinge, Cassel, 1815.

3 Tr. Linn. Soc. London, Zool. 2nd Ser., v. 1890, pp. 147, 148.

320 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 35 mer., but when the age is fifty days
their weight has increased to 541 mer., being then 2 of the whole
weight of the body. In the pupa they undergo a gradual atrophy,
and in the inoth they are, according to Helm, no longer to be
found, though earlier authors were of a contrary opinion.’ Ac-
cording to Joseph, the silk-vessels begin to develop. at an ex-
tremely early age of the embryo, and are very aifferent 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
seoment 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 applied to certain metallic butterfly pupae.
The Lepidopterous pupa differs from that of other Insects in the
fact that its outer skin forms a hard shell, all the appendages of

1 For information as to the structure and function of the silk-vessels, refer to
Helm, Zeitschr. wiss. Zool. xxvi. 1876, p. 484; and Gilson, La Cellule, vi. 1890,
paplelGe * Jahresber. Schlesisch. Ges. viii. 1881, p. 116.

ll i

VI IPOH ere 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

Fic. 167.—Section of
female pupa of
Anosia plexippus,
3-4 days old. J,
pro-, II, meso-, III,
meta-thorax ; 1-9,
abdominal seg-
ments; @, anten-
na(?); «ac, aortal
chamber ; ag! - aq’,
abdominal ganglia ;
agl, accessory
glands ; ao, aorta;
br, brain ; ¢, colon ;

cp, bursa copulatrix ; cv, cremaster ; 71, first femur ; 7, food-reservoir ; /, dorsal vessel ;

i, part of intestine ; mv, Malpighian tube; ma, base of maxilla; oe, oesophagus ; 0v, ovary ;

ph, pharynx; sd, salivary duct; sg/, salivary gland ; st, stomach; 7}, first tarsus; fg,

compound thoracic ganglion ; ¢s*, ¢s?, 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 Micropterye 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. The interior of the pupa contains a

' The student will find important information as to the varieties of external

form of pupae in Dr. T. A. Chapman’s writings ; see especially 7’r. ent. Soc. London,
1893, 1894, and 1896.

320 LEPIDOPTERA CHAP.

quantity of cream-like matter, including the results of histolysis
—hbut 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.' 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
Fic. 168.—Wing-rudiments of Pieris hypodermal cells that occupy

brassicae. A, Rudiments of a wing :

the spots where the spiracles

before the first moult of the cater-
pillar: ce, embryonic cells; ch, ex- of the second and third thor-
ternal cuticle; h, hypodermis ; 0, : : ‘
opening of the invagination; ¢, aci¢ segments might be looked
trachea. B, posterior wing-rudiment foy, (It will be recollected that
of full-grown caterpillar; }, semi- ‘
circular pad; c, a bundle of the there are no spiracles on these
rolled tracheae ; e, envelope ; 7, pedi- “ay pr ; yj]
Boe Gaies (after Cony two thoracic segments in Lepi
dopterous larvae). Gonin has

examined the wing-rudiments in the caterpillar, a few days old, of
1 Latter, Tr. ent. Soc. London, 1895, p. 399.

‘ascertained ; but 1t would appear from

VI DEVELOPMENT—WINGS—NERVURES 329

Pieris brassicae, 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
to the older entomologists, and gave
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

Gonin’s observations that it 1s not py¢, 169.—Anterior parts of a cater-
by a process of evagination, but by Pillar of P. brassicae, the body-

3 3 é wall having been dissected off,
destruction of the hypodermis lying immediately before pupation.

y

outside the wince, “However this may % %» Snterior and pasterior

cota wings ; st J, first spiracle ; p, p’,
be, 16 1s well known that, when the second and third legs. (After
caterpillar’s skin is finally shed and ©o™™)
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. 170, A). The
nervures apparently differ as to the structures found in them.
Spuler observed in a nervure of 7riphaena 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.! Schiffer?
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

Fia. 171.—Scales of male Lepidoptera.
A, Scale from upper surface of
Everes comyntas ; B, from upper

Fic. 170.—Structure of wing of imago. A, surface of Pieris rapae ; C, from

Transverse section of basal portion of wing
[of Vanessa ?] containing a nervure ; ¢,
cuticle ; fr, wing-rib ; g, wall of nervure
(“Grundmembran”’) ; h, hypodermis ; p,
connecting columns : 7, lumen of nervure ; B,
section of a rib ; 4, one of the chitinous pro-
jections ; str, central rod. (After Schiiffer.)

inner side of fold of inner margin
of hind wing of Laertias philenor ;
D, one of the cover-scales from the
costal androconium of Hudamus
proteus ; E, F, G, scales from andro-
conium 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 Zeitschr. wiss. Zool. liii. 1892, p. 623. 2 Zool. Jahrb. Anat. iii. 1889, p. 646.

|
|

VI SCALES Sei |

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
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 y,
the transitions between hairs and yg. 179. Insertion of scales. A.
true scales may be found on the — Socket holding the stalk in Galleria
‘ : 2 mellonella ; B, insertion of the scale
wings of certain Lepidoptera that of Polyommatus phloeas. 6, Base of
bear both hairs and scales, eg. Seale fee be 2D IESE (Os
Ithomia. It has Veen calculated es ct
that there are a million and a half of scales on the wings of an
individual of the genus Morpho. The scales are arranged on
the wing in an overlapping manner, somewhat like slates on the
roof of a house. ach 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 wing, 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 ' 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

7

i

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.

WN

il i
\

4
iii
LJ

Fia. 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. a, Upper covering-scales attached to the costal
portion of the under surface of the wing; }, 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; jf, 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 androconium 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
ot the thoracic segments the hind wing is brought to he under
the anterior one (ze. between it and the body), so that in the
newly formed pupa the arrangement is that shown in Fig. 174.
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

Ww
Los)
Ww

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 Com-
stock and Needham,’ 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
* ae Oe “1 Fic. 174.—Transverse section of
formation of the nervures ; possibly ans Gi dine. madk; Ghecuenl
this may be by setting up changes in chrysalis of Pieris brassicae,
tl ll nl ae eat showing the position and struc-
1 Cems hear them py means 9 ue ture of the wings, hanging
air they supply. Semper long ago from Che ee es ue body.
li ee | I i, d : l : li 1 i tr aa, Anterior wing ; &p, pos-
discovered hypodermal cylinders tra- terior wing; ¢, ¢, peritracheal
versed by a string (Fig. 170 B) spaces ; ¢, ¢, tracheae. (After
co) fo) ) ? ites
: Gonin.)

placed near the tracheae in the
pupa.” It appears probable that the “wing-ribs” found in
the nervures (Fig. 170, A fr 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 Amer. Natural., xxxii. 1898, p, 256. 2 Zeitschr. wiss. Zool. viii. 1857, p. 326.

334 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 Schiffer, 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!
that in some cases the colours are of the nature of urates; 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 Newbigin suggests that the organic
pigments used in scale-coloration will be found to be of two

Fia. 175.—#arly condition of scales and nervures. (After Semper.) A, Section of
portion of wing of pupa of Sphinx pinastri; a, basal membrane with trachea
beneath it ; c, scale-forming cell; d, early state of a scale; e, e, more advanced
stages; f, hypodermal cells. B, part of a cellular cylinder that excretes the
nervure [or more probably the rib or ‘‘ Rippe” of Schiiffer ; cf. Fig. 170, B]; 0.
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.” Marchal, who
has devoted a great deal of attention to the study of the Malpi-
ghian tubes, informs us that the subdermal pigments of cater-
pillars are frequently 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. <A similar condition as to
the melanin-pigments and the respiratory functions appears also

1 Phil. Trans. 186 B, 1896, No. 15. * Natural Science, viii. 1896, p. 94.
3 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
Nemeobius 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-four hours, and only commenced about
forty-eight hours before the imago emerged." 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
rariations” or “seasonal dimorphism.” It has, however, been
shown that, by careful management, the eges of a generation (say
form «@) may be made to produce form a, whereas in the usual
course of nature they would produce form $. 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.

3306 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 ayaa.
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
marecellus, 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 Lep1-
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. Poulton 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 Trans. ent. Soc. London, 1892, p. 2938, etc.

VI SENSITIVENESS—MIMICRY 3

ios)
N

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 known, 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.’ Usually the resembling
forms are actually associated during life. Bates, who 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 ;”* 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.

2 Trans. Linn. Soc. xxiii. 1862, p. 507.

WAOIEE \Val Z

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 behef that birds ‘have been but little concerned in
the matter. If we add to this that there is no sufticient 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.| 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 Brazihan Macroglossa
exactly resembles a humming-bird, in company with which it
flies ; and the same naturalist also tells us* of a Skipper butterfly
that greatly resembles a grasshopper of the genus 7ettix, 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,” £7d/.
Zool. 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.

2 P. Zool. Soc. London, 1883, p. 372.

° 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 Stett. ent. Zeit. li. 1891, p. 264; and lvi. 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 Rhopalocera ( = butterflies) and Hetero-

Fic. 176.—Clubs of butterflies’
antennae. Terminal portions
of antenna of, 1, Pieris bras-
sicae ; 2, Styx infernalis ; 3,
Hestia idea (sub-family Dan-
aides) ; 4, Hudamus proteus,
and 5, JLimochores tawmas
(Hesperiidae). (After Schatz
and Scudder.)

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 Pyralidae, Ptero-
phoridae and Orneodidae in Microlepidoptera. The division of all
Lepidoptera into two series 1s merely a temporary device necessi-
tated by imperfect acquaintance with morphology. The division
into Macro- and Micro- lepidoptera is entirely unscientific.

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. Heterocera or Moths.—Antennae various in form, only rarely
knobbed at the tip, and in such cases a frenulum present. In the
large majority a frenulum is present, and the costal nervure of the
hind-wing is either but little arched at the base (as in Fig. 161,
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 341

It may be inferred from these definitions that the distinc-
tion between the two sub-Orders is neither sharply defined nor of
great importance. The club of the antenna of the Rhopalocera
exhibits considerable variety in form (Fig. 176). 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. Reuter
considers them a distinct sub-Order with the name Grypocera.”

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
empodium in connection with them. Fam. PIERIDAE.

1 For an account of the antenne of butterflies, see Jordan, Nov. Zool. v. 1898
pp. 374-415.

* Haase first proposed the name Netrocera (Deutsche ent. Zeit. Lep. iv. 1891,
p- 1) for Hesperiidae, as a division distinct from all other butterflies; Karsch
replaced the name in the following year by Grypocera, because Netrocera is the
name of a genus.

b]

342 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. PAPmnioNipaer.

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. HesPERIIDAg.

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
“Rurales.” It is however difficult to go so far and no farther ;
for the relations between both divisions of Rurales and the
Nymphalidae are considerable. We shall subsequently find that
the genus Libythea 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
familes 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 Nymphalo-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 Hesperiids, such as Thanaos,
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 im 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."

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, while 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.— Zhe front pair of legs much reduced
in size in each sex, their tarsi in the male with but one joint,

1 The literature of butterflies has become extremely extensive. The following
works contain information as to general questions : 1, Scudder’s Butterflies of New
England, a beautifully illustrated work completed in 1889, and replete with
interesting discussions. 2, Staudinger, Schatz and Rober, Exotische Tagfalter,
in three folio volumes (Fiirth, 1884-1887), with illustrations of exotic butterflies
and a detailed sketch of their characters. 3, Enzio Reuter, ‘‘Uber die Palpen
der Rhopaloceren,”” in Acta Soc. Sci. Fenn. xxii. 1896, treating fully of classifica-
tion and phylogeny.

344 LEPIDOPTERA CHAP.

though in the female there are usually five but uithout any claws.
Pupa suspended by the tail so as to hang down freely. We
include in this family several sub-families treated by some
taxonomists as families; in this respect we follow Bates, whose
arrangement! 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-
Fic. 177.—Pupa of the Purple Emperor butterfly, creased by patches lke

pati tre Men Fora Intel, B gold or silver, placed om

with the suspensory hook; D, one hook still Varlous parts of the body.

mage arr It is believed that the
term chrysalid was first suggested by these golden pupe.
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 (Fig. 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,
hikernates 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.—/ront wing with inner-margin

* Journal of Entomology, i. 1862, p. 218: for early instars of South American
Nymphalidae see Miller, Zool. Jahrb. Syst. i. 1886, p. 417.

VI RHOPALOCERA—_DANAIDES 345

(submedian) nervure, with a short fork at the base. Cell.of hind
wing closed. Front foot of the female ending in a corrugate
knob. Caterpillars smooth, provided with a few long fleshy pro-
cesses. The claws are in a variable state, being sometimes simple,
as in Papilonidae, 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 Huploea 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, Cr. (unfortunately
also called Anosia menippe, Hb., and Danais archippus or even D.
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,
Asclepias. 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.’

Some, at least, of the Danaides are unpleasant to birds in
odour or in taste, or both. Among them there occur, according
to Moore” 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;* 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.

This is the subject of Scudder’s Life of a Butterfly, 1893.
P. Zool. Soc. London, 1883, p. 205.
Finn, J. Asiat. Soc. Bengal, Ixvi. 1896, p. 528 ; lxvii. 1897, p. 213.

we toe

340 LEPIDOPTERA CHAP.

Sub-Fam. 2. Ithomiides—Difers from Danaides by the
female front foot having a true, though somewhat abbreviate
tarsus. The caterpillers 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 Ituna 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. Jthomia and
Hymenitis ave remarkable for the
small area of their wings, which
bear remarkably few scales, these
ornaments being im many cases
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. Zithorea 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; 7ithorea has colour analogues in
Heliconius, and Ithomia in Dismorphia (formerly called Leptalis).
Crowds of individuals of certain species of J/homia are occasion-
ally met with, and mixed with them there are found a small ©
number of examples of Dismorphia coloured like themselves.
They are 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

Fic. 178.—Ithomia pusio. Brazil.

i aaaaaOOOOoaOOOOooOoeEEEE OEE eOoeow
= ———— eee eee

VE RHOPALOCERA—ITHOMIIDES—-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

1 The larvae are said to live on

the “theory of mimicry.”
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 together, 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 bifid. Pupa generally suspended
by the cremaster, 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 Hrebia 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
eround. 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 two 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 le on the ground; sometimes in a very
feeble cocoon or cell. There are no cremasteral hooks. The
pupae of the Grayling butterfly, Hipparchia semele, 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 Nee 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
Huetera, 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-lke forms are those characteristic of the family in general.
The genus ELlymnias has been separated by some authorities as
a sub-family, or even as a family, Elymniidae, chiefly on the
ground of a slight pecuharity im the termination of the branches
of the veins at the outer angle of the front wings. The Alymnias
are said to be of a mimetic nature, having a greater or less
resemblance to butterflies of various other divisions; there 1s
also a considerable difference in appearance between their own
sexes. The larva of #. wndularis 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.—7here is no cell on the hind wing,
the discocellular nervule being absent (Fig. 161,11. B). Caterpillars
smooth or spiny, with the extremity of the body divided ; frequently
gregarious. These Insects have become notorious from the extra-
ordinary brillancy of blue colour exhibited by the upper surface of
the wings of the typical genus Morpho. The species of Morpho 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 1s in JMorpho, 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 Morpho, 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 Morpho 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, W. Miiller tells us! 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 JZ achilles sit in companies, often
of more than 100 individuals, on trunks of trees, and so form a
conspicuous patch. The caterpillars of JL epistrophis hang to-
gether as red clumps on the twigs of their food-plants. Hence
it appears that in this genus we have an exception to the rule
that night-feeding caterpillars rest in a hidden manner during
the day.

Sub-Fam. 5. Brassolides.— Large butterflies, with the cell of
the hind wing closed, and usually with a small adjoining predis-
coidal cell. Larva not very spiny ; thinner at the two ends, the tail
bifid, 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 J/orpho, they are
adorned, @specially 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.

LEPIDOPTERA CHAP.

(After Burmeister.)

alt

Rio de Janeiro.

Fria. 179.—Larva of Caligo (Pavonia) ewrylochus.

attains its highest perfection in the
genus Caligo; they are enormous
Insects, and some of them not rare.
The larva of C. ewrylochus (Fig. 179)
during early life is green, and _ sits
on the leaf of a Musa, but after the
third moult it becomes brown and
hides itself among the dry leaves. It
is common in the gardens of Rio de
Janeiro, where its pupae are found on
the walls, hke those of our white
butterflies here.

Sub-Fam. 6. Acraeides. — Sub-
median nervure of fore wings not forked
at the base; the median without spur.
Cells closed. Palpi in section cylindric,
sparingly set with hairs. Larva armed
with branched spines. A somewhat
monotonous and uninteresting division ;
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 diver-
sified 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 1s
only one Eastern genus, but others
prefer to adopt seven or eight divisions.
Alaena 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 sub-family
are considered to be of the protected
kind.

a eee SC ET OIE ERTS Lr

VI RHOPALOCERA——-HELICONIIDES 35a

Sub-Fam. 7. Heliconiides.—Suwhmedian nervure of front wing
not forked ; median with a short spur near the base. Cell of hind
wing closed by a perfect nervule. Palpi compressed, with scales at
the sides, in front covered with hairs. Male with an elongate
unjointed, female with a four-jointed, front tarsus. Caterpillars set
with branched spines. This family is peculiar to tropical America
and consists of only two genera, Heliconius and Hueides, with
about 150 species; but it is one of the most characteristic of the
South American groups of Butterflies. It is very closely alhed 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 Zithorea, Melinaea, Melanitis ; there
being in several cases a great resemblance between species of the
two groups. <A frequent feature in one group of /eliconius 1s
that the hind wing bears a patch of red prolonged outwards by
angular radiating marks. The individuals of certain species—J/Z.
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. A. 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 Argynnis-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
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
wing, either closed only by imperfect transverse nervules or entirely
open. Front tarsus of the male unjointed and without spines,
of the female four- or five-jointed. Caterpillar either spined
or smooth; in the latter case the head more or less strongly
horned or spined, and the apex of the body bifid. 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? 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, among
which are included the Fritillaries, Admirals, Purple Emperor, and
the various Vanessa—Peacock, Camberwell 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 in
the Tortoise-shell butterfly; it is as if the creature knew that
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 P. ent. Soc. London, 1879, p. xxix.
? Allen’s Naturalists’ Library, Butterflies, i. 1896.

VI BUTTERFLIES—NYMPHALIDES 3

unr
Ww

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.

Vanessa, or Pyrameis, 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
now at home, though it is associated with a larger and more
powerful congener, P. tameamea. Another interesting Vanessid 1s
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
Nymphalides. 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, ete. 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,*7 we may presume that in the minute details of

1A most unfortunate diversity exists in the generic names applied to these
Vanessa, as well as in those of many other Lepidoptera.

2 Ann. Nat. Hist. (6), iv..1889, p. 212.

VOL. VI 2A

354 LEPIDOPTERA CHAP.

these resemblances we have a case of hypertely similar to that
of the resemblance to Insects’ minings exhibited by certain marks
on the tegmina of Pterochroza (mentioned in Vol. V. p. 322).

In South America there is a somewhat peculiar genus of
Nymphalides— 4geronia—that delights in settling on the trunks
of trees rather than on flowers or leaves. It was long since noticed
that the species of Ageronia 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
asa 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’ 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. 7

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 Apatura 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 ‘cris, 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).—7he
female has six perfectly formed legs, though the front pair is smaller.
The male has the coxae of the front legs forming a spine, and the
tarsi unjointed, without claws. This family consists of about 1000

' P. Zool. Soc. London, 1892, p. 191.

v1 BUTTERFLIES—ERYCINIDAE 265

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 Ouvrocnemis they are large and porrect. The
family is specially characteristic of tropical America, but there is
one small group of 30 or 40 species, Vemeobiides, in the Eastern
Hemisphere. We have one species in Britain, Nemeobius 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’
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 Zurygona 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 Humaeus connects the two.
Reuter 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.— Butterflies of average size, with
the palpi large and porrect: the front legs of the male small, the
tarsus reduced to one joint: the front leg of the female of the
normal structure, and but little reduced in size. This division
consists of the single genus Libythea, with only a score of species.
They are Insects somewhat like Vanessa in appearance, but can-
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

' Bull. Soc. ent. France, 1856, pp. ¢, ci.

356 LEPIDOPTERA CHAP.

in locating Libythea therein. If they be kept apart, it 1s almost
necessary to separate Libythea also; though possibly its claims
to be placed in Erycinidae slightly preponderate. The recently —
described genus Ourocnemis to some extent connects Erycinides
with Libythaeides.’

Fam. 3. Lycaenidae.—7Zhe front legs but little smaller than
the others: in the male, however, the tarsus, though elongate, is only
of one joint, and is terminated by a single claw. No pad on the
JSront 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, 7c. 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, Vanessula,
Mimacraea, etc.) resemble members of other sub-families of Nym-
phalidae, and even of Pieridae. Lycaenidae are well represented
wherever there are butterflies; 1 Britain we have 18 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. Gueénée had
previously called attention * to the existence of peculiar structures
contained in small cavities on the posterior part of the cater-
pillar of Lycaena 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, Zr. ent. Soc. London, 1887, p. 175, Pl. ix.
2 Ann. Soc. ent. France (4), vii. 1867, p. 665, Pl. xiii.

VI BUDTRERFLIES——EYCAENIDAE—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. Robson the pupa of Zajuria 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.’
Fam. 4. Pieridae.— Zhe six legs well developed, and similar
in the sexes; there is no pad on the front tibia. The claws of all

the feet are bifid, 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 has
recently been ascertained that the yellow and red pigments, as
well as the white, are uric acid or derivatives therefrom.” 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 Malpighian vessels during metamorphosis.

Our Garden-White, Brimstone, Clouded-yellows and Orange-tip
buttertles belong to this family ; as does also the South American
genus formerly called ZLeptalis., 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 Dismorphia.

The African Insect, Psewdopontia paradoxa, has nearly trans-
parent 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 J. Bombay Soc. ix. 1895, pp. 338-341.
* Hopkins, Phil. Trans. 186 B, 1895, p. 661.

358 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 infernalis, 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
Fic. 180.—Pupation of the 2° it does in Nymphalidae. It has been
Orange-tip butterfly, ascertained by experiment that if the
patients vee cr girdle round the larva be cut, the pupa-
the larva, with its girdle, tlon can nevertheless be accomplished by
prepared for the change. 4 considerable proportion of 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 FL. genutia has a
similar life-history, but the
larva leaves its Cruciferous
food-plant, wanders to an Fic. 181.—Newly-hatched larva of Euchloe car-
oak tree, and there turns to damines. A, The larva in profile; B, one

: é segment more magnified, showing the liquid-
a pupa, resembling in colour __ bearing setae ; ©, one of the setae still more

the bark of the tree magnified, and without liquid.

It is not 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 Huchloe 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 Fic. 182.—Larva of Euchloe cardamines in middle life.
A, the larva in profile ; B, one segment more magnified.

fluid-bearing spines
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.—Al/ 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 Ornithoptera is closely

1

olours, velvet- black,

uine

1
os
golden-yellow and green.)

fe

Ae
A
Le
>
=
S
Ss
ks
=
>
nN
S
=
Lea
2
=
S
=
ie)
3S
=I
—~
S
x
=~
j

ew

183.-

male,

Fra,

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

(‘AviS pur ‘ayy Sfortq ‘smMopoH “eoBJANS Jopun
ey} SUIMOYS “potovyep “opts }YUSLU oyy WO ‘sour aL)
"OTRULay ‘pasippiod (nibsaquaoyoy ) plapdoyjJUULO

362 LEPIDOPTERA CHAP.

consorts of the other sex, as to arouse in the student a feeling of
surprise, and astrong desire to fathom the mysteries involved. It
would be difticult to surpass the effective coloration of the males
in many of the species of Ornithoptera ; they are, too, very diverse
in this respect; O. 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 O. 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
O. brookiana 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 woos 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.'

In Africa Ornithoptera is to a certain extent represented
by two extremely remarkable forms, Papilio zalmoxis and P.
(Drurya) antimachus. 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 New
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 Leptocircus has parts of the front wings transparent, while
the hind pair form long tails. This genus is of interest in that

1 Ann. Nat. Hist. (6), iv. 1889, p. 2138. We trust there will not be many more

Kiinstlers, as this beautiful butterfly must certainly become extinct, if the female
be 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 Zhais, 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 (Shippers), Six perfect legs: metanotum
not free, largely covered by 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 Nymphalidae.
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,
we 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 apphed to the perfect Insects, because so many of them
indulge in a brief, jerky flight, mstead 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 Pyrrhopyge, Erycides, 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 Rhopalo-
cera in the tropics.

In their early stages the Skippers—so far as at present known
—depart considerably from the majority of butterflies, Imasmuch
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-

SL TS I I

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, but 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-

Fic. 185. —Pupation of
Badamia exclamationis.
(After Dudgeon. VJ.
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. a,
The pupa; 4, fastenings
of perpendicular threads
round pupa; ¢, cross
thread retaining the leaf
in cradle form ; d, mar-
gins of the leaf; e, mid-
rib of leaf.

genuity in the structure of the punparium, 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, ce. 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
shuftled 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.

empty 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 web, 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 mits 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, Microp-
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 sufti-
cient, as pointed out by Packard,’ 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 familes in question,
proposed a different combination. He considers that Eriocepha-
lidae should be 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 Neolepidoptera, comprising all Lepidoptera (in-

1 Mem. Ac. Washington, vii. 1895, p. 57.

VI HETEROCERA——-MOTHS 367

clusive 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 Neuroptera. Dr. Chapman, by a sketch of the
classification of pupae,’ and Dyar, by one on larval stages,? 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®-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, Noctuae,
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. Hampson’s recent work, Fauna of British India—Moths#
It includes nearly all the families at present recognised among
the larger Lepidoptera; certain families? not mentioned in this
key are alluded to in our subsequent remarks on the families :—

Tr. ent. Soc. London, 1893, p. 97, with Suppl. op. cit. 1896, pp. 129 and 567.

* Amer. Natural. xxix. 1895, p. 1066. See also Ann. N. York Ac. viii. 1895,
p- 194, and Ent. Record, 1897, pp. 186 and 196.

3 Handbook of British Lepidoptera, 1895.

* 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 oF Motus!

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, nde

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
Il. 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, see p. 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 with 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).
15. Tarsi as short as tibia, hairy; stoutly built moths
Fam. 11. Notodontidae,? sce 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 off 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 fora 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, see p. 435.
2. 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, Fam. 31. Pterothysanidae, see 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 1a 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 (Cat. 39-42).
39. Palpi rarely absent; @Q winged ;
larvae wood-borers
Fam. 20. Cossidae, see p. 395.
40. Palpi absent ; 2 apterous (Cat. 41,
42),
41. 9 rarely with legs; 2 and
larvae case-dwellers
Fam. 19. Psychidae, see p. 392.
42. ? and larvae free !
Fam. 18. Heterogynidae, see p. 392.

bo
bo

1 This is a mistake of Sir George Hampson’s. It has long been known that the
female of Heterogynis 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 2B

370 LEPIDOPTERA CHAP.

43. Hind wing with nervule 8 anastomosing

shortly with 7

Fam. 26. Limacodidae, see. p. 401.
44. Fore wing with nervure le 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 p. 400.
55. 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 le present
Fam. 41. Pyralidae, see p. 420.
©. 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 1b simple or with a very
minute fork at base
Fam. 14. Tinaegeriidae, see p. 387.
63. Fore wing with nervure la forming a large fork
with 1b at base Fam. 45. Tineidae, see p. 428.

VI HETEROCERA—CASTNIIDAE aye it
64. Wings divided into plumes (Cat. 65, 66).
65. Fore wing divided into at most two, hind wing into three
plumes . é . 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.— Zhe Jnsects of this family combine to a
large extent the characters of butterflies and moths. The antennae
are knobbed or hooked at the tip, there is a large precostal area to
the hind wing. The nervules of the front wing are complex and
anastomose so as to form one or more accessory cells (Fig. 162). 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, taxonomists are by no means
in agreement as to the limits of the family. The Castniidae
are diurnal Insects, and the North American genus J/egathymus
is by many con-
sidered to belong
to the Rhopalo-
cera. Huschemon
rapfilesiae (Fig. 186)
is extremely hke a
large Skipper with
long antennae, but
has a. well-marked
frenulum. The

Fic. 186.—Euschemon rafilesiae. Australia. (After
Doubleday. )

members of the
Australian genus
Synemon are much smaller, but they also look like Skippers.
Their habits are very like those of the Hesperiidae ; they
tht 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 might, 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
inoths. The male of H. fenestrata is provided with a sound-
producing organ similar to that of the Agaristid genus Aegocera.

The Castnia of South America are many of them like

37 2 LEPIDOPTERA CHAP.

Nymphalid butterflies, but exhibit great diversity, and resemble
butterflies of several different divisions of the family.!

The species are apparently great lovers of heat and can
tolerate a very dry atmosphere.” 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. therapon 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.? Castnia 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 TZascina
formerly placed in Castniidae has recently been separated by
Sir G. Hampson and associated with Neocastnia nicevillei, from
East India, to form this family. These Insects have the appear-
ance of Nymphald 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
hercules, 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 family is the occurrence in numerous forms of remarkable
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, Zr. Linn. Soe. London (2), i. 1877, p. 165, ete.
* For habits of some Brazilian Castnia see Seitz, Hnt. Zeit. Stettin, li. 1890, p. 258.
3 For pupa see Chapman, Ent. Rec. vi. 1895, pp. 286, 288.

VI HETEROCHERA—SATURNIIDAE 3

N
WwW

scaled. In <Aftacus these attain a large size. In other species,
such as the South African Ludia delegorquei, 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 eyes, which are
concealed when the Insect is reposing. In <Arceina, Copiopteryx,
Eudaemonia and others, the hind wings are prolonged into very
long tails, perhaps exceeding in length those of any other moths.

Fic. 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
open 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.’ The cocoon of Saturniidae
is more often continuous, 7.e. entirely closed. Packard says that

1 Souvenirs entomologiques, quatrieme série, 1891, pp. 39-46.

374 LEPIDOPTERA CHAP.

Actias luna 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." The cocoons of the species of Ceranchia
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 Loepa newara the cocoon is of a,
green colour and suspended by a stalk; looking lke the pod, or
pitcher, of a plant. The silk of the Saturniidae is usually coarse,
and is known as Tusser or Tussore ® 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, Saturnia 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 Wicken 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
Brahmaea 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

1 Amer. Natural. xii. 1878, p. 379.
2 Cotes, ‘‘ Wild Silk Insects of India,” Znd. Jfus. Notes, ii. No. 2, 1891,
15 plates.

.
|
.
.

VI HETEROCERA—BRAHMAEIDAE

BOMBYCIDAE 37,5

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 Citheronia. 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 ;
but 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 be helpless in certain
respects when placed on trees in the larval state; but the import-
ance of these points has been perhaps somewhat exaggerated.’
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 Pérez, Act. Soc. Bordeaux, xlvii. 1894, p. 236, etc.

376 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. 7. 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, Cnethocampa processionea, are
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 1s 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 Anaphe, as has been
described and illustrated by Dr. Fischer,’ Lord Walsingham, and
Dr. Holland.’ 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 of Hypsoides 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! Berlin. ent. Zeitschr. xxvii. 1883, p. 9.
2 Tr. Linn. Soc. ser. 2, ii. 1885, p. 421. 3 Psyche, vi. 1893, p. 385.

VI MOTHS—EUPTEROTIDAE—PEROPHORIDAE 397

The larvae of the South American genus Palustra, 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 amass.’ Another
species of Palustra, P. burmeisteri, Berg? 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.

Dirphia 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 Perophora
have for long been an enigma to systematists, and have been
placed as abnormal members of Psychidae or of Drepanidae,
but 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
Perophora are now known; they inhabit a large part of the
New World, extending from Minnesota to Buenos Aires. The
habits of P. melsheimeri have been described by Harris,
Packard * and Newman, and those of P. batesi by Newman.*. The
larva is very peculiar ; there is a flexible pair of appendages on the

? Bar and Laboulbéne, Ann. Soc. ent. France, (v:) iii. 1873, p- 300.
? Op cit. (5), vii. 1877, p. 181; and Ent. Zeit. Stettin, xxxix. 1878, p. 221; and
xliv. 1883, p. 402.

* Ann. New York de. viii. 1893, p. 48.
+ Tr. ent. Soc. London, n.s. iii. 1854, p. 1.

378 LEPIDOPTERA CHAP.

head, the use of which is unknown ;' they arise by slender stalks

behind and above the eyes, are about as long as the head, and
are easily broken off. 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
of a more tawny colour. Newman does not allude to any
cephalic appendages as existing in the larva of P.. batesi.
If we accept the eges figured and described by Snellen, as those
of P. batesi, 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. melsheimerc.
The genus Lacosoma is considered by Packard to be an ally of
Perophora. The caterpillar of LZ. 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 on the body of the caterpillar may be indications of this.*
The Argentinian Insect Mamillo curtisea* 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-
meri, and states that it belongs [according to the larva] to Tineidae ; the appendages
he considers to be enormously developed setae. J... York ent. Soc. iv. 1896, p. 92.

2 Tijdschr. Ent. xxxviii. 1895, p. 56, Pl. 4.

3 Ann. New 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 by Harris. The moth itself is very Geometrid-
like in colour and form. This species is now assigned to Pero-

Fic, 188.—-Larvae of Hammock-moth, Perophora sanguinolenta, projecting from their
Hammocks, built from their own excrement. South America. (After Jones. )

phora, 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-

380 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-
boseis 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 pecuhar, having a thick, sohd 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
Fic. 189. — Antennae

of Sphingidae. A,

One joint of an-

tenna of Choero-

campa elpenor é,

enlarged ; B, three

joints of antenna of

Sphinz ligust7t,

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 (Catocala, ete.). 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 Soc. xxxiii. 1879, p. lxxvii.

}

VI HETEROCERA—HAWK-MOTHS 381

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

Fic. 190.—Larva of
the Poplar Hawk-
moth, Smerinthus
PODULUa les WAS
in repose; B, in
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 have
been brought to light. We may refer the reader to the writings
of Weismann! and Poulton, without, however, recommending
him to place an implicit confidence in their somewhat
metaphysical disquisitions; for the views there shadowed wiil

' Studies in the Theory of Descent, part 2, London, 1881.
° Tr. ent. Soe. 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.’

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 ;

toe)

in this case it 1s highly probable that the method is the
friction of the palpi against the proboscis, as stated by Réaumur
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

Fic. 191.—Cocytia durvillii. New Guinea. (After Boisduval.)

the region of New Guinea, whence the first was brought by
D’Urville nearly a hundred years ago. They are still amongs
1 Tijdschr, Ent. x). 1897, pp. 27-103, 4 plates.

|
7

Vi HE TE ROCHE RA—— EO CY TIIDAE

NOTODONTIDAE 383

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’ 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 Noto-
donts are indeed considered to be very closely allied. The family
contains, however, some very remarkable forms. T'arsolepis has
an elongate body,
terminated (in the
female of 7. ful-
gurifera) bya very
conspicuous tuft
of enormously
long, _battledore
scales; while in
they male ef 7’.
sommert the hind
legs are provided
with an append-
age of beautiful,
roseate hairs. <A
few of the larger
kinds bear a con-

: i Fie. 192.—Cerura vinula (Puss-moth) caterpillar. A, Moult-
siderable resem- ing ; B, the same individual a few hours after the moult.

blance in form and

proportions to the Sphingidae. Some of the larvae are most inter-
esting objects; the Puss-moth caterpillar, the Lobster, and the
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. Soc. London, 1884, p. 351.

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.’
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 1s 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, as well as by: ejection (like so many other
larvae and insects generally) of fluid) There is no reason for
beheving 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 minnit 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 ike the spots, crevices, or other places amongst which
it is located; in this prison the creature remains for 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
thinner 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 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, Pl. x.
2 See Poulton, Zr. ent. Soc. London, 1886, etc. 3 Op. cit. 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, Stawropus fagi, 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-
pular, 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 Miiller states! 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” 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 Staphylinus is very striking. Others
say that this caterpillar resembles a dead and crumpled beech
leaf.

The larva of Hybocampa milhauserithe 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* 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 TYortria 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 fulerum

1 P. ent. Soc. London, 1880, p. iii. 2 Ent. Monthly Mag. xiii. 1877, p. 231.

3 Entomologist, xxiii. 1890, p. 92.
VOL. VI 2C

336 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 be
sundered.

Though many other larvae of Notodontidae are of most
curious form and asswne remarkable attitudes, yet this is not
the case with all, and some are quite ordinary and like the cater-
pillars of common Noctuidae. This is the case with the species
Rhegmatophila 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 North American
forms.’ The family has a very wide distribution, but is absent
from New 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
moths that connect the Bombyces with the Noctuids; 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 Noctuidae
the reverse is the case. The Insects, however, in certain respects
approach the Notodontidae, 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, Zhyatira batis, and the very
different Buff-arches, 7. derasa, being among them. Meyrick
denies any connexion of this group with Noctuidae, and in his
nomenclature Cymatophora becomes Polyploca, and the family,
consequently, Polyplocidae.

Fam. 13. Sesiidae or Aegeriidae (Clear-wings)— 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. Washington, vii. 1895, 290 pp., 49 plates.

vi HETEROCERA—SESIIDAE—-TINAEGERIIDAE 387

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,’ 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 Trochiliwm (called variously Sesia, Sphecia, and
Aegeria), with two species of comparatively large size, and Sesia
(called variously TZrochilium and
Aegeria), with nearly a dozen species
of smaller size. A third genus,
Sciapteron, 1s 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 Fic. 193. —Ocedematopoda princeps.

era ye ‘ Africa. (After Walsingham.)
due to Lord Walsingham.” Nothing zs
is known as to the life-histories, except that it has been stated
by Stainton 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. ent. Soc. London, 1878, p. 121, Pl. v. 2 Op. cit. 1889, pp. 1-40, 6 plates.

388 LEPIDOPTERA CHAP.

belong to the Tineid series of moths. The species we figure (Fig.
1935) has a character otherwise peculiar to Sesiidae in the wings
being inserted on the thorax remote from the head—a feature we
do not find in the Tineidae proper; while on the other hand it
has the long 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’ 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 lke 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 Walsingham, Op. cit., 1889. c. p. 21.

v1 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 ”
that a species of JMJacrocneme was observed by him to be exactly
like one of the blue wasps of the genus Pepsis.

One remarkable point in these Hymenopteroid 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 lke that of a Petiolate Hymenopteron; 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,’ 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. Zeit. Stettin, lvi. 1895, p. 233. 2 Op. cit. li. 1890, p. 261.
3 Ent. Zeit. Stettin, li., 1890, p. 263.

390 LEPIDOPTERA 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
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

Fic. 194.—Trichura, sp. x2. Demerara.

cocoon is formed. In Britain no member of this family is to
be met with, but Naclia ancilla may formerly have been a
native; Syniomis phegea has occurred here; probably an escaped
example that had been introduced in one of its earlier stages.
Fam. 16. Zygaenidae (Surnet-Moths)—This family is one
about the lmits 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;
bladder-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 intersegmental membranes

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

: : Fic. 195.—Hampsonia pulcherrima. Wings on
in a cabinet, the Kast right side detached and denuded to show

Indian Cadphises mooret nervuration. India. (After Swinhoe.)

eg.) 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.
Hampsonia pulcherrima (Kast 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
apart 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.

392 LEPIDOPTERA CHAP.

Fam. 17. Himantopteridae (Thymaridae of some autho-
rities) are placed by Hampson 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 lke moths.
They are peculiar to India and Africa. In the South African
genus Dianewra (belonging really to Phaudinae) also the wings
are scaleless and nearly transparent.

Fam. 18. Heterogynidae.—Consists of the single genus
Heterogynis 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 shows 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.’

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” 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, ete., see Rambur, Ann. Soe. ent. France, v. 1836,
p- 577 ; and Graslin, op. cit. xix. 1850, p. 396.

2 Monograph of European Psychidae, Ann. Soc. ent. Belgique, xxv. 1881,
j0s 7), Cwee

VI HETEROCERA—PSYCHIDAE 393

therein, and inside, also, their former pupa-skin; and here the
young hatch ; the pecuhar 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, eg.).' Great difficulties attend the study of
these case-bearing Insects, and several points require careful

Fic. 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 well 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,’ 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 hockingi
(Hig. 19:7, .@) consists <on
little pieces of wood cut
to the proper lengths, and
spirally arranged, so as to
A c form a construction that
FIia. 197.—Baskets, or cases, of Psychidae, A, would be quite a eredit to
dinicte quadreagdgris; B Aptrona(or Col ou» own species, Th. some
of the Canephorinae we
meet with long cylindrical cases, like those of Caddis-worms, or
of Tineid larvae.

Riley has given an account of several points in the struc-
ture and natural history of one of the North American basket-
or bag-worms, Zhyridopteryx ephemeraeformis ; one of his points
being the manner in which the newly hatched larva forms its
ease.” This question has also been discussed by Packard.’ The
larvae when hatched in unnatural conditions will make use of
fragments of paper, cork, ete., for the case; the act of construc-
tion takes one or two hours, and the larva does not eat till the
case 1s 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 remarked, 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

oO?
Britain) in the series Psyehina which includes Zygaenidae. He

1 Zool. Anz. xx. 1897, p. 473. This is probably Apterona crenulella, or one
of its varieties. 2 Bull. U.S. Dep. Agric. Ent. x. 1887, p. 22.
3 Ann. New York Ac. viii. 1898, p. 54.

VI . HETEROCERA—PSYCHIDAE—COSSIDAE 395

removes the other British genera, Fumea and EHpichnopteryx, 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-Schiiffer, 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.’ 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 Trypanidae) as a family of
the series Tortricina.

1 Kalender, Ent. Zeit. 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, Hollandiudae.

Fam. 22. Chrysopolomidae.—This family has been estab-
lished by Dr. Aurivillius* for an African genus, allied in wing-
nervuration to Cossidae ; the Insects are like Lasiocampidae.

Fam. 23. Hepialidae ((host- and Swift-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
alles; the conclusion that they are connected with the Micro-
pterygidae being certainly erroneous. Although but small in
numbers—only about 150 species bemg 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 glittermg metallic splashes on the wings,
making a barbaric but effective display. The South Australian
Zelotypia staceyt, of enormous size, 18 also a handsome moth ;
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” the young
larva of Hepialus mustelinus has the arrangement of setae that is
normal in Tineidae. The larva of H. humuli seems to be a very
simple form, but HH. heectus shows a considerable amount of
divergence from it. They probably live for several years; the
larva of H. argenteo-maculatus in North America lives for three
years, at first eating the roots of Alder and then entering the

1 Ent. Tidskr. xvi. 1895, p. 116.
? On larvae of Hepialidae, J. New York ent. Soc. iii. 1895, p. 69, Plates III. 1V.

VI HETEROCERA—HEPIALIDAE 397

stems. The pupae are also peculiar. They are of unusualiy

elongate, cylindrical form, with comparatively feeble integument,
but 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

)

corre corey
\ ail

MN

ff ears | \
pupa naturally placed 0 <n
amongst either the pupae BOW

obtectae or incompletae of Fic. 198.—Pupa of Hepialus lupulinus. Britain.
2 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
Hf. 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 hkely 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 New 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, Entomologist, xxviii. 1895, p. 114.

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... 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,? some being quite like the
females. In H. hectus 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 H. humuli brings its female. The hind legs of the male

1 Ent. Mag. xiii. 1876, p. 63; and xxiii. 1886, p. 164.
2? Weir, Entomologist, xiii. 1880, p. 249, plate; King, Ant. Record, vii. 1895,

job. JILL.

VI TET EROCERA—_HEPIALIDAE 399

are swollen, being filled with glands for secreting the
odorous matter." 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. /upulinus 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 Hf. sylvinus, H. velledu 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. Robson, 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 (HZ. humult) ;

(B) by odour (HZ. hectus).

2. The female attracts the male—(A) by vibration of wings
(#7. lupulinus and H. sylvinus); (B) without vibration,
but by some means acting at a distance (4. velleda,
Hi. 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, SB. Ver. Rheinland, xxxvi. 1879, p- 288; and Arch. Naturg.
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 labium 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 meta-notum being much less infolded and co-
ordinated. The wings are remarkable for the similarity of the
nervuration 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
jugum of Prof. Comstock. Brandt mentions the ‘following
anatomical peculiarities,’ 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 hfe:
The chain of nerve gangha consists of the supra- and infra-
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 Pterodecta are
very like butterflies of the Lycaenid genus Vhecla. 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). (//ook-tips)— The
larger moths of this family are of 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.
Oreta hyalodisca 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 Zool. Anz. iii. 1880, p. 186.

;
]

VI HETEROCERA—DREPANIDAE—LIMACODIDAE 401

find Insects with the scales imperfect, they being few and small
and approximating in form to hairs; in JD. hyalina scales are
nearly entirely absent. In other genera, e.g. Peridrepana, Strep-
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
shght 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 50 genera, and 150 species known, the
geooraphical distribution of the family being very wide. = In
Britain we have half a dozen species. Cilia glaucata (better
known as C. 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 ;
some, Rosema and Staetherinia,are of
extraordinary shapes; certain very
small forms, Gavara, Ceratonema,
resemble Tortricids or Tineids; a
few even remind one of Insects of Fi 199.—Mature larva of A poda

; 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 2D

402 LEPIDOPTERA CHAP.

in both hemispheres, but these Insects do not occur in insular
faunas. In Britain we have two genera, Heterogenea and Apoda
(better known as Limacodes'), each with a single species.

The early stages of these Insects are of great interest. The
eggs, so far as known, are peculiar flat 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 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 Apoda testudo has no conspicuous spines or processes,

just hatched. A, Dorsal view ;
of larva; B, C, D, a spine in and is an extremely soft, colourless
precy of evagination. creature” but it almost immediately

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. testudo is
variously styled Limacodes testudo (by Chapman and most naturalists), Apoda
limacodes (by Meyrick), or Apoda avellana (Kirby, Catalogue of Moths). The
family is called either Limacodidae, Apodidae, Cochliopodidae, or Heterogeneidae.

* See Chapman, 7’r. ent. Soc. London, 1894, p. 345, Plate VII., for our British
species ; for North American forms, Dyar, Life-histories of the New York Slug-cater-
pillars (in progress, with numerous plates), J. New Vork ent. 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,
but 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.’ 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 :>—
“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
vaterpillars 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 young shoot in drying turn a light tan-color, which

1 See Packard, P. Amer. Phil. Soc. xxxi. 1893, pp. 83, 108, Plates. (He uses
the term Cochliopodidae instead of Limacodidae) ; also Dyar, as above.
* Insects affecting the Orange, Washington, 1885, p. 143.

404 IEAM IODKONEME EVAN 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 hes 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.’ 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 Hpipyrops, 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 1t may be a Limacodid, and this
appears probable.

Fam. 27. Megalopygidae (or Lagoidae).—The American
genera, Megalopyge and Lagoa, are treated by Berg and by
Packard * 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 five pairs—the
usual number in Lepidoptera. When young the caterpillars of
Lagoa opercularis are white and resemble a flock 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, ent. Soc. London, 1894, p. 348. 2 Op. cit. 1876, p. 522 ; and 1877, p. 433.
3 P. Amer. Phil. Soc, xxxii. 1894, p. 275.

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 Rhodoneura, 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.

Fam. 29. Lasiocampidae (Lygers, 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-direected, 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 1s great in some
species. In the genus
Suana and its allies we meet
with moths in which the

difference in size of the Fic. 201.—Lappet-moth, Gastropacha querci-
folia, 2. Britain.

two sexes is extreme; the
males may be but 14 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, Borocera 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 differmg 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. 608.

406 LEPIDOPTERA CHAP.

some species the walls of the cocoons have a firm appeaz-
ance, looking very lke 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.’ 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 be prolonged for several years. There is not at
present any explanation of this fact. It may be 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,” EHndromis
versicolor, forms this family; it 1s a large and strong moth, and
flies wildly in the daytime in birch-woods. The larva has but
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. ent. Soc. London, 1891, 1b AA

\Git 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 Inany 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 Lymantriidae is one of the largest familes of the old group
Bombyces ; it includes some 180 genera and 800 species, and is
largely represented in Australia. Dasychira 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. Psilwra monacha—the Nun,
“die Nonne” of the Germans,'—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 Wachtl may be found in Wien. 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.' 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 inago ;
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.

Dasychiva pudibunda 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.” Zeara 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
buff 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 1s 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 Bombycid series of
moths, comprising something like 500 genera and 3000 species.
Hampson recognises four sub-families— Arctinae, Lithosiinae,

1 Wachtl and Kornauth, Mitt. forst. Versuchswesen Osterreichs, Heft xvi. 1893.
* Crahay, Ann. Soc. ent. Belyique, xxxvii. 1893, p. 282.

VI HETEROCERA—ARCTIIDAE 409

Nolinae, Nycteolinae,—to which may be added others from
America—Pericopinae, Dioptinae, 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 Ocnogyna) where the female is wingless
and moves but little, while the male flies with great rapidity.
Epicausis 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 Pericopinae are almost peculiar to South
America (two species of Gnophaela exist in North America) ;
some of this sub-family bear a great resemblance to Heliconid
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. Hampson 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

410 LEPIDOPTERA CHAP.

species are native in Britain. Packard maintains the family
Nolidae as distinct."

The sub-family Nycteolinae consists of a few small moths the
position of which has always been uncertain; Wycteola (better
known as Sarrothripus), Halias, and Harias are all British genera
that have been placed amongst Tortrices, to which they bear a
considerable resemblance. Sarrothripus is at present placed by
Hampson in Noctuidae, by others in Lithosidae, by Meyrick in
Arctiidae. The sub-family forms the family Cymbidae of Kirby;”
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, Harias, 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 ;
Donychopus niveus produces a sound by, it 1s believed, friction
of the wings, In the case of the genera Setina and Chelonia
the process is said to be pecuhar to the male sex: Laboulbene
believes it to proceed from drum-like vesicles situate one on each
side of the base of the metathorax.’

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 Husemia villicoides 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.
In 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.

Amer. Natural. xxix. 1895, p. 801.
2 Catalogue of Lepidoptera Heterocera, i. 1892.
3 Ann. Soc. ent. France (4), iv. 1864, p. 689.

VI HETEROCERA—AGARISTIDAE—GEOMETRIDAE 411

The male of the Indian Aegocera 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 otf
A, tripartita, yet it occurs in a very similar form in the genus
Hecatesia, already noticed under Castniidae.

Fam. 36. Geometridae (Carpets, Pugs, etc.)— 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 frenulumn
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-familes,
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.” 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 Himera

1 P. Zool. Soc. London, 1892, p. 188.
2 Tr. ent. Soc. London, 1892, pp. 53-140; for criticism on the nomenclature,
see Rebel, Ent. Zeit. Stettin, lili. 1892, p, °247.

AI2 LEPIDOPTERA CHAP.

pennaria 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-hke 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 1s 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
(Ephyra pendularia).
Certain of the larvae of
E 3 Geometridae vary in

colour, from shades of

brown to green; there

Fig. 202.—Larva of Amph idasis betularia, reposing is much diversity in
on a rose-twig. <1. Cambridge. 4 iy.

this variation. In some

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 Amphi-
dasis betularia it would appear that when the larva is hatched
the dimorphism is potential, and that the future colour, whether

ae GEOMETRIDAE ve

green or brown, is settled by some determining condition during
the first period of larval life and cannot be subsequently moditied.!
According to Poulton, the dark tint is due in A. betularia to
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 beimg 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
fight. 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 be 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.”

1 See Poulton, 77. ent. Soc. London, 1884, p. 51; op. cit. 1892, p. 293; and
Bateson, p. 213; Gould, p. 215.
2 Giraud, Ann. Soc. ent. France (4), v. 1865, p. 105; Fauvel, 7.c. 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
Cheimatobia 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-
inality.' Amphidasis betularia 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.”

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 out,
it has never received any thorough investigation. Few other
remarkable structures have yet been discovered: the most in-
teresting is perhaps the pecuhlarity in the hind wings of the
males of certain Ommatophorinae recently pointed out by Sir
G. F. Hampson *: in the genera Patula and Argiva 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, J.c.

’ Barrett, ‘‘ Increasing Melanism in British Geometridae,” Ent. Monthly Mag.
1895, p. 198.

3 P. Zool. Soc. 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
clistributed of all the genera of moths. Such caterpillars are
known as Cut-worms in North America.’ The great resemblance,
enter se, of certain of these Cut-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.
Huclidia 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. 7riphaena,
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 Fic. 203.—Brephos notha. Larva, newiy
appears in the form shown aa a aaa
in Fig. 204. This interesting larva joins together two or three

1 Although this term is widely used in North America, it is not in use in Eng-
land, though it may possibly have originated in Scotland. See Slingerland, Budi.
Cornell University Exp. Stat. 104, 1895, p, 555.

416 LEPIDOPTERA CHAP.

leaves of aspen and lives between them, an unusual habit for
Noctuid laryae. When about to pupate it bores imto bark or
soft wood to change to a pupa, Fig. 205;
the specimen represented closed the hole
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-

Fia. 205.—Brephos notha. A,
Pupa, ventral aspect; B,
extremity of body, magni-
tied ; C, the pupa in wocd ;
d, diaphragms constructed

Fic. 204.—Brephos notha. Adult larva. by the larva.

esting Insect is very uncertain: Meyrick and others associate it
with the Geometridae.

The larva of Leucania unipunctata is the notorious Army-
worm that commits great ravages on grass and corn in North
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—tfor 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, Aletia
aylinae is even more notorious on account of its ravages. Tiley
states ' that in bad years the mischief it commits'on the cotton
crop causes a loss of £6,000,000, 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
Heliothis armigera, which bores into the buds and pods. This

1 Fourth Rep. U.S Ent. 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 1s 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 Noctuid moth—AFrastria scitula
—are of great value. Its habits have been described by
Rouzaud.' On the shores of the Mediterranean the larva of this
little moth lives on a Scale-Insect—Lecaniwm 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 Hrastria 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 2E

418 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, 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
“ Apatela 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, Lryophila, possess the excep-
tional habit of feeding on lichens. Some of the American group
Erebides are amongst the largest Insects, measuring seven or
eight imches across the expanded wings.

The Deltoid moths are frequently treated as a distinct family,
Deltoidae, perhaps chiefly because of their resemblance 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 Polyxenus 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 See Chapman, The Genus Acronycta and its Allies, London, 1893.
” Insects Injurious, ete., Ed. 1862, 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 Nyctalemon, are like Swallow-tail butterflies and have
sunilar habits. The Madagascar moth, Chrysiridia madagascar-
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 Noctuidae! 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
Chrysiridia madagascariensis
has been widely spread; but
according to Camboué,’ the
account of the metamorphoses,
first. given by Boisduval, is
erroneous. The larva, it ap-
pears, resembles in general

form that of Coronidia, and Fic. 206.—Abdomen of Chrysiridia mada-
gascariensis. A, Horizontal section show-

has sixteen feet ; 1b 18, how- ing the lower part of the male abdomen :
ever, armed with long, spatu- 1, first segment; 2, spiracle of second

4 : segment ; 4-8, posterior segments. B,
late black hairs ; 1t changes to the abdomen seen from the side, with the

a pupa in a cocoon of open segments numbered, The section is that
of an old, dried specimen.
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-lke opening (usually

' See Westwood, Zr. Zool. Soc. London, x. pp. 507, ete., for discussion of this
question and for figures ; also E. Reuter, ct. Soc. Sci. Fenn. xxii. 1896, p. 202.
* Congr. Internat. 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 1s
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 familes 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 are
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 pecuhar 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 retinaculum, consisting of large
scales, and this, too, is connected with a distortion of the wing-
nervures. The Pyralidae—Pyralites of Ragonot, Pyralidina of

1 Ragonot, Ann. Soc. ent. France, 1890 and 1891; and Meyrick, Zr. 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 Ragonot does
not include the two groups named, but defines seventeen tribes
of the two familes—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 Ragonot’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 25 species ;
Madeira, 3); St. Helena, 6; South Atrica, 2° or 3-3. India,’ 9;
Malayan region, 3 or 4; Australia, 24; New Zealand, 64;
Hawaiian Islands, 50; North America, 17 (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 Hydrocampa nymphaeata were long since
described by Reaumur, and have more recently been dealt with
by Buckler,’ W. Miiller* and Prof. Miall.2 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. % Nat. Hist. Aquatic Insects, London, 1895.

4.22 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
araft; 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 Réaumur 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 ;
but 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 habits at some particular moult, accompanied by great
changes in the integument, and even in the size of the stigmata,
being of frequent occurrence.

VI HETEROCERA—PYRALIDAE 23

The larva of Nymphula stagnata, a close ally of H. nymphaeata,
has aquatic habits of a somewhat similar but simpler nature ;
while V. (Paraponyx) stratiotata 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, hke H. nymphaeata, 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 Paraponyz.

Musotimidae’ is a small group of two or three genera found in
Australia and Polynesia ; and the Tineodidae 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 Ragonot. Epipaschiinae (formerly
treated 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 Tortricidae
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

' For Bibliographic references connected with the divisions of Pyralidae see
Ragonot, Ann. Soc. ent. France (6), x. 1890, pp. 458, ete.

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
Ephestia infest groceries, and most children have become to a slight
extent acquainted with them amongst dried figs; that of JZ.
kuehniella 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 on Excoecaria biglandulosa, a Euphorbiaceous tree.
The female moth lays an egg on a gall, and the resulting larve
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 has thus
acquired. Another Phycitid, Dakrwma coccidivora, is very

1 Monograph, by Ragonot, in Romanoff, Mem. Lep. vii. 1893.
2 Ent. Zeit. Stettin, 1878, p. 230.

VI HETEROCERA—PYRALIDAE A425

beneficial in North America by eating large Scale-Insects of the
Lecanium group, somewhat after the fashion of Hrastria scitula ; 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.’ 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 on grass, and
construct silken tunnels about the roots at or near the surface.
The Ancylolominae are included in Crambidae by Meyrick and
Hampson. Schoenobiinae” 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.’ They have no trace of gills, and their

1 Howard, Insect Life, vii. 1895, p. 402.

* Monograph by Hampson, P. Zool. Soc. London, 1895, p. 897-974.

8 Disqué, Ent. Zeit. Stettin, li. 1890, p. 59. Cf. also Rebel, Zool. Jahrb. Syst.
xii. 1898, p. 3.

426 LEPIDOPTERA CHAP.

mode of respiration is unknown. A great deal has been 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! (P/wime-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 <Agdistinae
(of which we have a British representative of the only genus
Agdistes) have the wings undivided. Pterophorinae have the
hind wings trifid or (rarely) quadrifid, 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 lke 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
humerous 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, oceur in various regions. The larva and
pupa are less anomalous than those of the Pterophoridae, though
the imago is more anomalous. The caterpillar of our British A.
polydactyla feeds on the flower-buds of honey-suckle, and forms a

1 Classification ; Meyrick, Zr. ent. Soc. London, 1886, p. 1.

VI HETEROGCERA——TORTRICIDAE AZ 7,

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, 1b, 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 Tineidae. 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 Pyralidae, 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,’ Tortricidae, Grapholithidae, Conchy-
lidae ; subsequently,” he has adopted the names Tortricidae,
Epiblemidae, Phaloniadae. Lord Walsingham, who has devoted
a great deal of time and study to the elucidation of this most
difficult group, has suggested’ 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
bemg family or sub-family]; 2. Phaloniudae [=the formerly
used name, Conchylidae]; 3. Olethreutidae [ =the formerly used
name Grapholithinae = Epiblemidae, Meyr.]. We have upwards
of 500 species in Britain, nearly 200 of which belong to the
last division. The name Tortricidae refers to the habit the

1 P. Linn. Soc. N. S. Wales (2), vi. 1881, p. 410.
2 Handbook Brit. Lep. 1895, p. 493. 3 Tr. ent. 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:—1, 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 funebrana, “larva frequent in plum-pies”; the cater-
pillar of Carpocapsa pomonella (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.
juliana in Spanish chestnuts. Two, if not more, larvae live in
the seeds of Euphorbiaceous plants, and have become notorious
under the name of jumping-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
from skin to skin; in this the larva makes a movement sufficient
to alter the point of equilibrium 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.
Curpocapsa saltitans living in the seeds of Croton colliguaja and
Grapholitha sebastianiae living in the seeds of Sebastiania
bicapsularis.

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 head 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,

————a eee rr eee SES

VI HETEROCERA—TINEIDAE 429

if Mr. Meyrick be correct in supposing that a single one of
the divisions of the family— Oecophoridae—comprises 2000
species in Australia and New 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 classifi-
cation is in a comparatively
rudimentary state. We shall
not, therefore, deal with it here. a pat any
= : Fic. 207.—Diplosara lignivora (Gelechiides).
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 Fig. 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, eg. Thysanoptera 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. No 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

_1ts internal anatomy: the testes consist of four round follicles

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 quite free (as they are in Hepialus),
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. Anz. y. 1882. p. 262.

43.0 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, Bradypus 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. Occocecis
guyonella is said to be the cause of galls on Limoniastrum
guyomanum, 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 Guénée! 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.
Trichophaga tapetzella is perhaps entitled to be considered the
Clothes-moth ; its caterpillar not only feeds on clothes, but spins
webs and galleries amongst them. TZinea pellionella is also very
common; its larva lives in a portable case, while that of the
third species, 7ineola 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 Zinea have been recorded as viviparous, viz.
Tinea vivipara in Australia, and an undetermined species in
South America. The species of the genus Solenobia—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, pl. 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.’ 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
eases of various forms. Some are leaf-miners during their
early life, and subsequently change their habits by con-
structing a sortable case. The genus Coleophora 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, Vematobola orthotricha ;? 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 ;* 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 ana
trees. Before pupating, the larva folds down the edges of the
hood over the mouth of the tube, like an envelope, fastening
them with 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. 1-8.

2 P. Linn. Soc. N.S. Wales (2) vii. 1892, p. 593.

3 Durrant, Ent. Mag., xxxi. 1895, p. 107.

432 LEPIDOPTERA CHAP.

The family Prodoxidae 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
mouth, exhibit a
remarkable modifi-
cation of instinct.
The facts are chiefly
known from the
observations of
Riley ' on Pronuba
yuccasella, a moth
living on Yuweca
jilamentosa ; this
plant has been in-
troduced into our
gardens in this
country, where it
never, we believe,
produces seed. The
Yuceas 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
Fi 300. Pronie Gree a es eb pat
E, head and part of thorax of the female moth: a, plant, and as these

labial palp; 6, maxillary tentacle; c, maxillary palp ;
d, proboscis ; e, base of front leg. (After Riley.)

MGzy f: . a}
Aid ggyyyni!

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
moth first visits the stamens, and collects, by the aid of the

1 «The Yucca moth and Yucca Pollination,” Rep. Missouri Botanical Garden,
1892, pp. 99-158.

VI HETEROCERA——PRODOXIDAE——ERIOCEPHALIDAE A233

maxillae (which in this sex are very remarkably formed), 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
apples 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 Yuceas, 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 we figure is the largest
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.” 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

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, D- 161:

2 Chapman, 7’r. ent. Soc. London, 1894, p. 366.

VOL. VI ; AB

434 LEPIDOPTERA 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
mandibulate Insects. riocephala in both these respects connects
the Lepidoptera with Mandibulata: the mandibles have been
shown by Walter! 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 ttle 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.” The wing-characters of Eriocephala are repeated—as
to their main features—in Hepiahdae and Micropterygidae ; but
both these groups differ from Hriocephala as to the structure of
the mouth-parts, and in their metamorphoses. Although Z7v0-
cephala calthella 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, yet
its metamorphoses were till
recently completely unknown.
Dr. Chapman has, however,

been able to give us some FIG. 209.—Larva of Eriocephala cone
é ; ; (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, in or ball-like appendage: c, abdominal foot
‘ Fics of larva,

both of which points the crea-

ture is most interesting. The eggs and young larvae are “ quite

1 Walter, Jena. Zeitschr. Naturw. xviii. 1885. He did not distinguish Hrio-
cephala as a genus, as we have explained on p. 308.
2 Amer. Natural. 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, owing 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 (Limacodids), but Dyar is probably
correct in thinking the resemblances between the two are unim-
portant: the larva of Hriocephala 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,
trifohate 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-
mira is only imperfectly known.
Meyrick considers it the “most
ancient”? Lepidopteron yet dis-
covered ; and it would appear that
its relations are with Hriocephala
rather than with MWicropteryz. From
information he has kindly given to
us, we are able to say that this moth
possesses mandibles but no proboscis.

A : Fic. 210.
Fam. 47. Micropterygidae.— A, Ventral view of the larva,

Small moths of metallic colours, ™*gnified; B, the same, with

Larva of Micropteryx sp.

é = : setae unduly magnified. Britain.
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

430 LEPIDOPTERA CHAP.

veins. There are no 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 Z7io-
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 ;* 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
Fig. 211.—Pupa of Micropteryx (semipurpurella ?). A, greater PaEe of its life
Dorsal aspect ; B, C, D, views of head dissected off; aS a larva, changing to
ae 2 ae G as D, anterior aspects; m, 9 pupa very early in
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.” In the interval between

1 Wood, Ent. Mag. xxvi. 1890, p. 148.
* See Chapman, 7%. ent. Soc. London, 1893, p. 255.

v1 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 Eriocephala, 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, ike the deciduous,
supplementary mandibles found in Otiorhynchides (Coleoptera,
Rhynchophora). }

CHEAP TER Vil

DIPTERA—-OR FLIES ; APHANIPTERA——OR FLEAS; THYSANOPTERA
Oe, WSO ES)

Order VII. Diptera

Wings two, membranous, usually transparent and never very large ;
behind the wings a pair of small erect capitate bodies—
halteres —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.

Tuts 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-
ing to study them, or
to collect them for
observation. | Never-
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

Fic. 212.—A Dipteron (Fam. Syrphidae), Cheilosia chryso-

been most perfected. coma. Britain. A, Adult larva; B, the pupa; C,
y\ maggot, hatehine nymph, extracted from pupa; D, imago. (From
foto) 5

; Weyenbergh.)
from 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

.

440 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.” 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.® 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 Osten Sacken, Zr. ent. Soc. London, 1884, p. 501, and Berlin. ent. Zeitschr.
XXXVii. 1892, p. 423, etc.

2 Osten Sacken has recently discussed the intermediate conditions, and proposed
the name ‘‘ pseudholoptic”’ for some of them, Berlin. ent. Zeitschr. xli. 1896, p. 367.

3 Girschner, Berlin. ent. Zeitschr. xxxi. 1887, p. 155.

VIL ANTENNAE 44]

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. 215, 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 in
a hollow formed for their reception on the front of the head.
Fhes with this form of antennae are called Athericerous. Between
the two forms of antennae we have mentioned there exists what
may, speaking roughly, be called an intermediate condition, or

Fira. 213.—Antennae of flies. A, The two antennae of Glaphyroptera picta (Myceto-
philidae) ; B, antenna of Hexatoma pellucens (Tabanidae) ; C, of Asilus crabroniformis
(Asilidae) ; D, of Leptis scolopacea (Leptidae) ; E, of Dolichopus undulatus (Doli-
chopidae) ; F, of Volucella bombylans (Syrphidae). (After Wandolleck.)

rather a variety of intermediate conditions, associated in the
series Brachycera (Fig. 213, B to D).' Here there are three
(sometimes one or two) segments and a terminal appendage,
but the appendage is usually compound (often so distinctly com-
pound that it is evidently a series of partially, or even completely,
separate joints, Fig. 215, B): the appendage in these cases is ter-
minal, that is to say it is placed, not as in the Eumyiidae on the
front af the joint that bears it, but (in the great majority of
Brachycera) at the tip thereof; this appendage is often conical
and pointed, often hair-like. Exceptional forms of antenna are
found in the parasitic flies of the series Pupipara. In the Order
generally the two basal joints of the antennae are evidently
distinct in function from the others, and form the “scape”; the

1 It may be well to remark that this name was formerly applied to all Diptera
except Nemocera.

442 DIPTERA CHAP.

part of the antenna beyond the scape is called the “ flagellum ” ;
an appendage of the flagellum is called “arista” when bristle-
like, when thicker “style.” In the basal joint of the antenna
there is a complex nervous structure known as Johnston’s organ.
It is specially well developed in Culex and Chironomus, and is
larger in the male than it is in the female. Child has found
something of the kind present in all the Diptera he has
examined, and he considers that an analogous structure exists in
Insects of other Orders. He thinks it is concerned with the
perception of vibration, there being no sharp distinction between
auditory and tactile sensation."

About one-half of the Diptera possess a peculiar structure
in the form of a head-vesicle called “ptilinum.” In the fly
emerging from the pupa this appears as a bladder-like expansion
of the front of the head; being susceptible of great distension, it
is useful in rupturing the hard shell in which the creature is then
enclosed. In the mature fly the ptilinum is completely intro-
verted, and can be found only by dissection; a little space, the
“Junula,” just under an arched suture, extending over the point
of insertion of the antenae remains, however, and offers evidence
of the existence of the ptilinum. This structure is also of
importance in classification, though, unfortunately, it is difficult
to verify. |

No point of Insect morphology has given rise to more differ-
ence of opinion than the mouth of Diptera; and the subject is
still very far from being completely understood. The anatomy
and morphology of the mandibulate Insect-mouth are compara-
tively simple (though not without greater difficulties than are
usually appreciated) ; and it has been the desire of morphologists
to homologise the sucking mouth of Diptera with the biting
mouth; hence the view that the appendages of three segments
are separate and distinct in the fly’s mouth is taken for granted,
and it is further assumed that some of the secondary parts of the
appendages of the biting mouth can also be recognised in the
sucking mouth. The anatomy of the mouth-parts is, however,

' Zool. Anz. xvii. 1894, p. 35, and Ann. Nat. Hist. (6) xiii. 1894, p. 3872 ; Zeitschr.
wiss. Zool. lviii. 1895, p. 475.

2 Cf. Osten Sacken, Berlin. ent. Zeitschr. xxxviii. 1893 ; and Becher, Wien. ent.
Zeit. 1. 1882, p. 49. For an account of the condition, with diagrammatic figures,
of the fly emerging from the pupa, ef. Sasatti, J. Coll. Japan, i. 1887. p. 34;
pl. vi.

VI MOUTH-PARTS 443

subject to great diversity of structure within the limits of the
Order itself, even the two sexes in some species differing pro-
foundly in this respect. In the majority of the family Oestridae
the mouth-parts are practically absent, and no definite entry to
the alimentary canal can be perceived (Fig. 245). Besides this
condition and its antithesis (Fig. 214), the complex assemblage
of lancets seen in the Breeze-flies that draw blood, there is a
great variety of other anatomical conditions.

Fic. 214.—Mouth - parts of
the common blood-suck-
ing fly, Maematopota
pluvialis 9. A, Viewed
from beneath, the pro-
boscis removed; 4a,
labrum; 6, 6, cultelli
(mandibles of other
anatomists) ; c, ¢, scal-
pella (maxillae of other
anatomists) ; d, part of
ventral scutum of second
metamere ; @, é,,f,,f, parts
of palpi ; g, hypopharynx
and pellucid — salivary
duct ; h, salivary recep-
tacle ; 7, salivary duct ;
k, membranous part of
second metamere; J,
pharynx: B, labrum,
pharynx, hypopharynx,
separated, seen from
beneath ; @, labrum; 3,

hypopharynx ; ¢, salivary duct; d, pharynx; e, protractor muscles: ©, proboscis

(labium) from beneath ; a, scutum proboscidis ; ¢, ¢, labella; d, d, retractor muscles.

(After Meinert).

Although, as we have said, great diversity of opinion
exists, yet on the whole the majority of Dipterologists accept a
view something to the following effect :—the labrum, or the
labrum combined with the epipharynx, is frequently much pro-
longed ; the tongue—hypopharynx—nmay also be much prolonged,
and may form, in apposition with the labrum, a more or less im-
perfect tube for ingestion of the nutriment; the labium is more
or less membranous or fleshy, and acts as a sheathing organ, its
tips—called labella—being in some cases developed to a quite
extraordinary extent. As to the other parts of the mouth there
is less agreement; the pointed organs (Fig. 214, A, b b) are by

1 It is frequently said that one sex of a single species may be dimorphic in this
respect, but we shall subsequently mention (in Blepharoceridae) that this is not
yet sufficiently established.

444 DIPTERA CHAP.

many identified as mandibles, while another pair of pointed pro-
cesses (¢ ¢) are considered to be parts of a maxilla, and the palpi
(ff) are by some considered to be maxillary palps. The Danish
entomologist, Meinert, has published the best anatomical deserip-
tion of many of the diverse kinds of Dipterous mouth.’ He,
however, takes a different view of the morphology; he considers
that not only may parts of the appendages of the mouth be much
modified during the early stages of the individual development,
but that they may be differently combined, even parts of the
appendages of two segments being brought together in intimate
combination. He has also pointed out that the mandibulate and
sucking mouth are mechanical implements constructed on opposed
principles; the main object of a biting mouth being the fixing
and perfecting of the articulations of the mouth, so that great
power of holding may be attained with a limited but definite
power of movement. In the sucking mouth the parts are in-
timately associated for simple protrusion. Hence the two kinds
of mouth must have been distinguished very early in ‘the
phylogeny, so that we must not expect to find a great corre-
spondence between the parts of biting and sucking mouths.
He apparently also considers that not only the appendages of
a head-segment, but also part of the body of the segment, may
be used in the construction of the mouth-organs. Meinert’s
views allow a much greater latitude of interpretation of the parts
of the Dipterous mouth; had he contented himself with enun-
ciating them in the manner we have followed him in summarily
describing, they would have been recognised’ as a formidable
obstacle to the facile adoption of the ordinary views. He has,
however, accompanied his general statement with a particular
interpretation and a distinct nomenclature, neither of which is it
possible to adopt at present, as they have no more justification
than the ordinary view. So that instead of one set of doubtful
interpretations we have two.” In so difficult a question as homo-
logising the trophi of different Orders of Insects we ought to use

1 Fluernes Munddele, Copenhagen, 1881, 91 pp. 6 plates ; Ent. Tidskr. i. 1879,
p- 150; Becher having given (Denk. Ak. Wien. xlv. 1882, p. 123) an interpreta-
tion different from that of Meinert, this author set forth his general views in
Zool. Anz. v. 1882, pp. 570 and 599.

* The reader should not suppose that there are only two views as to the Dipterous
mouth, for actually there are several ; our object is here only to give a general idea
of the subject.

VII STRUCTURE AAs

exhaustively every method of inquiry: and from this point of
view the development is of great importance. This has, however,
as yet thrown but little light on the subject, this study being a
very difficult one owing to the profound changes that take place
during metamorphosis, the diversity of the parts in the early
stages of Diptera, and the possibility that the larval conditions
may themselves have been greatly changed in the course of the
phylogeny. Miall informs us, however, that in Chironomus as
well as in Corethra the new parts of the mouth of the imago are
developed within those of the larva.’ This may permit of an
identification of the main divisions of the mouth, at any rate in
these cases. Lowne has to some extent traced the development
in the blowfly, and he does not agree with the usual interpreta-
tion of the parts in the adult.

The mouth is of considerable importance in the classification
of Diptera. The Nemocera are remarkable from the linear de-
velopment and flexibility of the palpi, which are nearly always at
least three- or four-jointed ; this condition occurring in no other
Diptera. The palpi attain an extraordinary development in some
Culicidae; in the genus Megarrhina they are nearly as long as
the body, and project in front of the head after the fashion of
the palpi of Lepidoptera. In the Brachycera the sclerites or
hard parts of the mouth reach a maximum of development, and
in Tabanidae (Fig. 214), Nemestrinidae and Bombylidae are
often quite disproportionate to the size of the Insect. In many
of the Eumyid flies the soft parts are greatly developed, and
capable of a variety of movement, the proboscis as a whole being
protrusible, and having an elbow-joint in the middle.

The thorax is remarkable from the absence of distinct separa-
tion into the three divisions that may usually be so easily dis-
tinguished in Insects. The perfect combination of the three
segments adds much to the difficulty of arriving at general con-
clusions as to the identification of the parts; hence considerable
difference of opinion still prevails. It was formerly supposed
that a segment from the abdomen was added to the thorax of
Diptera as it is in Hymenoptera, but this has been shown by
Brauer to be erroneous. Indeed, according to Lowne, the abdo-
minal cavity 1s increased by the addition of the small posterior
area of the thorax ; it being the mesophragma that separates the

1 Tr. Linn. Soc. London (2).v. 1892, p. 271.

446 DIPTERA CHAP.

second and third great divisions of the body-cavity. The pro-
thorax is always small, except in a few of the abnormal wingless
forms (Melophagus); in Nycteribia (Fig. 248) the mesothorax
forms the anterior part of the body; the head and such parts of
the prothorax as may be subsequently discovered to exist being
placed entirely on the dorsum of the body. The mesothorax in
all the winged Diptera forms by far the larger portion of the
thoracic mass, the prominent part of it, that projects backwards
to a greater or less extent over the base of the abdomen, being
the scutellum. The first or prothoracic stigma is remarkably large
and distinct, and is by some called mesothoracic. Another large
stigma is placed very near to the halter (or balancer); the meta-
thorax being very small. An imperfect stigma is said by Lowne
to exist in the blowfly near the base of the wing. The number
of abdominal segments externally visible is very diverse ; there may
be as many as nine (in the male 7ipula), or as few as five, or even
four, when the basal segment is much concealed; the diminu-
tion is due to certain segments at the extremity being indrawn
and serving as a sort of tubular ovipositor in the female, or curled
under the body and altered in form in the other sex, so as to
constitute what is called a “hypopygium.” In the female of
Tipulidae the body is terminated by some horny pieces forming
an external ovipositor. In nearly all Diptera the feet are five-
jointed; the claws are well developed, there being placed under
each of them a free pad or membrane, the “ pulvillus”; there may
be also a median structure between each pair of claws, of diverse
form, the “ empodium.”

On the surface of the body of many flies there will be seen
an armature of pointed bristles; these flies are called “ chaeto-
phorous”; where no regularly arranged system of such bristles
exists the fly is “eremochaetous.” In some familes the arrange-
ment of these bristles is of importance in classification, and a
system of description has been drawn up by Baron Osten
Sacken: this branch of descriptive entomology is known as
chaetotaxy.’

The wings are of great importance in classifying Diptera ;
but unfortunately, like the other parts, they have not received an
exhaustive anatomical study, and Dipterologists are not agreed
as to the names that should be applied to their parts. We give

1 Tr. ent. Soc. London, 1884, p. 497.

VII WINGS 447

below. figures of two systems that have been used by eminent
Dipterologists for the description of the nervures and cells. The
comprehension of these features of the Dipterous wing will be facil-
tated by noticing that the wing—being extended at right angles to
the body—is divided by the longitudinal nervures into two great
fields, anterior and posterior, with an interval. between them :
this interval is traversed only by a short cross-vein (marked x in
Fig. 215 A,andiin B). This cross-vein may be placed near the
base or nearer to the tip of the wing; it is of importance because
no nervure in front of the median area traversed by it can corre-

A Fic. 215.—Nervuration of Dipterous

wing. A, Wing of a Tipulid, accord-
ing to Loew, who uses the following
nomenclature : a, costal nervure ; J,
mediastinal ; c, subcostal ; d, radial ;
e, cubital ; 7, discoidal ; g, postical ;
h, anal; 7, axillar; «x, transverse,
y, posterior transverse, nervure ;
1, 2, mediastinal areas; 38, sub-
costal ; 4, cubital ; 5, anterior basal ;
6, posterior basal; 7, anal; 8, pos-
terior marginal; 9, discoidal. B,
Wing of an Acalypterate Muscid
(Ortalis), according to Schiner, who
uses the following nomenclature :
(nervures, small letters ; cells, capi-
tal letters): a, transverse shoulder ;
6, auxiliary ; ¢ to h, first to sixth
longitudinal ; 7, middle transverse ;
k, posterior transverse ; 7, m, 7, 0,
costa ; p, anterior basal transverse ; g, posterior basal transverse ; 7, rudiment of a fourth
nervure ; s, axillary incision: A, B, C, first, second, and third costal cells ; D, marginal ;
E, sub-marginal ; F, G, H, first, second, and third posterior ; I, discal; K, L, M, first,
second, and third basal cells ; N, anal angle; O, alula.

spond with a nervure placed behind it in another wing. The
very different nature of the nervuration in the two wings we have
figured will readily be appreciated by an inspection of the parts
posterior to the little cross-vein. On the hind margin of the
wing, near the base, there is often a more or less free lobe (Fig.
215, B, O) called the “ alula”: still nearer to the base, or placed on
the side of the body, may be seen one or two other lobes, of which
the one nearer the alula is called the “tegula,” or (when a
lobe behind it is also present) the “upper tegula,” (the “anti-
tegula” of Osten Sacken); the other being the “lower tegula.”
These two terms are erroneous, the word tegula being definitely
applied to another part of the Insect-body. In speaking of this
structure in the following pages, we have preferred to call it the

448 DIPTERA CHAP,

“squama.”' Those Muscidae in which the squama covers the

halter like a hood are called“ calypterate.” In Fig. 216, we
represent these structures, and in
the explanation have mentioned the
synonyms, The terms we think
most applicable to the three lobes
are alula, antisquama, squama. The
squama may be called “calypter”
when it covers the halter.

The halteres—commonly called
Fia. 216. —Parts at the base of the balancers or poisers—are perhaps

wing in Calliphora. a, Anal Ds

angle or lobe of the wing; 6, the most characteristic of all the

alula; ¢, antisquama, squama T)ipterous structures, though they are

alaris, or antitegula ; d, squama, : 3

squama thoracicalis, tegula, caly- absent in most of the few wingless

pler,;or ealypiron's 6 posteriat forme or the Order. a Outside thei:

extremity (scutellum) of the Cota

mesothorax ; f, scutum of meso- tera similar organs appear to exist

ores only in male Coccidae. The pair of
halteres is placed on the metathorax, one on each of the pleural
regions. They are believed to be the homologues of the hind
wings; Weinland states * that certain canals existing in the interior
of the halter correspond to wing-nervures. The halter may be
described as a small rod-like body with a head hke a pin, this
terminal part being, however, rather variable in form. We
have already stated that in many Diptera the squama forms a
hood, the position of which leads to the belief that it is an
important adjunct to the halter. Although the exact functions
of the halteres are far from clear, it is certain that they are
highly complex bodies, of extremely delicate structure: they are
doubtless sense-organs, possessing as they do, groups of papillae
on the exterior and a chordotonal organ (a structure for assist-
ing the perception of sound) in the basal part; each halter
is provided with four muscles at the base, and can, like the
wings, execute most rapid vibrations. Seeing that they are the
homologues of wings, it is a remarkable fact that in no Diptera
are they replaced by wings, or by structures intermediate between

these two kinds of organs.
Internal Structure.—Information about the internal anatomy
1 Osten Sacken, although making use of the terms tegula and antitegula,

suggested the propriety of using squama and antisquama, as we have done,
2 Zeitschr. wiss. Zool. li. 1891, p. 55.

VII ANATOMY—LARVAE 449

is by no means extensive. The tracheal system is highly developed,
and has air-sacs connected with it; a large pair at the base of
the abdomen being called aérostats by Dufour. Inside the
thoracic spiracles there are peculiar structures supposed by some
to be voice-organs, while the abdominal spiracles are said to be
remarkably simple in structure. Lowne says that there are ten
or eleven pairs of spiracles in the Blow-fly ; one of these, near
the base of the wing, is peculiar in structure, and may not be
a true stigma; he calls it a tympanic spiracle; 1t seems doubtful
whether there are more than seven abdominal pairs. The alimen-
tary canal is very elongate, and is provided with a diverticulum,
the crop; this is usually called the sucking stomach, though its func-
tion is extremely doubtful. The Malpighian tubes are four in
number, and are very elongate; in several groups of Nemocera
there are, however, five Malpighian tubes, a number known to
occur in only very few other Insects. The nervous system is
remarkable on account of the concentration of gangha in the
thorax, so as to forma thoracic, in addition to the usual cephalic,
brain. For particulars as to the positions of the ganglia and the
great changes that occur in the lifetime, the student should
refer to Brandt, to Kiinckel, and to Brauer.1 Much information
as to the internal anatomy of the Blowfly is given by Lowne, but
it is doubtful to what extent it is applicable to Diptera in
general.”

The larvae of Diptera are—so far as the unaided eye is
concerned—without exception destitute of any kind of adorn-
ment, the vast majority of them being of the kind known as
maggots. None of them have
true thoracic legs; though in
the earlier groups, pseudopods
or protuberances of the body
that serve as aids in locomo- Fre, 217.—Acephalous larva or maggot of the
tion are common. Unlike eal with the head, a, extended. (After
what occurs in other Orders
the arrangement of these pseudopods on the body differs greatly
in various forms; in a few cases they are surmounted by

1 Brandt, Horae Soc. ent. Ross. xiv. 1878, p. vil.; xv. 1879, p. 20. Brauer,
Denk. Ak. Wien, xlvii. 1883, pp. 12-16. Kiinckel, C.R. Ac. Paris, \xxxtx. 1879,
p- 491.

2 Blow-fly, 1895: in two vols. For Anatomy of Volucelia, see Kiinckel
d’Herculais, Recherches sur Vorg. des Volucelles, Paris, 1875 and 1881.

VOL. VI 2G

450 \. -DIPTERA CHAP,

curved hairs. The most important distinction in external form
in Dipterous larvae is that while those that are thorough maggots
possess no visible head, others have a well-marked one (Fig.
225); these are therefore called “eucephalous”: they have a
mouth of the mandibulate type. In some other Dipterous larvae
the head is more or less reduced in size, and in the acephalous forms
there is only a framework of a few chitinous rods to represent
it. The nervous system in the most completely headless larvae
is very remarkable, all the ganglia being concentrated in a single
mass placed in the thorax. The tracheal system exhibits a great
variety ; some larvae have stigmata arranged along the sides of
the body after the fashion normal in Insect-larvae; these are
called “ peripneustic ” ; as many as ten pairs of stigmata may be
present in these cases, but nine pairs 1s much more common.
Other larvae have a pair of stigmata placed at the termination of
the body, and another pair near the anterior extremity, the two
pairs communicating by large tracheal trunks extending the
length of the body; these larvae are said to be “ amphipneustic ” :
this is the condition usual in the more completely acephalous
larvae. Others have only the terminal pair of spiracles, and are
styled “ metapneustic.” Some begin life in the metapneustic state
and afterwards become amphipneustic. In the aquatic larva
of Corethra there are no spiracles, though there is an imperfect
tracheal system. Many Dipterous larvae that live in water
or in conditions that prevent access of air to the body have
remarkable arrangements for keeping the tip of the body in
communication with the atmosphere. The stigmata in meta-
pheustic and amphipneustic larvae are very remarkable compound
structures, exhibiting however great diversity ; their peculiarities
and uses are not well understood; it appears very doubtful
whether some of them have any external opening. Reference
may be made, as to the variety of structure, to Meijere’s paper ’
from which we take the accompanying figure of a posterior
stigmatic apparatus in Lipara lucens. It appears that there is a
compound chamber—* Filzkammer ”—terminating externally in
lobes or fingers —“ Knospen” and appearing as marks on the
outer surface: this chamber is seated on a tracheal tube, and is,
Meijere thinks, probably a secondary growth of the trachea
coming to the outer surface. It is traversed by what may be
1 Tijdschr. Ent. xxxviii. 1895, pp. 65-100.

VII LARVA——PUPA

451

considered the original tracheal tube, opening externally as an

external stigmatic scar—‘*Stigmennarbe ”

or inner scar placed internally. We may
conclude from what is already known
that these structures will be found to
differ in the same larva according to the
stage of its development.

An extremely valuable summary of
the characters and variety of Dipterous
larvae has been given by Brauer,’ from
which it appears that the larvae of the
first half of the family exhibit great
variety and have been much studied,
while the more purely maggot-like forms
of the Muscidae have, with one or two
exceptions, been little investigated.

The pupal instar is of two distinct
kinds. First, we meet with a pupa like that

and with a second

Ges

Fie. 218.—The posterior

stigma of the larva of Lip-
ara lucens. a, One of the
three “ Knospen”’ or lobes ;
b, external stigmatic scar ;
c, internal scar; d, stig-
matic chamber (Filzkam-
mer); é, trachea. (After
Meijere.)

of Lepidoptera, viz. a mummy-lke object,

or pupa obtecta, in which there is a crisp outer shell, formed
in part by the adherent cases of the appendages of the future
imago. This condition, with a few exceptions to be subsequently
noticed, obtains in the Nemocera and Brachycera. It is exhibited
in various degrees of perfection, being most complete in Tipulidae :
in other forms the shell is softer and the appendages more pro-
tuberant. The second kind of pupa is found in the Cyclor-
rhaphous flies; it has externally no marks except some faint
circular rings and, frequently, a pair of projections from near
one extremity of the body; occasionally there is a single pro-
minence at the other extremity of the body. This condition is
due to the fact that the larva does not escape from the skin
at the last ecdysis, but merely shrinks within it, so that the
larval skin, itself contracted and altered by an excretion of
chitin, remains and forms a perfect protection to the included
organism. This kind of pupa looks like a seed, and is well
exemplified by the common Blow-fly. The capacity for entering
on such a condition is evidently correlative with the absence of a
larval head. The metamorphosis in this curious little barrel
goes on in a different manner to what it dces in the pupa

1 Denk. Ak. Wien, x\vii. 1883, pp. 1-100, pls. i.—v.

452 DIPTERA CHAP.

obtecta. A good name for the whole structure of this instar
has not been found. — Older authors called it “pupa coarctata,”
or “nympha inclusa”; Brauer speaks of it as a “compound
pupa”; ordinarily in our language it is called a “puparium,” a
term which is more applicable to the case alone.

In species having a pupa obtecta the larval skin is cast after
the chief processes of the external metamorphosis have occurred,
and then an exudation of chitin hardens the general surface.
In the “compound pupa” of the Blow-fly there is for a consider-
able period no formed pupa at all, but merely a shell or case
containing the results of histolysis and the centres for regenera-
tion of new organs; the chitin-exudation to the exterior of the
larval skin occurs in the early part of the series of metamorphic
changes, and the organism breaks down to a cream within the shell
thus formed, and then gradually assumes therein the condition of a
soft, nymphoid pupa. The exceptional conditions previously re-
ferred to as exhibited by a few forms are certain cases in which a
more or less perfect pupa obtecta is found within the last larval
skin, as is the case in Stratiomys. Another highly remarkable
condition exists in the Hessian fly,and a few other Cecidomyids,
where the Insect apparently makes an exudation which it uses
as a covering case, independent of the larval skin; this latter
being subsequently shed inside the case, so that this condition
of coarctate pupa differs from that we have described as exist-
ing in Cyclorrhaphous flies, although the two are superficially
similar. In the Pupipara the larval stage is passed in the body
of the mother, which produces a succession of young, nourished
one at a time by the secretion of glands; this young is born as
a full-grown larva that becomes at once a pupa.

Metamorphosis.—As it is in Diptera that the phenomena
of Insect-metamorphosis have reached their highest development
we endeavoured to give some idea of their nature in the previous
volume, therefore we need give only a brief sketch of the chiet
features of Dipterous metamorphosis. The Blow-fly undergoes <
rapid embryonic development, the later stages of which are, on
the whole, of a retrogressive nature. On the emergence of the
young maggot it feeds up rapidly, the rapidity varying greatly
according to circumstances, and then when full-grown rests.
While resting, a process of internal liquefaction, called histolysis,
is going on, and the maggot contracts and exudes an excretion

vII METAMORPHOSIS 453

that hardens its skin. At the time this hard skin has become
complete, or soon after, the maggot inside has dissolved into a
cream contained in a sac inside the shell; this cream becomes
reconstituted into a fly by a gradual process of growth and
development of certain minute portions of the body—the
imaginal discs or folds, the histoblasts and neuroblasts that
were exempt from the histolytic process: in the early stages
of the reconstitution the general structure is, of course, altogether
vague, and this condition—purely one of transition—is called
the pronymph; the nymph becomes gradually developed: it
corresponds vaguely with the pupa obtecta of the early groups
of Diptera, but is soft like the pupa of Hymenoptera. This
nymph gradually develops into the fly itself, the external
parts being first completed and the internal organs elaborated
subsequently. The sexual organs do not undergo metamorphosis
lke other internal organs, there being a gradual (though ir-
regular or interrupted) growth of them in the young larva, till
they are completed some time after the emergence of the perfect
fly. The processes in the Blow-fly have been studied by numerous
able histologists of various nationalities, and have recently been
described by Lowne in our own language.’ Comparatively little
has been done in studying the corresponding phenomena in other
Diptera. Weismann has investigated the development of Corethra,
and Miall that of Chironomus. These two flies belong to a division
of Diptera different from that which includes the Blow-fly, and
they display a condition of the metamorphic processes allied to
what occurs in Lepidoptera, as well as to that which takes place in
the Blow-fty. Imaginal folds are formed, but they only appear
much later in the life, and they are much less distant from the
positions they will, when developed, occupy in the imago. In
Chironomus, according to Miall, the imaginal folds only appear
in the last larval instar, but they grow with such rapidity that
the legs and wings of the future fly can be distinguished in the
larva, even before pupation ; thus when the activity of the larva
ceases but lttle change is required to complete the obtected
pupa. In the Blow-fly some of the imaginal folds have been

1 Since our brief and imperfect sketch of metamorphosis appeared in Vol. V. of
this series, Packard has treated the subject more fully in his Text-book.of Entomology,
New York, 1898 ; and Pratt has summarised the state of knowledge as to imaginal
dises in Psyche, viii. 1897, p. 15, ete.

454 DIPDERA CHAP.

traced back to the embryo; how many centres for the new
growth there may be is uncertain, for though there are upwards
of sixty for the outer body, the number of regenerative centres
for the internal organs is not ascertained. The peculiar central
nervous mass, mentioned in our remarks on the larva, consists of
two kinds of tissue mixed together in a complex manner; one
of these kinds is functionally active during the larval life and
at the metamorphosis undergoes histolysis, while the other, or
embryonic, portion develops into the nervous system of the fly.
It forms no part of our task to deal with general subjects,
but we may be pardoned for calling attention to the bearing the
metamorphosis of the higher Diptera has on our ideas of heredity
in Insects. The fly bears no resemblance whatever to the larva,
and is only obtained by the organic destruction of the latter,
which occurs before the perfection of the sexual organs takes
place, and yet the fly reproduces itself only secondarily, but
primarily gives rise to the totally different larva. It is supposed
that the larval structures have been gradually acquired, and yet
they are transmitted with the utmost faithfulness by the totally
different fly. We can only conclude that that which is bequeathed
in each species is the early state of a particular process of develop-
ment from which the subsequent stages follow necessarily if the
developing organism be placed in conditions having on it in-
fluences like to those that influenced the ancestors.
Classification ——The classification of Diptera is as yet very
imperfect. Formerly they were divided into two great groups,
Nemocera and Brachycera, according to the structure of the
antennae, as previously mentioned. This division has been
abandoned, and the term Brachycera is now applied to only a
small part of the old section that bore the name. The primary
division usually adopted at present is into Orthorrhapha and
Cyclorrhapha. The characters of these two groups are based
on the nature of the metamorphosis, and have been gradually
elaborated by Brauer in various memoirs.' The Orthorrhapha
includes the forms with obtected pupae, the Cyclorrhapha those
with a nymph-compound, as previously described. This distinction
is of great importance, but unfortunately it is difficult to apply
to the fly itself; the only character that can be used in connec-

1 Monograph of Oestridae, Verh. Ges. Wien, 1863, and other papers op. cit.
1864, 1867, 1869 ; also Denk. Ak. Wien, xlii. 1880, xlvii. 1883.

vil CLASSIFICATION ass

tion with the imago is the existence of a suture over the insertion
of the antennae in a portion, but not all, of the Cyclorrhapha.'
The next set of divisions used by Brauer divides the Order into
four sections, viz. 1. Orthorrhapha Nematocera, 2. O. Brachycera,
3. Cyclorrhapha Aschiza, 4. C. Schizophora. As these four
eroups are recognised more readily than the two major groups
the student will do well at first to disregard the primary division
and consider the Diptera as divisible into four great groups. To
these four divisions we, however, add temporarily a fifth, viz.
Pupipara. This is included by Brauer in Schizophora, but it
appears to be really an unnatural complex, and had better be
kept separate till it has been entirely reconsidered. These
ereat sections may be thus summarised :—

Series 1, Orthorrhapha Nemocera.— Antennae with more than 6 segments,
not terminated by an arista; with the segments of the flagellum
more or less similar to one another. Palpi slender and flexible,
four- or five-jointed.?

Series 2. Orthorrhapha Brachycera. — Antennae variable, but never truly
Nemocerous nor like those of Cyclorrhapha; when an arista is
present it is usually placed terminally, not superiorly ; when an
arista is not present the flagellum terminates as an appendage
consisting of a variable number of indistinctly separated segments ;
thus the flagellum is not composed of similar joints ; [rarely are the
antennae as many as seven-jointed], Palpi only one- or two-
jointed? Around the insertion of the antennae there is no definite
arched suture enclosing a small depressed space. The nervuration
of the wings is usually more complex than in any of the other
divisions.

Series 3. Cyclorrhapha Aschiza. Antennae composed of not more than three
joints and an arista; the latter is not terminal. Front of head
without definite arched suture over the antennae, but frequently
with a minute area of different colour or texture there. This group
consists of the great family Syrphidae, and of four small families,
viz. Conopidae, Pipunculidae, Phoridae, and Platypezidae. The
section is supposed to be justified by its being Cyclorrhaphous in
pupation, and by the members not possessing a ptilinum (or having
no trace of one when quite mature). The Syrphidae are doubtless

1 Becher, Wien. Ent. Zeit. i. 1882, p. 49; for observation on connecting forms
see Brauer, Verh. Ges. Wien, xl. 1890, p. 272.

2 The palpi are said to be of only one segment in some genera of Cecidomyiidae.
The Cecidomyiidae are easily distinguished by the minute size—body not more
than a line long—and by there not being more than six nervules at the periphery
of the wing. Aédes (Culicidae) has also short palpi.

3 It is said by Schiner that in the anomalous genus Nemestrina the palpi are
of three segments.

456 DIPTERA CHAP.

a natural group, but the association with them of the other
families mentioned is a mere temporary device. The greatest
difficulty is experienced in deciding on a position for Phoridae, as
to which scarcely any two authorities are agreed. :

Series 4. Cyclorrhapha Schizophora, or Eumyiid flies. The antennae consist
of three joints and an arista. In the Calyptratae the frontal suture,
or fold over the antennae, is well marked and extends downwards
along each side of the face, leaving a distinct lunule over the
antennae. In the Acalyptrate Muscids the form of the head and
of the antennae vary much and are less characteristic, but the
wings differ from those of Brachycera by their much less complex
nervuration.

Series 5. Pupipara. These are flies of abnormal habits, and only found in
connection with living Vertebrates, of which they suck the blood
(one species, Braula caeca, lives on bees). Many are wingless,
or have wings reduced in size. The young are produced alive,
full grown, but having still to undergo a metamorphosis. This
group consists of a small number of flies of which some are
amongst the most aberrant known. This is specially the case with
the Nycteribiidae. This Section will probably be greatly modified,
as it is far from being a natural assemblage.!

The Sub-Order Aphaniptera, or Fleas, considered a distinct Order by many
entomologists, may for the present be placed as a part of Diptera.

It must be admitted that these sections are far from satis-
factory. Brauer divides them into Tribes, based on the nature
of the larvae, but these tribes are even more unsatisfactory than
the sections, hosts of species being entirely unknown in the
larval state, and many of those that are known having been very
inadequately studied. We must admit that the classification of
Diptera has at present advanced but little beyond the stage of
arranging them in natural families capable of exact definition.
We may, however, draw attention to the attempt that is being
made by Osten Sacken to remodel the classification of the Nemo-
cera and Brachycera by the combination of families into super-
families.” He proposes to divide the Nemocera into two super-
families: 1. Nemocera Vera, including all the families from
Cecidomyiidae to’ Tipulidae ; 2. Nemocera Anomala, consisting of
the small families Bibionidae, Simuliidae, Blepharoceridae, Rhy-
phidae and Orphnephilidae.

For Orthorrhapha Brachycera he adopts the following arrange-

1 For tables of the families of flies the student may refer to Loew, Smithson-
Misc. Coll. vi. Art. i. 1862; to Brauer, Denk. Ak. Wien, xlii. 1880, p. 110
(Orthorrhapha only); to Williston, Manual of N. American Diptera, 1896; to

Schiner, Fauna austriaca, Diptera, Vienna, 1860, etc.
2 Berlin, ent. Zeitschr. xxxvii. 1892, p. 365, and xli. 1897, p. 365.

vr BLOOD-SUCKING FLIES 457

ment: 1: Super-family Eremochaeta, for Stratiomyidae, Tabanidae,
Acanthomeridae and Leptidae ; 2. Tromoptera, for Nemestrinidae,
Acroceridae, Bombylidae, Therevidae, and Scenopinidae ; 3. Ener-
gopoda, for Asilidae, Dolichopidae, Empidae and Lonchopteridae,
Phoridae being included with doubt; 4. Mydaidae remains isolated.

This classification is based on the relations of the eyes and
bristles of the upper surface, and on the powers of locomotion,
aérial or terrestrial. At present it is not sufficiently precise to
be of use to any but the very advanced student.

Blood-sucking Diptera.—The habit of blood-sucking from
Vertebrates is, among Insects, of course confined to those with
suctorial mouth, and is exhibited by various Diptera. It is,
however, indulged in by but a small number of species, and
these do not belong to any special division of the Order. It is
remarkable that as a rule the habit is confined to the female sex,
and that a large proportion of the species have aquatic larvae.
This subject has many points of interest, but does not appear to
have yet received the attention it merits. We give below a
brief summary of the facts as to blood-sucking Diptera.

Series I. Nemocera.—In this section the habit occurs in no less
than five families, viz. :

Blepharoceridae. Curupira ; in the female only ; larva aquatic.

Culicidae. Culex, Mosquitoes; in the female only; other
genera, with one or two exceptions, do not suck blood;
larvae aquatic.

Chironomidae. Ceratopogon, Midge; in the female only; ex-
ceptional even in the genus, though the habit is said to
exist in one or two less known, allied genera; larval habits
not. certain ; often aquatic; in C. bipwnetatus the larva lives
under moist bark.

Psychodidae. Phlebotomus: in the female only (?); quite ex-
ceptional in the family; larva aquatic or in hquid filth.

Simuliidae. Stmuliwm, sand-flies; general in the family (7),
which, however, is a very small one; larva aquatic, food
probably mixed vegetable and animal microscopic organisms.

Series II. Brachycera. Tabanidae. Gad-flies: apparently general
in the females of this family; the habits of the exotic forms
but little known; in the larvai state, scarcely at all known;
some are aquatic.

458 DIPTERA CHAP.

Series IV. Cyclorrhapha Schizophora: Stomoxys, Haematobia ;
both sexes (7); larvae in dung. [The Tse-tse flies, Glossina,
are placed in this family, though their mode of parturition is
that of the next section].

Series V. Pupipara. The habit of blood-sucking is probably
common to all the group and to both sexes. The flies, with
one exception, frequent Vertebrates; in many cases living
entirely on their bodies, and apparently imbibing much blood ;
the larvae are nourished inside the flies, not on the imbibed
blood, but on a milky secretion from the mother.

Sub-Order Aphaniptera. Fleas. The habit of blood-sucking is
common to all the members and to both sexes. The larvae
live on dried animal matter.

Fossil Diptera— A considerable variety of forms have been
found in amber, and many in the tertiary beds; very few members
of the Cyclorrhaphous Sections are, however, among them; the
Tipulidae, on the other hand, are richly represented. In the
Mesozoic epoch the Order is found as early as the Lias, the
forms being exclusively Orthorrhaphous, both Nemocera and
Brachycera being represented. All are referred to existing
families. Nothing has been found tending to connect the
Diptera with other Orders. No Palaeozoic Diptera are known.

Series 1 Orthorrhapha Nemocera

Fam. 1. Cecidomyiidae.— An extensive family of very minute
and fragile flies, the wings of which have very few nervures; the
antennae are rather long, and are furnished with whorls of hair.
In the case of some species the antennae are beautiful objects;
in Aylodiplosis some of the hairs have no free extremities, but
form loops (Fig. 220). In the males of certain species the
joints appear to be double, each one consisting of a neck and a
body. Although comparatively little is known as to the flies
themselves, yet these Insects are of importance on account of
their preparatory stages. The larvae have very diverse habits ;
the majority lve in plants and form galls, or produce defor-
mations of the leaves, flowers, stems, buds, or roots in a great
variety of ways; others live under bark or in animal matter ;
some are predaceous, killing Aphidae or Acari, and even other

VII CECIDOMYIIDAE 459

Cecidomyids. The North American Diplosis resinicola lives in
the resin exuded as the results of the attacks of a caterpillar.
The larva burrows in the semi-liquid resin, and, according to
Osten Sacken,' is probably amphipneustic. Cecidomyiid larvae
are short maggots, narrowed at the two ends, with a very small
head, and between this and the first thoracic segment (this bears a
stigma), a small supplementary segment ; the total number of seg-
ments is thirteen, besides the head; there are eight pairs of stigmata

Fic. 219.—Cecidomyia (Diplosis) buzi. Fic. 220.—One segment of an-
Britain. A, Larva, magnified ; B, tenna of Xylodiplosis sp. ; a,
pupa; C, imago; D, portion of an- Tip of one segment ; 5, base of
tenna. (After Laboulbene.) another. (After Janet.)

on the posterior part of the body. Brauer defines the Cecido-
myiid larva thus, “ peripneustic, with nine pairs of stigmata, the
first on the second segment behind the head; two to nine on fifth
to twelfth segments; body as a whole fourteen - segmented
without a fully-formed head.” The most remarkable peculiarity
of Cecidomyiid larvae is that those of many species possess a
peculiar organ—called breast-bone, sternal spatula, or anchor-
process—projecting from the back of the lower face of the pro-
thoracic segment. The use of so peculiar a structure has been
much discussed. According to Giard, in addition to the part

1 Tr. Amer. ent. Soc. iii. 1871, p. 345. 2 Bull. Soc. ent. France, 1893, p. 1xxx.

460 DIPTERA CHAP.

that protrudes externally, as shown in Fig. 219, A, there is a longer
portion concealed, forming a sort of handle, having muscles
attached to it. Some of these larvae have the power of executing
leaps, and he states that such larvae are provided on the terminal
segment with a pair of corneous papillae; bending itself almost
into a circle, the larva hooks together the breast-bone and
the papillae, and when this connection is broken the spring
occurs. This faculty is only possessed by a few species, and it
is probable that in other cases the spatula is used as a means for
changing the position or as a perforator. Some of the larvae
possess false feet on certain of the segments. Williston says
they probably do not moult. In the pupal instar (Fig. 219, B), the
Cecidomyiid greatly resembles a minute Lepidopterous pupa.
The Hessian fly, Cecidomyia destructor, is frequently extremely
injurious to crops of cereals, and in some parts of the world
commits serious depredation. The larva is lodged at the point
where a leaf enwraps the stem; it produces a weakness of the
stem, which consequently bends. This Insect and C. tritica (the
larva of which attacks the flowers of wheat) pupate in a very
curious manner: they form little compact cases like flax-seeds ;
these have been supposed to be a form of pupa similar to what
occurs in the Blow-fly ; but there are important distinctions. The
larva, when about to undergo its change, exudes a substance from
its skin, and this makes the flax-seed ; the larval skin itself does
not form part of this curious kind of cocoon, for it may be found,
as well as the pupa, in the interior of the “ flax-seed.” Other
Cecidomyiids form cocoons of a more ordinary kind; one species,
described by Perris as living on Pinus maritima, has the very
remarkable faculty of surrounding itself, by some means, with a
cocoon of resin. Walsh describes the cocoon-forming process of
certain Cecidomyiids as one of exudation and inflation ; Williston
as somewhat of the nature of crystallisation. Some Cecidomyids
are said to possess, in common with certain other Diptera, the
unusual number of five Malpighian tubes; and Giard says that
in the larva there is only a pair of these tubes, and that their
extremities are united so as to form a single tube, which is
twisted into an elegant double loop.

Thirty years or more ago the Russian naturalist, Wagner, made
the very remarkable discovery that the larva of a Cecidomyiid
produces young; and it has since been found by Meinert and

VII CECIDOMYIIDAE 461

—

others that this kind of paedogenesis occurs in several species of
the genera MJiastor and Oligarces. The details are briefly as
follows :—A female fly lays a few, very large, eggs, out of
each of which comes a larva, that does not go on to the

perfect state, but produces in its interior young larvae that,

after consuming the interior of the body of the parent larva,
escape by making a hole in the skin, and thereafter subsist
externally in a natural manner. This larval reproduction may
be continued for several generations, through autumn, winter,
and spring till the following summer, when a generation of the
larvae goes on to pupation and the mature, sexually perfect. fly
appears. Much discussion has taken place as to the mode
of origination of the larvae; Carus and others thought they
were produced from the rudimental, or immature ovaries of the
parent larva. Meinert, who has made a special study of the
subject,’ finds, however, that this is not the case; in the repro-
ducing larva of the autumn there is no ovary at all; in the re-
producing larvae of the spring-time rudimentary ovaries or testes,
as the case may be, exist ; the young are not, however, produced
from these, but from germs in close connection with the fat-
body. In the larvae that go on to metamorphosis the ovaries
continue their natural development. It would thus appear that
the fat-body has, like the leaf of a Begonia, under certain circum-
stances, the power, usually limited to the ovaries, of producing
complete and perfect individuals.

Owing to the minute size and excessive fragility of the Gall-
midge flies it is extremely difficult to form a collection of them ;
and as the larvae are also very difficult of preservation, nearly
every species must have its life-history worked out as a special
study before the name of the species can be ascertained. Not-
withstanding the arduous nature of the subject it is, however, a
favourite one with entomologists. " The number of described and
named forms cannot be very far short of 1000, and each year sees
some 20 or 50 species added to the list. The number of unde-
scribed forms is doubtless very large. The literature of the subject
is extensive and of the most scattered and fragmentary character.

The Cecidomyiidae have but little relation to other Nemocera,
and are sometimes called Oligoneura, on account of the reduced
number of wing-nervures. Their larvae are of a peculiar type

1 Naturhist. Tidskr. (3) viii. 1874, p. 34, pl. xii.

462 DIPTERA CHAP.

that does not agree with the larvae of the allied families having
well-marked heads (and therefore called Eucephala), nor with the
acephalous maggots of Eumyiidae.

Fam. 2. Mycetophilidae—7hese small flies are much less
delicate creatures than the Cecidomyiidae, and have more nervures
in the wings ; they possess ocelli, and frequently have the coxae
elongated, and im some cases the legs adorned with complex arrange-
ments of spines: their antennae have not whorls of hair. Although
very much neglected there are probably between 700 and 1000
species known; owing to many of their larvae living in fungoid
matter the flies are called Fungus-gnats. We have more than
100 species in Britain. Hpidapus is remarkable, inasmuch as
the female is entirely destitute of wings and halteres, while
the male has the halteres
developed but the wings of
very reduced size. #. scabiei
is an excessively minute fly,
smaller than a common flea,
and its larva is said to be
very lnjurious to stored
potatoes. The larvae of
Mycetophilidae are usually
very elongate, worm - lke
maggots, but have a distinct,

Fig. 221.—Mycetobia pallipes. Britain. small head ; they are perl-
A, Larva; B, pupa; C, imago. (After Dufour.)

pheustic, having,according to
Osten Sacken, nine pairs of spiracles, one pair prothoracic, the others
on the first eight abdominal segments. They are usually worm-
like, and sometimes seem to consist of twenty segments. Some of
them have the faculty of constructing a true cocoon by some sort
of spinning process, and a few make earthen cases for the purpose of
pupation. The pupae themselves are free, the larval skin having
been shed. The Mycetophilidae are by no means completely
fungivorous, for many live in decaying vegetable, some even in
animal, matter.

The habits of many of the larvae are very peculiar, owing to
their spinning or exuding a mucus, that reminds one of snail-
slime; they are frequently gregarious, and some of them have
likewise, as we shall subsequently mention, migratory habits.
Perris has described the very curious manner in which Sezophila

VII MYCETOPHILIDAE 463

unimaculata forms its slimy tracks;’ it stretches its head to one
side, fixes the tip of a drop of the viscous matter from its
mouth to the surface of the substance over which it is to
progress, bends its head under itself so as to affix the matter to
the lower face of its own body; then stretches its head to the
other side and repeats the operation, thus forming a track on
which it glides, or perhaps, as the mucus completely envelops
its body, we should rather call it a tunnel through which the
maggot slips along. According to the description of Hudson?
the so-called New Zealand Glow-worm is the larva of Boleto-
phila luminosa ; it forms webs in dark ravines, along which it
glides, giving a considerable amount of light from the peculiarly
formed terminal segment of the body. This larva is figured as
consisting of about twenty segments. The pupa is provided
with a very long, curiously-branched dorsal structure: the fly
issuing from the pupa is strongly luminous, though no use can
be discovered for the property either in it or in the larva. The
larva of the Australian Ceroplatus mastersi is also luminous.
Another very exceptional larva is that of Epicypta seatophora ;
it is of short, thick form, like Cecidomyiid larvae, and has a very
remarkable structure of the dorsal parts of the body ; by means of
this its excrement, which is of a peculiar nature, is spread out and
forms a case for enveloping and sheltering the larva. Ultimately
the larval case is converted into a cocoon for pupation. This larva
is so different from that of other Mycetophilidae, that Perris was
at first unable to believe that the fly he reared really came from
this unusually formed larva. The larva of Mycetobia pallipes
(Fig. 221) offers a still more remarkable phenomenon, inasmuch
as it is amphipneustic instead of peripneustic (that is to say, it
has a pair of stigmata at the termination of the body and a pair
on the first thoracic segment instead of the lateral series of pairs
we have described as normal in Mycetophilidae). This larva lives
in company with the amphipneustic larva of Rhyphus, a fly of
quite another family, and the Mycetobia larva so closely resembles
that of the Rhyphus, that it is difficult to distinguish the two.
This anomalous larva gives rise, like the exceptional larva of
Epicypta, to an ordinary Mycetophilid fly.®
1 Ann. Soc. ent. France (2) vii. 1849, p. 346.
* Trans. New Zealand Inst. xxiii. 1890, p. 48.

* Osten Sacken, Berlin. ent. Zeitschr. xxxvii. 1892, p. 442; and Perris, Ann. Soc.
ent. France (2) vii. 1849, p. 202.

464 DIPTERA CHAP,

But the most remarkable of all the Mycetophilid larvae are
those of certain species of Sciara, that migrate in columns, called
by the Germans, Heerwurm. The larva of Sciara militaris
lives under layers of decomposing leaves in forests, and under
certain circumstances, migrates, sometimes perhaps in search of a
fresh supply of food, though in some eases it is said this cannot be
the reason. Millions of the larvae accumulate and form them-
selves by the aid of their viscous mucus into great strings or
ribbons, and then glide along like serpents: these aggregates are
said to be sometimes forty to a hundred feet long, five or six inches
wide, and an inch in depth. It is said that if the two ends of
one of these processions be brought into contact, they become
jomed, and the monstrous ring may writhe for many hours before
it can again disengage itself and assume a columnar form.
These processional maggots are met with in Northern Europe
and the United States, and there is now an extensive literature
about them.’ Though they sometimes consist of almost incredible
numbers of individuals, yet it appears that in the Carpathian
mountains the assemblages are usually much smaller, being from
four to twenty inches long. A species of Sciara is the “ Yellow-
fever fly” of the Southern United States. It appears that it
has several times appeared in unusual numbers and in unwonted
localities at the same time as the dreaded disease, with which it
is popularly supposed to have some connection.

Fam. 3. Blepharoceridae..— Wings with no discal cell, but
with a secondary set of crease-like lines. The flies composing
this small family are very little known, and appear to be
obscure Insects with somewhat the appearance of Empidae,
though with strongly iridescent wings; they execute aerial
dances, after the manner of midges, and are found in Europe
(the Pyrenees, Alps and Harz mountains) as well as in North
and South America. Their larvae are amongst the most re-
markable of Insect forms; indeed, no entomologist recognises
them as belonging to a Hexapod Insect when he makes a first

1 See Guérin-Méneville, Ann. Soc. ent. France (2) iv. 1846; Bull. p. 8; and
Nowicki, Verh. Ges. Wien, xvii. 1867, SB. p. 23.

? For details as to the family cf. Osten ‘Sacken, Berlin. ent. Zeitschr. x1.
1895, p. 148; and for the larvae F. Miiller, Arch. Mus. Rio-Jan. iy. 1881,
p. 47. The name ‘‘Liponeuridae” was formerly applied by some authorities
to this family, but it is now generally recognised that Blepharoceridae is more
legitimate.

vir BLEPHAROCERIDAE 465

+ ——

acquaintance with them. The larva of Curupira (Fig. 222)
lives in rapid streams in Brazil, fixed by its suckers to stones or
rocks. It consists only of six or seven divisions, with project-
ing side-lobes; the usual segmentation not being visible. There
are small tracheal gills near the suckers, and peculiar scale-like
organs are placed about the edges of the lobes. Miller considers
that the first lobe is “cephalothorax,” corresponding to head,
thorax and first abdominal segment of other larvae, the next four
lobes he considers to correspond each to an
abdominal segment, and the terminal mass
to four segments. He also says that
certain minute points existing on the sur-
face, connected with the tracheal system by
minute strings, represent nine pairs of
spiracles. These larvae and their pupae
can apparently live only a short time after
being taken out of the highly aérated
water in which they exist, but Miiller
succeeded in rearing several flies from a
number of larvae and pupae that he
collected, and, believing them to be all one
species, he announced that the females
exhibited a highly developed dimorphism,
some of them being blood-suckers, others
honey-suckers. It is however, more prob-
able that these specimens belonged to two

Fic. 222.—Under surface of

or three distinct species or even genera. the larva of Curupira
é : Be ya! : 1 (Paltostoma) torrentium,
This point remains to be cleared up. The shawitietha siiceers alone
larva we have figured is called by Miiller the middle of the body,
me : ae Hawo : much magnified. Brazil.

Paltostoma torrentium. It is certain, how- (After Fritz Miiller.)

ever, that the Brazilian Insect does not
belong to the genus Paltostoma, and it will no doubt bear the
name used by Osten Sacken, viz. Curupira.

The metamorphoses of the European Liponeura brevirostris
have been partially examined by Dewitz, who found the Insects
in the valley of the Ocker in September.’ He does not consider
the “cephalothorax ” to include an abdominal segment; and he
found that two little, horn-lke projections from the thorax of the

1 Berlin. ent. Zeit. xxv. 1881, p. 61; and cf. Brauer, Wien: ent. Zeit. i. 1882,
jib dle
VOL. VI 2H

4060 DIPTERA CHAP.

ee

pupa are really each four-leaved. The pupa is formed within
the larval skin, but the latter is subsequently cast so that the
pupa is exposed; its dorsal region is horny, but the under sur-
face, by which it clings firmly to the stones of the rapid brook, is
white and scarcely chitinised, and Dewitz considers that the
chitinous exudation from this part is used as a means of fastening
the pupa to the stones. Blepharoceridae possess, in common
with Culex, Psychoda and Ptychoptera, the peculiar number of
five Malpighian tubes, and it has been proposed by Miiller to
form these Insects into a group called Pentanephria.

Fam. 4. Culicidae (Mosquitoes, Gnats)—Antennae with whorls
of hair or plumes, which may be very dense and long in the male,
though scanty in the female; head with a long, projecting pro-
boscis. Although there are few Insects more often referred to
in general literature than Mosquitoes, yet the ideas in vogue
about them are of the vaguest character. The following are the
chief points to be borne in mind as to the prevalence of Mos-
quitoes :—The gently humming Gnat that settles on us in our
apartments, and then bites us, is a Mosquito; there are a large
number of species of Mosquitoes ; in some countries many in one
locality ; in Britain we have ten or a dozen ; notwithstanding the
multiplicity of species, certain Mosquitoes are very widely diffused ;
the larvae are all aquatic, and specially frequent stagnant or
quiet pools; they are probably diffused by means of the water
in ships, it being known that Mosquitoes were introduced for
the first time to the Hawaiian Islands by a sailing vessel about
the year 1828. Hence it is impossible to say what species the
Mosquitoes of a given locality may be without a critical examina-
tion. No satisfactory work on the Mosquitoes of the world
exists. Urich states that he is acquainted with at least ten
species in Trinidad. The species common in our apartments in
Central and Southern England is Culea pipiens, Linn., and this
species is very widely distributed, being indeed one of the
troublesome Mosquitoes of East India. The term Mosquito is a
Spanish or Portuguese diminutive of Mosca. It is applied to a
variety of small flies of other families than Culicidae, but should
be restricted to these latter. The irritation occasioned by the
bites of Mosquitoes varies according to several circumstances,
viz. the condition of the biter, the condition or constitution of
the bitten, and also the species of Mosquito. Réaumur and

vII MOSQUITOES 467

others belheved that some irritating fluid is injected by the Mos-
quito when it bites. But why should it want to irritate as well
as to bite? Macloskie, considering that the Mosquito is really
a feeder on plant-substances, suggests that the fluid injected may
be for the purpose of preventing coagulation of the plant-juices
during the process of suction. It is a rule that only the female
Mosquito bites, the male being an inoffensive creature, and pro-
vided with less effectual mouth-organs; it has, however, been
stated by various authors that male Mosquitoes do occasionally
bite. It is difficult to understand the blood-sucking propensities
of these Insects; we have already stated that it is only the
females that suck blood. There is reason to suppose that it is an
acquired habit; and it would appear that the food so obtained
is not essential to their existence. It has indeed been asserted
that the act is frequently attended with fatal consequences to
the individual that does it. The proper method of mitigating
their nuisance is to examine the stagnant waters in localities
where they occur, and deal with them so as to destroy the larvae.
These little creatures are remarkable from the heads and thorax
being larger and more distinct than in other Dipterous larvae.
Their metamorphoses have been frequently described, and recently
the numerous interesting points connected with their life-histories
have been admirably portrayed by Professor Miall,’ in an accessible
form, so that it is unnecessary for us to deal with them.
Corethra is placed in Culicidae, but the larva differs totally from
that of Culex ; it is predaceous in habits, is very transparent, has
only an imperfect tracheal system, without spiracles, and has two
pairs of air-sacs (perhaps we should rather say pigmented struc-
tures possibly for aerostatic purposes, but not suppliers of oxygen).
The kungu cake mentioned by Livingstone as used on Lake
Nyassa is made from an Insect which occurs in profusion there,
and is compressed into biscwt form. It is believed to be a
Corethra. One of the peculiarities of this family is the pre-
valence of scales on various parts of the body, and even on the
wings: the scales are essentially similar to those of Lepidoptera.
Though Mosquitoes are generally obscure plain Insects, there
are some—in the South American genus Megarrhina—that are
elegant, beautifully adorned creatures. Swarms of various species
of Culicidae, consisting sometimes of almost incalculable numbers

1 Natural History of Aquatic Insects, London, 1895, chap. ii.

468 DIPTERA CHAP.

of individuals, occur in various parts of the world; one in New
Zealand is recorded as having been three-quarters of a mile long,
twenty feet high, and eighteen inches thick. There is good
reason for supposing that Mosquitoes may act as’ disseminators
of disease, but there is no certain evidence on the subject. The
minute Milaria that occurs in great numbers in some patients,
is found in the human body only in the embryonic and adult
conditions. Manson considers that the intermediate stages are
passed in the bodies of certain Mosquitoes.’

Fam. 5. Chironomidae ((nats, Midges)—Small or minute
flies of slender form, with narrow wings, without projecting
rostrum, usually with densely feathered antennae in the male, and
long slender legs. The flies of this family bear a great general
resemblance to the Culcidae. They are much more numerous
in species, and it is not improbable that we have in this country
200 species of the genus Chironomus alone. They occur in
enormous numbers, and frequently form dancing swarms in the
neighbourhood of the waters they live in. The species are
frequently extremely similar to one another, though distinguished
by good characters; they are numerous about Cambridge. Many
of them have the habit of using the front legs as feelers rather
than as means of support or locomotion. This is the opposite of
what occurs in Culicidae, where many of the species have a habit
of holding up the hind legs as if they were feelers. The eggs of
Chironomus are deposited as strings surrounded by mucus, and are
many of them so transparent that the development of the embryo
can be directly observed with the aid of the microscope. They are
said to possess a pair of air-sacs. The larvae, when born, are
aquatic in habits, and are destitute of tracheal system. They
subsequently differ greatly from the larvae of Culex, inasmuch as
the tracheal system that develops is quite closed, and in some
cases remains rudimentary. There is, however, much diversity
in the larvae and also in the pupae. The little Blood-worms,
very common in many stagnant and dirty waters, and used by
anglers as bait, are larvae of Chironomus. They are said to be
ai Eprides of Aristotle. The red colour of these larvae is due to
haemoglobin, a substance which has the power of attracting and
storing oxygen, and giving it off to the tissues as they require
it. Such larvae are able to live in burrows they construct

\ Tr. Linn. Soc. Lond. (2) ii. 1884, p. 367.

VII GNATS—MIDGES 469

amongst the mud. Some of them, provided plentifully with
haemoglobin, are in consequence able to live at great depths, it
is said even at 1000 feet in Lake Superior, and come to the
surface only occasionally. A few are able even to tolerate salt
water, and have been fished up from considerable depths in the
sea. It is a remarkable fact that these physiological capacities
differ greatly within the limits of the one genus, Chironomus,
for some of these species are destitute of haemoglobin, and have
to live near the surface of the water; these have a superior
development of the tracheal system. The pupae of Chironomus
have the legs coiled, and the thorax, instead of being provided
with the pair of tubes or trumpets for breathing that 1s so
common in this division of Diptera, have a pair of large tufts
of hair-like filaments." A very curious form of parthenogenesis
has been described by Grimm? as existing in an undetermined
species of Chironomus, inasmuch as the pupa deposits eggs.
Although this form of parthenogenesis is of much interest, it is
not in any way to be compared with the case, already referred to,
of Miastor (p. 461). The“ pupa” is at the time of oviposition prac-
tically the imago still covered by the pupal integument ; indeed
Grimm informs us that in some cases, after depositing a small
number of ova, the pupa became an imago. This partheno-
genesis only occurs in the spring-generation ; in the autumn the
development goes on in the natural manner. The case is
scarcely entitled to be considered as one of paedogenesis.

Gnats of this family, and believed to be a variety of Chiro-
nomus plumosus, are subject to a curious condition, inasmuch as
individuals sometimes become luminous or “ phosphorescent” ; this
has been noticed more specially in Eastern Europe and Western
Asia. The whole of the body and legs may exhibit the luminous
condition, but not the wings. It has been suggested by Schmidt
that this condition is a disease due to bacteria in the body of
the gnat.’

Ceratopogon is a very extensive genus, and is to some
extent anomalous as a member of Chironomidae. The larvae
exhibit considerable variety of form. Some of them are aquatic

1 For an extremely interesting account of Chironomus refer to Miall’s book,
already cited, and, for the larva, to the valuable work of Meinert on Eucephalous
larvae of Diptera, Danske Selsk. Skr. (6) iii. 1886, p. 436.

2 Ann. Nat. Hist. (4) viii. 1871; p. 31. 3 Ibid. (6) xv. 1895, .p. 1338.

470 DIPTERA _ CHAP.

in habits, but the great majority are terrestrial, frequenting
trees, etc. The former larvae are very slender, and move after
the manner of leeches; they give rise to imagos with naked
wings, while the terrestrial larvae produce flies with hairy
wings. There are also important distinctions in the pupae of
the two kinds; the correlation between the habits, and the dis-
tinctions above referred to, is, however, far from being absolutely
constant.’ Certain species of midges are in this country amongst
the most annoying of Insects; being of very minute size, scarcely
visible, they settle on the exposed parts of the body in great
numbers, and by sucking blood create an intolerable irritation.
Ceratopogon varius is one of the most persistent of these annoyers
in Scotland, where this form of pest is much worse than it is in
England ; in Cambridgeshire, according to Mr. G. H. Verrall,
the two troublesome midges are the females of C. pulicaris and
C. bipwnctatus.

Fam. 6. Orphnephilidae.—Simall, brown or yellowish flies,
bare of pubescence, with very large eyes contiguous in both sexes,
and with antennae composed of two joints and a terminal bristle ;
both the second joint and the bristle are, however, really complex.
One of the smallest and least known of the families of Diptera,
and said to be one of the most difficult to classify. The nervures
of the wings are very distinct. Nothing is known of the habits
and metamorphoses; there is only one genus—Orphnephila ; it
is widely distributed ; we have one species in Britain.

Fam. 7. Psychodidae (JJoth-flies)— Extremely small, helpless
jlies, usually with thickish antennae, bearing much hair, with
wings broader than is usual in small flies, and also densely clothed
with hair, giving rise to a pattern more or less vague. These
flies are very fragile creatures, and are probably numerous in
species. In Britain forty or fifty species have been recognised.”
A South European form is a blood-sucker, and has received the
appropriate name of Phlebotomus. The life-history of Pericoma
canescens has receritly been studied by Professor Miall.’ The
larva is of aquatic habits, but is amphibious, being capable of
existing in the air; it has a pair of anterior spiracles, by means

1 For metamorphoses of aquatic species of Ceratopogon, see Miall and Meinert,
already quoted: for examples of the terrestrial species, and their illustrations,
refer to Mik, Wien. ent. Zeit. vii. 1888, p. 183.

2 Monograph, Eaton, Ent. Mag. xxix. and xxx. 1893, 1894: supplement op. cit.
1896, etc. 3 Tr. ent. Soc. London, 1895, p. 141.

eee ee

VII ESYCHODIDAE——_DIXIDAE——TIPULIDAE 47%

of which it breathes in the air, and a pair at the posterior
extremity of the body, surrounded by four ciliated processes,
with which it forms a sort of cup for holding air when it is
in the water. The favourite position is amongst the filaments
of green algae on which it feeds. A much more extraordinary
form of larva from South America, doubtless belonging to
this family, has recently been portrayed by Fritz Miiller,
under the name of Maruwina.' These larvae live in rapid waters
In company with those of the genus Curupira, and like the
latter are provided with a series of suctorial ventral discs.
Fritz Miiller’s larvae belong to several species, and probably to
more than one genus, and the respiratory apparatus at the
extremity of the body exhibits considerable diversity among
them.

Fam. 8. Dixidae.—The genus Diva must, it appears, form a
distinct family allying the Culicid series of families to the
Tipulidae. The species are small, gnat-like Insects, fond of
damp places in forests. We have four British species (D.
maculata, D. nebulosa, D. aestivalis, D. aprilina). The genus is
very widely distributed, occurring even in Australia. The
larvae are aquatic, and have been described by Réaumur,
Miall, and Meimert. The pupa has the legs coiled as in the
Culicidae.

Fam. 9. Tipulidae (Daddy-long-legs, or Crane-flies).—Slender
Insects with elongate legs, a system of wing-nervures, rather com-
plex, especially at the tip; an angulate, or open V-shaped, suture
on the dorsum of the thoraxin front of the wings: the female
with the body terminated by a pair of hard, pointed processes,
concealing some other processes, and forming an ovipositor. The
curious, silly Insects called daddy-long-legs are known all
over the world, the family being a very large one, and found
everywhere, some of its members extending their range even
to the most inclement climates. It includes a great variety
of forms that would not be recognised by the uninitiated,
but can be readily distinguished by the characters mentioned
above. It is impossible to assign any reason of utility for
the extreme elongation of the legs of these Insects; as
everyone knows, they break off with great ease, and the Insect
appears to get on perfectly well without them. It is frequently

1 Tr. ent. Soc. London, 1895, p. 479.

472 DIPTERA CHAP.

the case that they are much longer in the males than in
the females: Other parts of the body exhibit a_ peculiar
elongation; in some forms of the male the front of the head
may be prolonged into a rostrum. In a few species the
head is separated by a great. distance from the thorax, the
gap being filled by elongate, hard, cervical sclerites; imdeed
it is in these Insects that the phenomenon, so rare in Insect-
structure, of the elongation of these sclerites and their be-
coming a part of the actual external skeleton, reaches its
maximum. In several species of Eriocera the male has the
antennae of extraordinary length, four or five times as long as
the body, and, strange to say, this elongation is accompanied
by a reduction in the number of the segments of which the
organ is composed, the number being in the male about six,
in the female ten, in place of the usual fourteen or sixteen.
In Youxorrhina and EHlephantomyia the proboscis is as long as
the whole body. In other forms the wings become elongated to
an unusual extent by means of a basal stalk. It is probable
that the elongation of the rostrum may be useful to the Insects.
Gosse,’ indeed, describes Limnobia intermedia as having a rostrum
half as long as the body, and as hovering lke a Syrphid, but
this is a habit so foreign to Tipulidae, that we may be pardoned
for suspecting a mistake. The larvae exhibit a great variety of
form, some being terrestrial and others aquatic, but the ter-
restrial forms seem all to delight in damp situations, such
as shaded turf or rotten tree-stems. They are either amphi-
pheustic or metapneustic, that is, with a pair of spiracles placed
at the posterior extremity of the body; the aquatic species
frequently bear appendages or projections near these spiracles.
The pupae in general structure are very like those of Lepidoptera,
and have the legs extended straight along the body ; they possess
a pair of respiratory processes on the thorax in the form of
horns or tubes. |

There are more than 1000 species of these flies known,
and many genera. They form three sub-families, which are by
some considered distinct families, viz.: Ptychopterinae, Limno-
biinae or Tipulidae Brevipalpi, Tipulinae or Tipulidae Longi-
palpi.

The Ptychopterinae are a small group in which the angulate

1 A Naturalist’s Sojourn in Jamaica, London, 1853, p. 284.

VII TIPULIDAE

suture of the mesonotum is indistinct; the larvae are aquatic and

have the head free, the terminal two segments
of the body enormously prolonged (Fig. 225),
forming a long tail bearing, in the North
American Littacomorpha, two respiratory fila-
ments. Hart describes this tail as possessing
a stigmatal opening at the extremity ; no doubt
the structure is a compounded pair of spiracles.
The pupa (Fig. 223, B) has quite lost the respira-
tory tube at the posterior extremity of the
body, but has instead quite as long a one at
the anterior extremity, due to one tube of the
pair normal in Tipulidae being enormously
developed, while its fellow remains small. This
is a most curious departure from the bilateral
symmetry that is so constantly exhibited in
Insect-structure. Our British species of Ptych-
optera have the pupal respiratory tube as extra-
ordinary as it is in Littacomorpha, though the
larval tail is less peculiar.” This group should
perhaps be distinguished from the Tipulidae
as a separate family, but taxonomists are not
yet unanimous as to this. Brauer considers
that the head of the larva, and the condition
of five Malpighian tubules in the imago,
require the association of Ptychopterinae with
the preceding families (Chironomidae, etc.),
rather than with the Tipulidae.

The great majority of the Tipulidae are com-
prised in the sub-family Limnobiinae—the
Tipulidae Brevipalpi of Osten Sacken:? in
them the last joint of the palpi is shorter or
not much longer than the two preceding
together. They exhibit great variety, and many
of them are types of fragility. The common
winter gnats of the genus 7Zrichocera are a
fair sample of this sub-family. The species

>

setae
.

eee,

eavene.
Weewean

i

if

il

ii

cit

3ii\\

|

weer
ve

@

\

Fic. 223. — Bittaco-

mor pha

cla vipes.

North America x 3.

(After
Larva ;
7, the
right
tube.

Hart.) A,
B, pupa:
lett, 7, the
respiratory

of this genus mostly inhabit high latitudes, and delight in

1 Bull. Illinois Lab., iv. 1895, p. 193. 2 Miall’s Aquatic Insects, 1895, p. 174.

3 “Studies,” etc., Berlin. ent. Zeitschr. xxxi.

1887.

474 DIPTERA CHAP.

a low temperature; it has been said that they may be seen
on the wing’ in the depth of winter when the temperature is
below freezing, but it is pretty certain that the spots chosen by
the Insects are above that temperature, and Eaton states that the
usual temperature during their evolutions is about 40° or 45°
Fahr. They often appear in the damp conditions of a. thaw
when much snow is on the ground. 7. simonyi was found at an
elevation of 9000 feet in the Tyrol, crawling at a temperature
below the freezing-point, when the ground was deeply covered with
snow. 7. regelationis occurs commonly in mines even when they
are 500 feet or more deep. The most extraordinary of the
Limnobiunae is the genus Chionea, the species of which are totally
destitute of wings and require a low temperature. C. araneoides
inhabits parts of northern Europe, but descends as far south as
the mountains near Vienna; it is usually said to be only really
active in the depth of winter and on the surface of the snow.
More recently, however, a large number of specimens were found
by Professor Thomas in the month of October in his garden in
Thuringia; they were caught in little pit-falls constructed to
entrap snails. The larva of this Insect is one of the interesting
forms that display the transition from a condition with spiracles
at the sides of the body to one where there is only a pair at the
posterior extremity.

A very peculiar Fly, in which the wings are reduced to mere
slips, Halirytus amphibius, was discovered by Eaton in Kerguelen
Land, where it is habitually covered by the rising tide. Though
placed in Tipulidae, it is probably a Chironomid.

The group Cylindrotomina is considered by Osten Sacken? to
be to some extent a primitive one having relationship with the
Tipulinae; it was, he says, represented by numerous species in
North America during the Oligocene period. It is of great
interest on account of the larvae, which are in several respects
similar to caterpillars of Lepidoptera. The larva of Cylindrotoma
distincta lives upon the leaves of plants—Anemone, Viola,
Stellaria—almost like a caterpillar; it is green with a crest
along the back consisting of a row of fleshy processes. Though
this fly is found in Britain the larva has apparently not been
observed here. The life-history of Phalacrocera replicata has
been recently published by Miall and Shelford.? The larva eats

1 Tr. ent. Soc. London, 1897, p. 362. 2 Tr. ent. Soc. London, 1897, pp. 343-361.

VII TIPULIDAE——-BIBIONIDAE 475

submerged mosses in the South of England, and bears long forked
filaments, reminding one of those of caterpillars. This species
has been simultaneously discussed by Bengtsson, who apparently
regards these Tipulids with caterpillar-like larvae—he calls them
Erucaeformia'—as the most primitive form of existing Diptera.
The Tipulinae—Tipulidae Longipalpi, Osten Sacken*—have
the terminal joint of the palpi remarkably long, longer than the
three preceding joints together. The group includes the largest
forms, and the true daddy-long-legs, a Chinese species of which,
Tipula brobdignagia, measures four inches across the expanded
wings. The group contains some of the finest Diptera. Some
of the exotic forms allied to Ctenophora have the wings coloured
in the same manner as they are in certain Hymenoptera, and
bear a considerable resemblance to members of that Order.
- Fam. 10. Bibionidae.—/Vies of moderate or small size, some-
times of different colours in the two sexes, with short, thick, straight,
antennae; front tibiae usually with a long pointed process ,;
coxae not elongate. Eyes of male large, united, or contiguous in

Fic. 224.—Head of Bibio. «10. A, Of male, seen from the front ; C, from the side ;
a, upper, b, lower eye ; B, head of female.

front. The flies of the genus BAibio usually appear in England
in the spring, and are frequently very abundant; they are of
sluggish habits and poor performers on the wing. The differ-
ence in colour of the sexes is very remarkable, red or yellow
predominating in the female, intense black in the male; and

1 Acta Univ. Lund. xxxiii. (2) No. 7, 1897.
2 «« Studies,” etc., Berlin. ent. Zeitschr. xxx. 1886, p. 153.

476 DIPTERA CHAP.

it is a curious fact that the same sexual distinction of colour
reappears in various parts of the world—England, America, India,
and New Zealand; moreover, this occurs in genera that are by
no means closely allied, although allied species frequently have
concolorous sexes. The eyes of the males are well worth study,
there being a very large upper portion, and,
abruptly separated from this, a smaller, differ-
ently faceted lower portion, practically a
separate eye; though so largely developed the
upper eye 1s In some cases so hairy that it
must greatly interfere with the formation
of a continuous picture. Carriere con-
siders that the small lower eye of the male
corresponds to the whole eye of the female.
The larvae of Bibio (Fig. 225) are caterpillar-
like in form, have a horny head, well de-
veloped, biting mouth-organs, and spine-like
processes on the body-segments. They are
certified by good authorities’ to possess the ex-
tremely unusual number of ten pairs of spiracles;
a larva found at Cambridge, which we refer to
Bibio (Fig. 225) has nine pairs of moderate
spiracles, as well as a large terminal pair
separated from the others by a segment without
spiracles. The genus Dilophus is closely
allied to Bibio, the larvae of which (and those
of Bibionidae in general) are believed to feed
on vegetable substances; the parasitism of
Dilophus vulgaris on the larva of a moth, Zpino-—
Fic. 225.—Larva of tia (Chaetoptria) hypericana, as recorded by
see sp. Cam Meade,” must therefore be an exceptional case.
In the genus Scatopse there is a very im-

portant point to be cleared up as to the larval respiratory
system; it is said by Dufour and Perris* to be amphipneustic ;
there are, however, nine projections on each side of the body that
were considered by Bouché, and probably with good reason, to

1 Osten Sacken, Berlin. ent. Zeitschr. xxxvii. 1892, p. 450.

2 Entomologist, xiv. 1881, p. 287. This observation has never, we believe, been
confirmed.

3 Ann. Soc. ent. France (2) v. 1847, p. 46.

VII BIBIONIDAE—SIMULIIDAE AV,

be spiracles. The food of Scatopse in the larval state is
principally vegetable. The larva of Scatopse changes to a
pupa inside the larval
skin; the pupa is pro-
vided on the thorax with
two branched respiratory
processes that project
outside the larval skin.!
Lucas has given an in-
teresting account of the
occurrence of the larva
of Bibio marci in enorm-
ous numbers at Paris;
they lived together in
masses, there being ap-

parently some sort of Fic. 226.—Portion of integument of Bibio sp. Can-
bridge. yp, Intersegmental processes ; s, spiracle.

connection between the
individuals.” In the following year the fly was almost equally
abundant.

Owing to the great numbers in which the species of Bibionidae
sometimes appear, these Insects have been supposed to be very
injurious. Careful inquiry has, however, generally exculpated
them as doers of any serious injury, though Dilophus febrilis—a
so-called tever-fly—appears to be really injurious in this country
When it multiplies excessively, by eating the roots of the hop-
plant.

Fam. 11. Simuliidae (Sand-jties, Buffalo-gnats).—Small obese
flies with humped back, rather short legs and broad wings, with short,
straight antennae destitute of setue; proboscis not projecting.

- There is only one genus, Simulium, of this family, but it is very
widely spread, and will probably prove to be nearly cosmo-
poltan. Some of the species are notorious from their blood-suck-
ing habits, and in certain seasons multiply to an enormous extent,
alight in thousands on cattle, and induce a disease that produces
death in a few hours; it is thought as the result of an instilled
poison. S. columbaczense has occasioned great losses amongst
the herds near the Danube; in North America the Buffalo- and
Turkey-gnats attack a variety of mammals and birds. In Britain

* Perris, in Ann. Soc. ent. France (2) v. 1847, p. 37, pl. i.
2 Ann. Soc. ent. France (5) i. 1871, Bull. p. lxvii.

478 DIPTERA CHAP.

and other parts of the world they do not increase in numbers to
an extent sufficient to render them seriously injurious: their bite
is however very annoying and irritating to ourselves. In their
early stages they are aquatic and require well aérated waters:
the larvae hold themselves erect, fixed to a stone or some other
object by the posterior extremity, and have on the head some
beautiful fringes which are agitated in order to bring food within
reach; the pupae are still more remarkable, each one being
placed in a pouch or sort of watch-pocket, from which projects
the upper part of the body provided with a pair of filamentous
respiratory processes. For an account of the interesting circum-
stances connected with the metamorphoses of this species the
reader should refer to Professor Miall’s book; and for the life-
history of the American Buffalo-gnat to Riley."

Fam. 12. Rhyphidae.—This is another of the families that
have only two or three genera, and yet are very widely distributed.
These lttle flies are distinguished from other Nemocera Anomala
(cf. p. 456) by the presence of a discal cell; the empodia of the
feet are developed as if they were pulvilli, while the true pulvilli
remain rudimentary. The larvae are lke little worms, being
long and cylindric; they are amphipneustic, and have been
found in decaying wood, in cow-dung, in rotten fruits, and
even in dirty water. The “petite tipule,” the metamorphoses of
which were described and figured by Réaumur, is believed to
be the common Rhyphus fenestralis. R. fenestralis is often
found on windows, as its name implies.

Series 2. Orthorrhapha Brachycera

Fam. 13. Stratiomyidae.—<Antennae with three segments
and a terminal complex of obscure joints, frequently bearing
an arista: tibiae not spined; wings rather small, the an-
terior nervures usually much more strongly marked than those
behind. The median cell small, placed near to the middle of the
wing. Scutellum frequently spined ; terminal appendages of the
tarsi small, but pulvilli and a pulvilliform empodium are

1 Rep. Dep. Agric. Ent. Washington, 1886, p. 492.

2 Cf. Réaumur, Mem. v. 1740, p. 21; and Perris, Ann. Soc. ent. France (4) x.
1870, p. 190.

VII STRATIOMYIDAE——-LEPTIDAE 479

present. This is a large family, whose members are very
diversified, consequently definition of the whole is difficult. The
species of the typical sub-family Stratiomyinae generally have the
margins of the body prettily marked with green or yellow, and
the scutellum spined. In the remarkable American genus,
Hermetia, the abdomen is much constricted at the base, and the
scutellum is not spined; in the division Sarginae the body is
frequently of brilliant metallic colours. The species all have
an only imperfect proboscis, and are not blood-suckers. The
larvae are also of diverse habits; many of those of the Stratio-
myinae are aquatic, and are noted for their capacity of living in
salt, alkaline, or even very hot water. Mr. J. C. Hamon found
some of these larvae in a hot spring in Wyoming, where he could
not keep his hand immersed, and he estimated the temperature
at only 20° or 30° Fahr. below the boiling-point. The larva
of Stratiomys is of remarkably elongate, strap-like, form, much
narrowed behind, with very small head; the terminal segment
is very long and ends in a rosette of hairs which the creature
allows to float at the surface. After the larval skin is shed the
pupa, though free, is contained therein; the skin alters but
little in form, and has no organic connection with the pupa,
which merely uses the skin as a shield or float. These larvae
have been very frequently described; they can live out of the
water. Brauer describes the larvae of the family as “ perip-
neustic, some perhaps amphipneustic.” Miall says there are, in
Stratiomys, nine pairs of spiracles on the sides of the body which
are not open, though branches from the longitudinal air-tubes
pass to them. There are probably upwards of 1000 species of
Stratiomyiidae known, and in Britain we have 40 or 50 kinds.
The American genus Chiromyza, Wied., was formerly treated by
Osten Sacken as a separate family, Chiromyzidae, but Williston
places it in Stratiomyidae.

Fam. 14. Leptidae, including Xylophagidae and Coenomyi-
idae.—The Leptidae proper are flies of feeble build; antennae
with three joints and a terminal bristle ; in the Xylophagidae the
antennae are longer, and the third joint is complex. The wings
have five posterior cells, the middle tibiae are spined. Pulvilli and a
pulvilliform empodium present. The three families are considered
distinct by most authors, but there has always been much difficulty
about the Xylophagidae and Coenomyiidae, we therefore treat them

480 DIPTERA CHAP.

as sub-families. The Xylophaginae are a small group of slender
Insects, perhaps most like the short-bodied kinds of Asilidae ;
the third joint of the antenna is vaguely segmented, and there
is no terminal bristle. &hachicerus is a most anomalous
little fly with rather long stiff antennae of an almost nemo-
cerous character, the segments of which give off a short
thick prolongation on each side, reminding one of a two-edged
saw. The three or four British species of Xylophaginae
are forest Insects, the larvae of which live under bark, and”
are provided with a spear-hke head with which they pierce
other Insects. The Coenomyiinae consist of the one genus
Coenomyia, with two or three European and North American
species. They are remarkably thick-bodied, heavy flies, reminding
one somewhat of an imperfect Stratiomyid destitute of orna-
mentation. The metamorphosis of C. ferruginea has been
described by Beling.’ The larva is not aquatic, but lives in
burrows or excavations in the earth where there are, or have
recently been, rotten logs; it is probably predaceous. It is
cylindric, with an extremely small head and eleven other segments,
the stigma on the first thoracic segment distinct; the terminal
segment is rather broad, and the structures surrounding the
stigma are complex. The pupa
has stigmata on each of ab-
dominal segments 2 to 8. Not-
withstanding that the fly is so
different to LXylophagus, the
larvae indicate the two forms
as perhaps really allied. One
of the Leptinae, Atheriz bis,
has a singular mode of ovi-
position (Fig. 227), the females
of the species deposit their eggs
in common, and, dying as they
do so, add their bodies to the

common mass, which becomes Fie. 227.—Atherix ibis. A, The fly, nat.

an agolomeration, it may be size ; B, mass of dead flies overhanging
v DE ied Ne water, much reduced.

of thousands of individuals, and

of considerable size. The mass is attached to a branch of a

bush or to a plant overhanging water, into which it ulti-

1 Verh. Ges. Wien, xxx. 1880, p. 343.

Vi LEPTIDAE—TABANIDAE 481

mately falls. These curious accumulations are occasionally
found in England as well as on the Continent, but no reason
for so peculiar a habit is at present forthcoming. Still more
remarkable are the habits of some European Leptids of the
genera Vermileo (Psammorycter of some authors) and Lamp-
romyia, slender rather small flies of Asilid-like appearance, the
larvae of which form pit-falls after the manner of the Ant-lion.
According to Beling' the larva of Leptis is very active, and 1s
distinguished by having the stigmatic orifice surrounded by four
quite equal, quadrangularly placed prominences ; and at the other

Fic. 228.—Larva of
Vermileo degeeri

(Psammorycter ver-
mileo). A, lateral,
B, dorsal view:
p, an abdominal
pseudopod; st,
stigma. Europe.
(After Réaumur
and Brauer. )

extremity of the body a blackish, naked, triangular plate; on the
under side of each of seven of the abdominal segments there is a
band of spines. The larva of Atherix has seven pairs of abdominal
feet. Altogether there are some two or three hundred known
species of Leptidae; our British species scarcely reach a score.
They are destitute of biting-powers and are harmless timid
creatures. Leptis scolopacea, the most conspicuous of our native
species, a soft-bodied fly of rather large size, the wings much
marked with dark colour, and the thick, pointed body yellowish,
marked with a row of large black spots down the middle, is a
common Insect in meadows.

Fam. 15. Tabanidae (Breeze - flies, Cleggs, or Horse - flies,
also frequently called Gad-jlies)—Proboscis fleshy, distinct, en-
wrapping pointed horny processes, palpi distinct, terminal joint
inflated, pendent in front of proboscis. Antennae projecting, four-
jointed, second joint very short, third variable in form, fourth
forming an indistinctly segmented continuation of the third, but
not ending in a bristle. A perfect squama in front of the halter.
Eyes large, very large in the males, but laterally extending, rather
than globose. This large and important family of flies, of which

1 Arch. Naturges. xii. i. 1875, p. 48.
VOU vs 21

482 DIPTERA CHAP.

Williston states that 1400 or 1500 species are named, is
well known’ to travellers on account of the blood - sucking
habits of its members; they have great powers
of flight, and alight on man and animals, and
draw blood by making an incision with the
proboscis; only the females do this, the
males wanting a pair of the lancets that
enable the other sex to inflict their for-
midable wounds. They are comparatively large
Insects, some of our English species of Zabanus
attaining an inch in length. The smaller, grey
Huematopota, is known to every one who has
walked in woods or meadows in the summer, as
it alghts quietly on the hands or neck and
bites one without his having previously been
made aware of its presence. The larger Tabani
hum so much that one always knows when an
individual is near. The species of Chrysops, in
habits similar to Hwematopota, are remarkable
for their beautifully coloured golden-green eyes.
In Brazil the Motuca fly, Hadrus lepidotus,
Perty, makes so large and deep a cut that con-
siderable bleeding may follow, and as it some-
times settles im numbers on the body, it is
deservedly feared. The most remarkable forms

Pangonia —% : . 2 . :
longirostris, x Of Tabanidae are the species of the widely dis-

Fic, 229.

1. Nepal. (After

tributed genus Pangonia (Fig. 229). The pro-
Hardwicke. ) 5 J (Fig. 229) e |

boscis in the females of some of the species is
three or four times the length of the body, and as it is. stiff and
needle-like the creature can use it while hovering on the wing, and
will pierce the human body even through clothing of considerable
thickness. The males suck the juices of flowers. The Seroot
fly, that renders some of the districts of Nubia uninhabitable
for about three months of the year, appears, from the figure and
description given by Sir Samuel Baker, to be a Pangonia.
Tabanidae are a favourite food of the fossorial wasps of the
family Bembecidae. These wasps are apparently aware of the
blood-sucking habits of their favourites, and attend on travellers
and pick up the flies as they are about to settle down to their
phlebotomic operations. The larvae of the Tabanidae are some

VII TABANIDAE——ACANTHOMERIDAE 483

g
wood; they are of predaceous habits, attacking and sucking
Insect-larvae, or worms. Their form is cylindric, attenuate
at the two extremities; the slender small head is retractile, and
armed with a pair of conspicuous, curved black hooks. The
body is surrounded by several promi-
nent rings. The breathing apparatus is
apparently but little developed, and con-
sists of a small tube at the extremity of
the body, capable of being exserted or
withdrawn ; in this two closely approxi-
mated stigmata are placed. In a larva,
probably of this family, found by the
writer in the shingle of a shallow stream
in the New Forest, the annuli are re-
placed by seven circles of prominent
pseudopods, on the abdominal segments
about eight in each circle, and each of
these feet is surmounted by a crown of
small hooks, so that there are fifty or
sixty feet distributed equally over the
middle part of the body without refer-
ence to upper or lower surface. The

of them aquatic, but others live in the earth or in decaying

Fic. 230.—Larva of a Taba-

figures of the larva of 7. cordiger, by nid. [? Atylotus fulvus.
6 che A (Ge ae 3; B
ee Mater . . Fs) foe a Cen eee 5 tne larva, xo): :
Brauer, and of Haematopota pluvialis, by so: esate OM cull at bal,
Perris, are something like this, but have D, one of the pseudopods.

New Forest.
no setae on the pseudopods. The meta- oA chee

morphoses of several Tabanidae are described and figured by Hart ;?
the pupa is remarkably like a Lepidopterous pupa. We have five
genera and about a score of species of Tabanidae in Britain.

Fam. 16. Acanthomeridae.—A very small family of two
genera (Acanthomera and Rhaphiorhynchus) confined: to America,
and including the largest Diptera, some being two inches long.
The antenna is terminated by a compound of seven segments and
a style; the proboscis is short, and the squama rudimentary.
The general form reminds one of Tabanidae or Oestridae. <A
dried larva exists in the Vienna collection ; it is amphipneustic,
and very remarkable on account of the great size of the anterior
stigma.

1 Bull. Illinois Lab. iv. 1895.

484 DIPTERA CHAP.

the appearance of short Asilidae. They have, however, only a
feeble fleshy proboscis, and minute claws, with pulvilli but no
empodium ; the antennae project, are short,
three - jointed, pointed—The flies of this
family are believed to be predaceous like
the Robber-flies, but they appear to be very
feebly organised for such a life. We have
about ten species in Britain, and there
are only some 200 known from all the
world. But little is known as to the meta-
morphoses. Meigen found larvae of 7.
Tig O31 eT aeun (eat: nobilitata im rotten stumps, but other larvae
cephala) confinis. A, have been recorded as devouring dead pupae
(Alter aa Europe. or larvae of Lepidoptera. The larvae are said
to be elongate, very slender, worm-like, and
to have nineteen body-segments, the posterior pair of spiracles
being placed on what looks like the seventeenth segment, but is
really the eighth of the abdomen. The pupa is not enclosed
in the larval skin; that of Psilocephala is armed with setae and
spinous processes, and was found in rotten wood by Frauenfeld.
Fam. 18. Scenopinidae.— Rather small flies, without bristles.
Antennae three-jointed, the third joint rather long, without ap-

pendage. Proboscis not projecting. Empodium absent. These
unattractive thes form one of the smallest families, and are
chiefly found on windows. SS. fenestralis looks like a tiny
Stratiomyid, with a peculiar, dull, metallic surface. The larva
of this species has been recorded as feeding on a variety of
strange substances, but Osten Sacken is of opinion! that it is
really predaceous, and frequents these substances in order to find
the larvae that are developing in them. If so, Scenopinus is useful
in a small way by destroying “moth,” etc. The larva is a little
slender, cylindrical, hard, pale worm of nineteen segments, with a
small brown head placed like a hook at one extremity of the body
and with two short, divergent processes at the other extremity,
almost exactly like the larva of 7hereva. Full references to the
literature about this Insect are given by Osten Sacken.

Fam. 19. Nemestrinidae.—These Insects appear to be allied
to the Bombyhidae. They are of medium size, often pilose, and

1 Ent. Mag. xxiii. 1886, p. 51.

VII NEMESTRINIDAE—BOMBYLIIDAE 4

ee)
ul

sometimes with excessively long proboscis; antennae short, with a
simple third joint, and a jointed, slender, terminal appendage ; the
tibiae have no spurs, the empodium is pulvillus-like. The wing-
nervuration is perhaps the most complex found in Diptera, there
being numerous cells at the tip, almost after the fashion of
Neuroptera. With this family we commence the aerial forms
composing the Tromoptera of Osten Sacken. Nemestrinidae is a
small family of about 100 species, but widely distributed.
Megistorhynchus longirostris 1s about two-thirds of an inch long,
but has a proboscis at least four times as long as itself. In
South Africa it may be seen endeavouring to extract, with this
proboscis, the honey from the flower of a Gladiolus that has a
perianth just as long as its own rostrum ; asit attempts to do this
when it is hovering on the wing, and as the proboscis is, unlike
that of the Bombyli, fixed, the Insect can only succeed by con-
trolling its movements with perfect accuracy ; hence it has great
difficulty in attaining its purpose, especially when there 1s much
wind, when it frequently strikes the earth instead of the flower.
M. Westermann thinks ' the life of the Insect and the appearance
and duration of the flower of the Gladiolus are very closely con-
nected. The life-history of Hirmoneura obscura has recently been
studied in Austria by Handlirsch and Brauer.” The larva is par-
asitic on the larva of a Lamellicorn beetle (2hizotroqus solstitialis) ;
it is metapneustic, and the head is highly modified for predaceous
purposes. The young larva apparently differs to a considerable
extent from the matured form. The most curious fact is that the
parent fly does not oviposit near the Lamellicorn-larva, but places
her eggs in the burrows of some wood-boring Insect in logs; the
larvae when hatched come to the surface of the log, hold them-
selves up on their hinder extremity and are carried away by the
wind ; in what manner they come into contact with the Lamellicorn
larva, which feeds in turf,is unknown. The pupa is remarkable
on account of the prominent, almost stalked stigmata, and of two
pointed divergent processes at the extremity of the body. This
life-history is of much interest, as it foreshadows to some extent
the complex parasitic life-histories of Bombyliidae. The Nemes-
trinidae are not represented in the British fauna.

Fam, 20. Bombyliidae— Body frequently fringed with down,

1 Ann. Soe. ent. France, ii. 1838, p. 492:
2 Wien. ent. Zeit. ii. 1883, pp. 11 and 24, pl. i.

486 DIPTERA CHAP.

or covered in large part with hair, Legs slender, claws small,
without distinct empodium, usually with only minute pulvilli.
Proboscis very long or moderate, antennae three-jointed, terminal
joint not distinctly divided, sometimes large, sometimes hair-like.
This is a very large family, including 1500 species, and is of
great importance to both naturalist and economist. Two well-
marked types, formerly treated as distinct families, are included in
it—(1) the Bombylides with very long exserted rostrum, and
humped thorax; and (2) Anthracides, with a short beak, and of
more slender and graceful form. None of these flies are blood-
suckers, they frequent flowers only, and use their long rostrums in
a harmless manner. The members of both of these groups usually
have the wings ornamented with a pattern, which in Anthrax
is frequently very remarkable; in both, the clothing of the body
is frequently variegated. Their powers of flight are very great,
and the hovering Bombylius of early spring is endowed with an
unsurpassed capacity for movement, remaining perfectly still on
the wing, and darting off with lhghtning rapidity; Anthrax is
also most rapid on the wing. In Britain we have but few
species of Bombylidae, but in warm and dry climates they are
very numerous. The life-history of these Insects was till recently
unknown, but that of Argyro-
moeba (Anthrax) trifasciata has
been described by the French
naturalist, Fabre, who ascertained
that the species is parasitic on the
Mason-bee, Chalicodoma muraria,
that forms nests of solid mas-
onry. He endeavoured to dis-
cover the egg, but failed; the
parent-fly oviposits, it appears,
by merely dropping a minute
egg while flying over the surface
of the mass of masonry by which
ti os, ea A ee 8 a | the grubs of the Ch alicodome are
Young larva; B, adult larva; ©, protected. From this egg there
Wea Sa ibnay aie is hatched a minute delicate ver-
miform larva (Fig. 232, A). In

order to obtain its food, it is necessary for this feeble creature to
penetrate the masonry; apparently a hopeless task, the animal

ee BOMBYLIIDAE 487

being scarcely a twentieth of an inch long and very slender ; it
is, however, provided with a deflexed horny head, armed in front
with some stiff bristles, while on the under surface of the body
there are four pairs of elongate setae serving as organs of loco-
motion; thus endowed, the frail creature hunts about the sur-
face of the masonry, seeking to find an entrance; frequently it
is a long time before it is successful; but though it has never
taken any food it 1s possessed of great powers of endurance.
Usually, after being disclosed from the egg, it remains about
fifteen days without stirring; and even after it commences its
attempts to enter the nest it is still capable of a long life without
taking any food. Possibly its organisation will not permit it to
feed (supposing any food were obtainable by it) without its
growing somewhat thereafter, and if so, its chance of obtaining
entrance through the masonry would be diminished. Abstention,
it would appear, is the best policy, whether inevitable or not ;
so the starving little larva continues its endeavours to find a
chink of entrance to the food contained in the interior of the
masonry. It has plenty of time for this, because it is better for
it not to get into the cell of the bee until the grub is quite full
grown, and is about to assume the pupal form, when it is quite
incapable of self-defence. Finally, after greater or less delay,
the persevering little larva succeeds in finding some tiny gap in
the masonry through which it can force itself. M. Fabre says
that the root of a plant is not more persistent in descending
into the soil that is to support it than is this little Anthrax
in insinuating itself through some crack that may admit it to its
food. Having once effected an entrance the organisation that
has enabled it to do so is useless; this primary form of the larva
has, in fact, as its sole object to enable the creature to penetrate to
its food. Having penetrated, it undergoes a complete change of
form, and appears as a creature specially fitted for feeding on the
quiescent larva of the bee without destroying it. To accomplish
this requires an extreme delicacy of organisation and instinct ;
to bite the prey would be to kill it, and if this were done, the
Anthrax would, Fabre supposes, ensure its own death, for it
cannot feed on the dead and putrefying grub; accordingly, the
part of its body that does duty as a mouth is merely a delicate
sucker which it apples to the skin of the Chalicodoma-grub ;

and thus without inflicting any perceptible wound it sucks day

488 DIPTERA CHAP.

after day, changing its position frequently, until it has completely
emptied the pupa of its contents, nothing being left but the skin.
Although this is accomplished without any wound being inflicted,
so effectual is the process that all the Chalicodoma is gradually
absorbed. The time requisite for completely emptying the victim
is from twelve to fifteen days; at the end of this time the
Anthraz-larva is full grown, and the question arises, how is it to
escape from the cell of solid masonry in which it is imprisoned ?
It entered this cell as a tiny, slender worm through a minute
orifice or crack, but it has now much increased in size, and exit
for a creature of its organisation is not possible. For some
months it remains a quiescent larva in the cell of the Chali-
codoma, but in the spring of the succeeding year it undergoes
another metamorphosis, and appears as a pupa provided with a
formidable apparatus for breaking down the masonry by which
it is imprisoned. The head is large and covered in front with
six hard spines, to be used in striking and piercing the masonry,
while the other extremity of the body bears some curious horns,
the middle segments being armed with rigid hairs directed back-
wards, and thus facilitating movement in a forward direction and
preventing shpping backwards. The pupa is strongly curved,
and fixes itself by the aid of the posterior spines; then, unbend-
ing itself, it strikes with the armour of the other extremity
agaist the opposing wall, which is thus destroyed piecemeal
until a gallery of exit is formed; when this is completed the
pupa-skin bursts and the perfect fly emerges, leaving the
pupa-case still fixed in the gallery. Thus this species appears
in four consecutive forms—in addition to the egg—each of
which is highly specialised for the purposes of existence in that
stage.

The habits of our British Bombylius major have been partially
observed by Dr. Chapman,’ and exhibit a close analogy with
those of Anthrax trifasciata. The bee-larva that served as food
was in this case Andrena labialis, and the egg was deposited
by the fly, when hovering, by jerking it against the bank in
which the nest of the bee was placed.

It has recently been discovered that the larvae of various
species of Bombylhidae are of great service by devouring the
eggs of locusts. Riley found that the egeg-cases of Caloptenus

1 Ent. Mag. xiv. 1878, p. 196.

VII BOMBYLIIDAE—ACROCERIDAE 489

spretus are emptied of their contents by the larvae of Systoechus
oreas and Triodites mus. A similar observation has been made
in the Troad by Mr. Calvert, who found that the Bombyliid,
Callostoma fascipennis, destroys large quantities of the eggs of
Caloptenus italicus. Still more recently M. Kiinckel d’Herculais
has discovered that the destructive
locust Stauwronotus maroccanus is
kept in check in Algeria in a
similar manner, as many as 80
per cent of the eggs of the locust
being thus destroyed in certain
localities. He observes that the
larva of the fly, after being full
fed in the autumn, passes the
winter in a state of lethargy—
he calls it “ hypnody ”—in the
ege-case of the locust, and he Ae Ge

ee : : Fig. 233.—Systropus crudelis. South
further informs us. that in the Africa. A, Pupa; B, imago, ap-

case of Anthrax fenestralis, which — pendages of the left side removed.
(After Westwood.)

devours the eggs of the large
Ocnerodes, the lethargy may be prolonged for a period of three
years. After the pupa is formed it works a way out of the case
by means of its armature, and then again becomes for some days
immobile before the perfect fly appears. Lepidopterous larvae ’
are also attacked by Bombyliid flies. A species of Systropus has
been recorded as destroying the larva of Limacodes. Several of
the Bombylhids of the genus just mentioned are remarkable for
the great resemblance they display to various Hymenoptera, some
of them being very slender flies, like the thin bodied fossorial
Hymenoptera. The difference between the pupa and imago in
this case is very remarkable (Fig. 233).

Fam. 21. Acroceridae or Cyrtidae.—/Vies of the average size,
of peculiar form, the small head consisting almost entirely of the
eyes, and bent down under the humped thorax: wings small, halteres
entirecy concealed by the very large horizontal squamae ; antennae
very diverse. The peculiar shape of these flies is an exaggeration of
that we have already noticed in Bombyliuvs. The mouth in Acro-
ceridae is very variable ; there may be a very long, slender proboscis
(Aerocera), or the mouth-parts may be so atrophied that it is
doubtful whether even an orifice exists (Ogcodes). There are but

490 DIPTERA CHAP.

few species known, and all of them are rare;' in Britain we

have but two (Ogcodes gibbosus, Acrocera globulus). The genus
Pterodontia, found in North America and Australia, an inflated
bladder-like form with a minute head, is
amongst the most extraordinary of all the
forms of Diptera. The habits are very
peculiar, the larvae, so far as known, all
living as parasites within the bodies of
spiders or in their egg-bags. It appears,
however, that the flies do not oviposit in
appropriate places, but place their eggs on
Fic. 234.—Megalybus gra. stems of plants, and the young larvae have
Be ( oa dee to find their way to the spiders. Brauer has
described the larva of the European Asto-
mella lindeni, which lives in the body of a spider, Cteniza
arvana ; it is amphipneustic and maggot-like, the head being
extremely small. The larva leaves the body of the spider for
pupation ; the pupa is much arched, and the head is destitute of
the peculiar armature of the Bombyliidae, but has a serrate ridge
on the thorax. Emerton found the larvae of an Acrocera in the
webs of a common North American spider, Amaurobius sylvestris,
they having eaten, it was supposed, the makers of the cobwebs.
Fam. 22. Lenchopteridae.—Smual/, slender flies, with pointed
wings, short, porrect antennae, with a simple, circular third joint,
bearing a bristle ; empodium very small, pulvilli absent—Only one
genus of these little flies is known, but it is apparently widely
distributed, and its members are common Insects. They have
the appearance of Acalyptrate Muscidae, and the nervuration of
the wing is somewhat similar, the nervures being simple and
parallel, and the minute cross-nervures placed near the base.
The systematic position is somewhat doubtful, and the meta-
morphoses are but incompletely known, very little having been
added to what was discovered by Sir John Lubbock in 1862.°
The larva lives on the earth under vegetable matter; it is very
transparent, amphipneustic, with a peculiar head, and with fringes
on the margins. This larva changes to a semi-pupa or apterous
maggot-lke form, within the larval skin; the true pupa was

| For figures, etc., cf. Westwood, 7'r. ent. Soc. London, 1876, p. 507, pls. v. vi.

2 VerhGes. Wien, xix. 1869, p. 7d”; pl. xii.
3 Tr. ent. Soc. London (3) i. 1862, p. 338, pl. xi.

VII LONCHOPTERIDAE——MYDAIDAE—ASILIDAE 491

not noticed by Lubbock, but Frauenfeld' has since observed it,
though he only mentions that it possesses differentiated limbs
and segments. The metamorphoses appear to be very peculiar.
This fly requires a thorough study.

Fam. 23. Mydaidae.— Large flies of elongate form ; the hind
femora long and toothed beneath ; the antennae knobbed at the tip,
projecting, rather long, the basal joint definite, but the divisions
of the subsequent joints more or less indistinct. Empodium small.
Wings frequently heavily pigmented ; with a complex nervuration.
These fine flies are exotic; a few species occur in the Mediter-
ranean region, even in the South of Europe; the chief genus,
Mydas, is South American, but most of the other genera are
Australian or African. But little is known as to the life-
histories. The larvae are thought to live in wood, and to prey
on Coleopterous larvae.

Fam. 24, Asilidae (Robber -jflies)—Mouth forming a short,
projecting horny beak, the palpi usually only small; the feet
generally largely developed ; the claws large, frequently thick and
blunt, the pulvilli generally elongate, the empodium a bristle ;
halteres free; no squama. The Asilidae is one of the largest
families of flies, and probably includes about 5000 described
species: as will readily be believed, there is- much variety of
form; some are short and thick and extremely hairy, superficially
resembling hairy bees, but the majority are more or less elongate,
the abdomen being specially long, and having eight segments
conspicuously displayed. The antennae are variable, but are
three-jointed with a terminal appendage of diverse form and
structure. They belong to the super-family Energopoda of Osten
Sacken, but the association of Empidae and Dolichopidae with them
does not seem to be very natural. In their perfect state these
flies are most voracious, their prey being Insects, which they
seize alive and impale with the rostrum. They are amongst
the most formidable of foes and fear nothing, wasps or other
stinging Insects being attacked and mastered by the stronger
species without difficulty. They have been observed to capture
even dragon-flies and tiger-beetles. As is the case with so many
other Insects that prey on living Insects, the appetite in the
Asilidae seems to be insatiable; a single individual has been
observed to kill eight moths in twenty minutes. They have

1 Verh. Ges. Wien, xix. 1869, p. 941.

492 DIPTERA CHAP.

been said to suck blood from Vertebrates, but this appears to be
erroneous. ‘The metamorphoses of a few species have been ob-
served. Perris has called attention to the close alliance between
the larvae of Tabanidae and of Asilidae,’ and it seems at present
impossible to draw a line of distinction between the two. So
far as is known, the larvae of Asilidae are terrestrial and
predaceous, attacking more particularly the larvae of Coleoptera,
into which they sometimes bore; in Laphria there are numerous
pseudopods, somewhat of the kind shown in Fig. 230, but less
pertect and without hairs; the head and breathing organs appear
to be very different. According to Beling’s descriptions of the
larvae of Asi/us, the head in this case is more like that of the
figure, but there are no pseudopods. The flies of Asilidae and
Tabanidae are so very distinct that these resemblances between
their larvae are worthy of note.

Fam. 25. Apioceridae.—Moderate-sized flies marked with black
and white, with an appearance like that of some Muscidae and
Asilidae ; with clear wings, the veins not deeply coloured ; antennae
short, with a short, simple appendage ; no empodium. But little
is known as to the flies of this family, of which only two genera,
consisting of about a dozen species, are found in North America,
Chili, and Australia. Osten Sacken is inclined to treat them
as an aberrant division of Asilidae. Brauer looks on them as
primitive or synthetic forms of much interest, and has briefly
described a larva which he considers may be that of Apiocera,
but this is doubtful; it is a twenty-segmented form, and may be
that of a Thereva.?

Fam. 26. Empidae.—Simall or moderate-sized flies of obscure
colours, grey, rusty, or black, with small head, somewhat globular
in form, with three-jointed antennae, the terminal joint long and
pointed ; usually there is a long slender beak ; the legs are elongate,
frequently hairy; the tarsi bear long pulvilli and a small em-
podium. The Empidae are an extensive family of flies, with
predaceous habits, the rostrum being used by the female as
an instrument for impaling and sucking other flies. They are
occasionally very numerous in individuals, especially in wooded
districts. There is great variety; there are nearly 200 species
in Britain, The forms placed in the sub-family Hybotinae are
curious slender little Insects, with very convex thorax and

1 Ann. Soc. ent. France (4) x. 1870, p. 221. 2 SB. Ak. Wien, xci. 1885, p. 392.

VII EMPIDAE—DOLICHOPIDAE 493

large hind legs. In Hemerodromia the front legs are raptorial,
the femora being armed with spines on which the tibiae
close so as to form a sort of trap. Many Empidae execute
aérial dances, and some of the species of the genus Hilara are
notorious for carrying veils or nets in the form of silken webs
more or less densely woven. This subject is comparatively new,
the fact having been discovered by Baron Osten Sacken in
1877, and it is not at all clear what purpose these peculiar
constructions serve; it appears probable that they are carried
by means of the hind legs, and only by the males. Mik thinks
that in Hf. sartor the veil acts as a sort of parachute, and is of
use In carrying on the aérial performance, or enhancing its
effect ; while in the case of other species, H. maura and H. inter-
stincta, the object appears to be the capture or retention of prey,
after the manner of spiders. The source of the silk is not
known, and in fact all the details are insufficiently ascertained.
The larvae of Empidae are described as cylindrical maggots, with
very small head, and imperfect ventral feet; the stigmata are
amphipneustic, the thoracic pair being, however, excessively
small; beneath the posterior pair there is nearly always a tooth-
or spine-like prominence present.

Fam. 27. Dolichopidae.— Graceful flies of metallic colours,
of moderate or small size, and long legs ; usually with bristles on the
thorax and legs, the halteres exposed, squamae
being quite absent ; antennae of two short stout
joints (of which the second is really two, its
division being more or less distinet), with a
thread-like or hair-like appendage. Proboscis
short, fleshy. Claws, pulvilli, and empodium
small ; wings with a simple system of nervures,
those on the posterior part of the wing are but
few, there is no anterior basal eross-vein be-

tween the discal and second basal cells, which
ye Jae 2 eae Fic. 235.—A, Larv:
therefore form but one cell. This is also a sas ae Dans
e at ; : 5 . pupa of Mede-
very extensive family of flies, of which we — terus — ambiguus.
‘ : : Sree France. (After
av “ghahliv ah 9 q Jac Ipite
have probably about 200 species in Britain. Pain)
They are conspicuous on account of their
golden, or golden-green colours, only a few being yellow or black.

1 Ent. Mag. xiv. 1877, p. 226; for a discussion of the subject see Mik, Wien.
ent. Zeit. xili. 1894, p. 273.

494 DIPTERA CHAP.

The males are remarkable for the curious special characters they
possess on the feet, antennae, face, or wings. These characters
are not alike in any two species; they are believed to be of the
nature of ornaments, and according to Professor Aldrich and others
are used as such in courtship.’ This family of flies approaches
very closely to some of the Acalyptrate Muscidae in its char-
acters. It is united by Brauer with Empidae to form the tribe
Orthogenya. Although the species are so numerous and
abundant in Europe, ttle is known as to their metamorphoses.
Some of the larvae frequent trees, living under the bark or in
the overflowing sap, and are believed to be carnivorous; they are
amphipneustic ; a cocoon is formed, and the pupa is remark-
able on account of the existence of two long horns, bearing the
spiracles, on the back of the thorax; the seven pairs of abdominal
spiracles being excessively minute.”

Series 3. Cyclorrhapha Aschiza

Fam, 28. Phoridae.—Small flies, with very convex thorax,
small head, very small two-jointed antennae, bearing a long seta ;
semora more or less broad; wings with two dark, thick, approx-
mate veins, meeting on the front margin near its middle, and
besides these, three or four very fine veins, that run to the margins
in a sub-parallel manner without forming any cells or forks. This
obscure family of flies is of small extent, but its members are ex-
tremely common in Europe and North
America, where they often occur in
numbers running on the windows of

houses. It is one of the most isolated
Tic. °36.—Wing of Zrinewra groups of Diptera, and great difference

werrima, one of the Phor- of opinion prevails as to its classifica-
idae. Britain.

tion. The wing-nervuration is peculiar

(but varies somewhat in the species), the total absence of

any cross-veins even on the basal part of the wing being

remarkable. There are bristles on the head and thorax, but

they are not arranged in a regular manner. The larvae

live in a great variety of animal and vegetable decaying matter,
1 Amer. Natural, xxviii. 1894, p. 35.

2 Perris, Ann. Soc. ent. France (4) x. 1870, p. 321, pl. 4; and Laboulbene, op.
(othe (G3) thas USB, jos IOS oll 16

vir PHORIDAE 495

and attack living Insects, and even snails, though probably
only when these are in a sickly or diseased condition. The
metamorphoses of several species have been described.t The
larvae are rather slender, but sub-conical in form, with eleven
segments and a very small head, amphipneustic, the body behind
terminated by some pointed processes. The pupa is remarkable ;
it is contained in a case formed by the contracted and hardened
skin of the larva; though it differs much in form from the
larva the segmentation is distinct, and from the fourth segment
there project two slender processes. These are breathing organs,
attached to the prothorax of the imprisoned pupa; in what
manner they effect a passage through the hardened larval skin
is by no means clear. Perris supposes that holes for them
pre-exist in the larval skin, and that the newly-formed pupa by
restless movements succeeds in bringing the processes into such a
position that they can pass through the holes. The dehiscence
of the puparium seems to occur in
a somewhat irregular manner, as
in Jierodon; it is never Cyclor-
rhaphous, and according to Perris
is occasionally Orthorrhaphous ;
probably there is no ptilinum.

The Insect recently described by
Meinert as Aenigmatias blattoides,
is so anomalous, and so little is
known about it, that it cannot at
present be classified. It is com-
pletely apterous; the arrangement
of the body-segments is unlike that “yy
of Diptera, but the antennae and
mouth-parts are said to be like those of Phoridae. The Insect
was found near Copenhagen under a stone in the runs of Formica
fusca. Meinert thinks it possible that the discovery of the
male may prove Aenigmatias to be really allied to Phoridae,
and Mik suggests that it may be the same as Platyphora lubbocki,
Verrall, known to be parasitic on ants. Dahl recently described
a wingless Dipteron, found living as a parasite on land-snails in
the Bismarck archipelago, under the name of Puliciphora lucifera,

Fic. 237.—Aenigmatias blattoides.

Denmark. (After Meinert.)

1 Perris, Ann. Soc. ent. France (4) x. 1870, p. 354.
2 Ent. Meddelelser, ii. 1890, p. 213.

490 DIPTERA CHAP.

and Wandolleck has recently made for this and some allies the
new family Stethopathidae. It seems doubtful whether these
forms are more than wingless Phoridae.

Fam. 29. Platypezidae.— Small flies, with porrect three-
jointed antennae, first two joints short, third longer, with a
terminal seta; no bristles on the back; hind legs of male, or of
both sexes, with peculiar, broad, flat tarsi; the middle tibiae bear
spurs; there is no empodium. Platypezidae is a small family
of flies, the classification of which has always been a matter of
considerable difficulty, and is still uncertain. The larvae are
broad and flat, fringed at the margin with twenty-six spines ; they
live between the lamellae of Agaric fungi. At pupation the form
alters but little ; the imago emerges by a horizontal cleft occurring
at the margins of segments two and four.' We have four genera
(Opetia, Platycnema, Platypeza, Callomyia), and nearly a score of
species of Platypezidae in our British list, but very little seems to
be known about them. There is much difference im the eyes of
the sexes, in some at any rate of the species, they being large and
contiguous in the male, but widely separated in the female.

Fam. 30. Pipunculidae..— Small flies, with very short antennae
bearing a long seta that is not terminal ; head almost globular,
formed, except at the back, almost entirely by the large conjoined
eyes; the head is only slightly smaller in the female, but in
the male the eyes are more approzimate at the top. This is
another of the small fami-
lies of flies, that seems dis-
tinct from any other, though
possessing no very im-
portant characters. In many
of the flies that have very
large. eyes, the head is
Fie. 238.—Head of Pipwaculus sp. A, Seen from either flattened (7.e. com-

mete ase showing an antenna pressed from before back-

; wards, as- in Tabanidae,

Asilidae), or forced beneath the humped thorax (as in Acro-
ceridae), but neither of these conditions exists in Pipunculus ;
in them the head extends far forwards, so that the area of the

1 Frauenfeld, Verh. Ges. Wien, xx. Parole pleats
2 For monograph of Pipunculidae, see Becker, Berlin. ent. Zeitschr. xlii. 1897,
pp. 25-100.

vil PIPUNCULIDAE—CONOPIDAE 497

eye compared with the size of the body is perhaps greater than
in any other Diptera. The general form is somewhat that of
Anthrax, but the venation on the hind part of the wing is much
less complex. There is a remarkable difference between the facets
on the front and the back of these great eyes. We have three
genera and about a dozen species of Pipunculidae in Britain but
apparently they are far from common Insects. What is known
about the life-history is almost confined to an imperfect observa-
tion by Boheman, who found the larva of P. fuscipes living after
the manner of a Hymenopterous parasite in the body of a small
Homopterous Insect." The pupa seems to be of the type of that
of Syrphidae.

Fam. 31. Conopidae.—Hlegant flies of moderate size, of varied
colours, with abdomen slender at the base, at the tip strongly
incurved and thicker; antennae inserted close together on a
prominence, three-jointed, first joint sometimes very short. The
upper surface of the body without bristles or with but few. There
is a slender, elongate proboscis, which is retractile and usually
invisible. This rather small family of flies includes some of the
most remarkable forms of Diptera; it includes two divisions,
the Conopinae with long antennae terminated by a very minute
pointed process, and Myopinae with shorter antennae bearing a
hair that is not placed at the end of the third joint. The
former are the more wasp-like and elegant; the Myopinae being
much more like ordinary flies, though they frequently have
curious, inflated heads, with a white face. The mode of life of
the larva of Conops is peculiar, it being parasitic in the interior
of Bombus, or other Hymenoptera. They have been found te
attack Bombus, Chalicodoma, Osmia, Vespa, Pompilus, and other
Aculeates. | Williston says that Orthoptera are also attacked.
Conops has been seen to follow Bumble-bees and alight on them,
and Williston says this act is accompanied by oviposition, the
larva that is hatched boring its way into the body of the bee.
Others have supposed that the flies enter the bees’ nests and
place their eggs in the larvae or pupae; but this is uncertain, for
Conops has never been reared from a bee-larva or pupa, though
it has frequently been procured from the imago: cases indeed
having been recorded in which Conops has emerged from the body

1 Ofv. Ak. Forh. xi. 1854, p. 302, pl. v., since confirmed by others, see Giard,

C.R. Ac. Sct. cix. 1889, pp. 79 and 708.
VOL, vil 2K

498 DIPTERA CHAP.

of a Lombus several months after the latter had been killed and
placed in an entomologist’s collection. The larva is broad, and
when full grown apparently occupies nearly all the space of
the interior of the abdomen of the bee; it has very peculiar
terminal stigmata. The pupa is formed in the larval skin,
which is greatly shortened and indurated for the purpose; this
instar bears, in addition to the posterior stigmata, a pair of
shghtly projecting, anterior stigmata. We have several species
of Conopidae in Britain; those belonging to the division
Conopinae are all rare Insects, but the Myopinae are not so
scarce ; these latter are believed to be of similar habits with
the Conopinae, though remarkably little is known about them.
This is another of the numerous families, the relations of which
are still a subject for elucidation. . Brauer places the Conopidae in
his section Schizophora away from Syrphidae, but we do not com-
prehend on what grounds; an inspection of the head shows that
there is no frontal lunule as there is in Eumyiidae ; both I/yopa and
Conops agreeing fairly well with Syrphus as to this. We therefore
place the family in its old position near Syrphus till the relations
with Acalypterate Muscidae shall be better established.

Fam. 32. Syrphidae (/Hover-jlies)—Of moderate or rather
large size, frequently spotted or banded with yellow, with a thick
fleshy proboscis capable of being withdrawn into a cleft on the
under side of the head ; antennae not placed in definite cavities,
three-jointed (usually very short), and leaving a seta that is not
terminal in position, and may be feathered. Squama variable, never
entirely covering the halteres; the chief (third to fifth) longitudinal
veins of the wings connected near their termination by cross-veins
and usually thus forming a sort of short margin parallel with the
hind edge of the wing ; a more or less imperfect false nervure run-
ning between the third and fourth longitudinal nervures ; no em-
podium and generally no distinct system of bristles on the back of
the body. The Syrphidae (Fig. 212) form one of the largest and
best known of all the families of flies ; they abound in our gardens
where, in sunny weather, some species may be nearly always
seen hovering over flowers, or beneath trees in places where the
rays of the sun penetrate amidst the shade. There are two or
three thousand species’ known, so that of course much variety
exists ; some are densely covered with hair (certain Volucella and
others), many are of elegant form, and some bear a consider-

Vil SYRPHIDAE——-HOVER-FLIES 499

able resemblance to Hymenoptera of various groups. The
peculiar veining of the wings permits of their easy identification,
the line of two nervules, approximately parallel with the margin
of the distal part of the wing (Fig. 212, D), and followed by a deep
bay, being eminently characteristic, though there are some excep-
tions; there are a few forms in which the antennae are exceptional
in having a terminal pointed process. The proboscis, besides
the membranous and fleshy lips, consists of a series of pointed
slender lancets, the use of which it is difficult to comprehend, as
the Insects are not known to pierce either animals or vegetables,
their food being chiefly pollen; honey is also doubtless taken
by some species, but the lancet-lke organs appear equally
ill-adapted for dealing with it. The larvae are singularly
diversified ; first, there are the eaters of Aphidae, or green-fly ;
some of these may be generally found on our rose-bushes or on
thistles, when they are much covered with Aphids; they are soft,
maggot-like creatures with a great capacity for changing their
shape and with much power of movement, especially of the
anterior part of the body, which is stretched out and moved
about to obtain and spear their prey: some of them are very
transparent, so that the movements of the internal organs and
their vivid colours can readily be seen: like so many other
carnivorous Insects, their voracity appears to be insatiable. The
larvae of many of the ordinary Hover-flies are of this kind.
Hristalis and its allies are totally different, they lve in water
saturated with filth, or with decaying vegetable matter (the
writer has found many hundreds of the larvae of Myiatropa
florea in a pool of water standing in a hcilow beech-tree). These
rat-tailed maggots are of great interest, but as they have been
described in almost every work on entomology, and as Professor
Miall’ has recently given an excellent account of their pecu-
liarities, we need not now discuss them. Some of the flies of
the genus Hristalis are very like honey-bees, and appear in old
times to have been confounded with them; indeed, Osten Sacken
thinks this resemblance gave rise to the “ Bugonia myth,” a
fable of very ancient origin to the effect that Honey-bees could
be procured from filth, or even putrefying carcases, by the aid of
certain proceedings that savoured slightly of witchcraft, and
may therefore have increased the belief of the operator in the

1 Natural History of Aquatic Insects, 1895, p. 198.

500 DIPTERA CHAP.

possibility of a favourable result. It was certainly not bees that
were produced from the carcases, but Osten Sacken suggests that
Hristalis-flies may have been bred therein.

In the genus Volucella we meet with a third kind of Syrphid
larva. These larvae are pallid, broad and fleshy, surrounded by
numerous angular, somewhat spinose, outgrowths of the body ;
and have behind a pair of combined stigmata, in the neighbour-
hood of which the outgrowths are somewhat larger; these larvae
live in the nests of Bees and Wasps, in which they are abundant.
Some of the Volucella, like many other Syrphidae, bear a con-_
siderable resemblance to Bees or Wasps, and this has given rise to
a inodern fable about them that appears to have no more legiti-
mate basis of fact than the ancient Bees-born-of-carcases myth.
It was formerly assumed that the Volucella-larvae lived on the
larvae of the Bees, and that the parent flies were providentially
endowed with a bee-like appearance that they might obtain
entrance into the Bees’ nests without being detected, and then
carry out their nefarious intention of laying eggs that would
hatch into larvae and subsequently destroy the larvae of the Bees.
Some hard-hearted critic remarked that it was easy to understand
that providence should display so great a solicitude for the welfare
of the Volucella, but that it was difficult to comprehend how it
could be, at the same time, so totally indifferent to the welfare of
the Bees, More recently the tale has been revived and cited as
an instance of the value of deceptive resemblance resulting from
the action of natural selection, without reference to providence.
There are, however, no facts to support any theory on the subject.
Very little indeed is actually known as to the habits of Volu-
cella in either the larval or imaginal instars; but the little that
is known tends to the view that the presence of the Volucella
in the nests is advantageous to both Fly and Bee. Nicolas has
seen Volucella zonaria enter the nest of a Wasp; it settled at a
little distance and walked in without any fuss being made. Erné
has watched the Volucella-larvae in the nests, and he thinks that
they eat the waste or dejections of the larvae. The writer kept
under observation Volucella-larvae and portions of the cells of
Bombus, containing some larvae and pupae of the Bees and some
honey, but the fly-larvae did not during some weeks touch any of
the Bees or honey, and ultimately died, presumably of starvation.
Subsequently, he experimented with Volucella-larvae and a portion

vu SYRPHIDAE—-HOVER-FLIES 5OI

of the comb of wasps containing pupae, and again found that the
flies did not attack the Hymenoptera; but on breaking a puna
of the Wasp in two, the fly-larvae attacked it unmediately and
eagerly; so ‘that the evidence goes to show that the Volucella-
larvae act as scavengers in the nests of the Hymenoptera.
Kiinekel d’Herculais has published an elaborate work on the
European Volucella; it is remarkable for the beauty of the
plates illustrating the structure, anatomy and development, but
throws little direct light on the natural history of the Insects.
V. bombylans, one of the most abundant of our British species,
appears in two forms, each of which has a considerable resem-
blance to a Bombus, and it has been supposed that each of the
two forms is specially connected with the Bee it resembles, but
there is no evidence to support this idea; indeed, there is some
little evidence to the contrary. The genus Merodon has larvae
somewhat similar to those of Volucella, but they live in bulbs
of Narcissus; M. equestris has been the cause of much loss to the
growers of Dutch bulbs; this Fly is interesting on account of its
great variation in colour; it has been described as a whole series
of distinct species.

The most remarkable of the numerous forms of Syrphid larvae
are those of the genus Microdon (Fig. 239), which live in ants’
nests. They have no resemblance to Insect-larvae, and when
first discovered were not only supposed to be little Molluscs, but
were actually described as such under the generic names of
Parmula and Scutelligera. There is no appearance of segmenta-
tion of the body; the upper surface is covered by a sort of
network formed by curved setae, which help to retain a coating
of dirt; there is no trace externally of any head, but on the
under surface there is a minute fold in which such mouth-organs
as may be present are probably concealed; the sides of the body
project so as to form a complex fringing arrangement; the ter-
minal stigmata are very distinct, the lateral processes connected
with them (the “Knospen” of Dr. Meijere), are, however, very
irregular and placed at some distance from the stigmatic scar.
Pupation occurs by the induration of the external covering and
the growth from it, or rather through it, of two short horns in
front. Inside this skin there is formed a soft pupa, of the kind
usual in Cyclorrhaphous flies; the dehiscence of the external
covering is, however, of unusual nature, three little pieces being

502 DIPTERA CHAP.

separated from the anterior part of the upper surface, while the
lower face remains intact. The account of the pupation given
by Elditt’ is not complete: the two horns that project are, it
would appear, not portions of the larval skin, but belong to
the head of the pupa, and according to Elditt are used to effect
the dehiscence of the case for the escape of the fly; there does
not appear to be any head-vesicle. Nothing is known as to
the details of the life of these anomalous larvae. M. Poujade
has described two species found in France in the nests of the
ant Lasius niger” The larva we figure was found by Colonel
Yerbury in nests of an Atta in Portugal, and an almost identical

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Fic. 239.—Larva of Microdon sp. Portugal. A, Dorsal view of the larva, x 4; 1,
the stigmatic structure ; B, posterior view of stigmatic structure; C, a portion of
the marginal fringe of the body.

larva was recently found by Mr. Budgett in Paraguay. The

flies themselves are scarce, Microdon mutabilis (formerly called

M. apiformis) being one of the rarest of British flies. They have

the antennae longer than is usual in Syrphidae, and the cross-

veins at the outside of the wing are irregularly placed, so that
the contour is very irregular: the resemblance to bees is very
marked, and in some of the South American forms the hind legs
are flattened and hairy like those of bees. The oviposition of

Microdon has been observed by Verhoeff ;° he noticed that the fly

was frequently driven away by the ants—in this case, Mormica

sanguinea—but returned undiscouraged to its task.

A brief résumé of the diverse modes of life of Syrphid larvae
fas been given by Perris,’ and he also gives some information as
to the curious horns of the pupae, but this latter point much

1 Ent. Zeit. Stettin, vi. 1845, p. 384, pl. i.

2 Ann. Soc. ent. France (6) iii. 1883, p. 23, pl. i.
* Ent. Nachr. xviii. 1892, p. 13. 4 Ann. Soc. ent. France (4) x. 1870, p. 330.

VII MUSCIDAE ACALYPTRATAE 503

wants elucidation. Whether the Syrphidae, or some of them,
possess a ptilinum that helps them to emerge from the pupa is
more than doubtful, though its existence has been aftirmed by
several authors of good repute."

Series 4. Cyclorrhapha Schizophora

Fam. 33. Muscidae acalyptratae—This group of flies has
been the least studied of all the Diptera; it is generally treated
as composed of twenty or thirty different families distinguished

Fia. 240.—Diopsis
apicalis. Natal.
A, The fly ; B, ex-
tremity of cephalic
protuberance, more
magnified. a,
The eye; 0b, the
antenna; C, middle
of head, front
view ; ¢, ocelli.

se =

by very slight characters. It is, however, generally admitted by
systematists that these assemblages have not the value of the
families of the other divisions of Diptera, and some even go so
far as to say that they are altogether only equivalent to a single
family. We do not therefore think it necessary to define each
one seriatim ; we shall merely mention their names, and allude
to certain points of interest connected with them. Taken collec-
tively they may be defined as very small flies, with three-jointed
antennae (frequently looking as if only two-jointed), bearing a
bristle that is not terminally placed ; frequently either destitute of
squamae or having these imperfectly developed so as not to cover the
halteres ; and possessing a comparatively simple system of nervura-
tion, the chief nervures being nearly straight, so that consequently
few cells are formed. These characters will distinguish the group

1 See on this difficult subject, Becher, Wien. ent. Zeit. i. 1882, p. 49.

504 DIPTERA CHAP.

from all the other Diptera except from forms of Aschiza, and
from certain Anthomyiidae, with both of which the Acalyptratae
are really intimately connected. Considerable difference of
opinion prevails as to the number of these divisions, but the
families usually recognised are :—

1. Doryceridae. 12. Dryomyzidae. 21. Trypetidae.
2. Tetanoceridae. 13. Borboridae. 99 JSapromyzidae
3. Sciomyzidae. 14. Phycodromidae. ““\incl.. Lonchaeidae.
4, Diopsidae. 15. Thyreophoridae. 23. Rhopalomeridae.
5. Celyphidae. 16. J Scatophagidae. 24. Ortalidae.
6 /Sepsidae * \(=Scatomyzidae). ( Agromyzidae
* (incl. Piophilidae. . f Geomyzidae 25. incl. Phytomyzi-
~ f Chloropidae ts \eiavels Opomyzidae. dae.
Ms \ (= Oscinidae). 18 { Drosophilidae ; 26. Milichiidae.
8. Ulidiidae. “* \inel. Asteidae. 27. Octhiphilidae.
9. Platystomidae. 19. Psilidae. 28. Heteroneuridae.
10. Ephydridae. 20 f Tanypezidae 29. Cordyluridae.
11. Helomyzidae. “~~ \ (= Micropezidae).

Brauer associates Conopidae with Acalyptrate Muscids, and
calls the Group Holometopa; applying the term Schizometopa
to the Calyptrate Muscidae.

No generalisation can yet be made as to the larvae of these
divisions, neither can any characters be pointed out by which
they can be distinguished from the larvae of the following
families. In their habits they have nothing specially distinctive,
and may be said to resemble the Anthomyiidae, vegetable
matter being more used as food than animal; many of them
mine in the leaves or stems of plants; in the genus Dorycera the
larva is aquatic, mining in the leaves of water-plants, and in
Ephydridae several kinds of aquatic larvae are found, some of
which are said to resemble the rat-tailed larvae of Syrphidae ;
certain of these larvae occur in prodigious quantities in lakes,
and the Insects in some of their early stages serve the Mexicans
as food, the eggs being called Ahuatle, the larvae Pusci, the
pupae Koo-chah-bee. Some of the larvae of the Sciomyzidae are
also aquatic: that of Tetanocera ferruginea is said by Dufour to
consist only of eight segments, and to be metapneustic; Brauer
considers the <Acalyptrate larvae to be, however, in general,
amphipneustic, ike those of Calyptratae. The Chloropidae are
a very important family owing to their occasional excessive
multiplication, and to their living on cereals and other grasses,
various parts of which they attack, sometimes causing great losses
to the agriculturist. The species of the genus Chlorops are

VII MUSCIDAE ACALYPTRATAE 505

famous for the curious habit of entermg human habitations in
great swarms: frequently many millions being found in a single
apartment. Instances of this habit have been recorded both in
France and England, Cambridge being perhaps the place where the
phenomenon is most persistently exhibited. In the year 1831 an
enormous swarm of C. /ineata was found in the Provost’s Lodge
at King’s College and was recorded by Leonard Jenyns; in
1870 another swarm occurred in the same house if not in the
same room.’ Of late years such swarms have occurred in certain
apartments in the Museums (which are not far from King’s
College), and always in the same apartments. No clue whatever
can be obtained as to their origin; and the manner in which
these flies are guided to a small area in

numbers that must be seen to be be- we:

lieved, is most mysterious. These swarms
always occur in the autumn, and it has
been suggested that the individuals are

seeking winter quarters. by we :
Several members of the Acalyptratae Nh) 3 |

have small wings or are wingless, as in th, Pi

some of our species of Borborus. The a ony yee

Diopsidae—none of which are European
have the sides of the head produced
into long horns, at the extremity of which
are placed the eyes and antennae; these
curiosities (Fig. 240) are apparently com-
mon in both Hindostan and Africa. In
the horned flies of the genus Hlaphomyia,
parts of the head are prolonged into
horns of very diverse forms according t0 fy¢, 241.—Céelyphus (Para-
the species, but bearing on the whole a ere) ee (earn
great resemblance to miniature stag- hover a, Ree ttica b,
horns. A genus (Giraffomyia) with along — D8 i eee Pre
neck, and with partially segmented appen- — downwards; a, scutellum ;
dages, instead of horns on the head, has sitomen. Cy Pate
been recently discovered by Dr. Arthur

Willey in New Britain. Equally remarkable are ae species of
Celyphus ; they do not look like flies at all, owing to the scutellum
being inflated and enlarged so as to cover all the posterior parts

1 Loudon’s Magazine, v. 1832, p. 302; P. ent. Soc. London, 1871, p. x.

506 DIPTERA CHAP.

of the body as in the Scutellerid Hemiptera: the wings are
entirely concealed, and the abdomen is reduced to a plate, with
its orifice beneath, not terminal; the surface of the body is
highly polished and destitute of bristles. Whether this is a
mimetic form, occurring in association with similar-looking
Bugs is not known. The North American genus Yowotrypana
is furnished with a long ovipositor; and in this and in the shape
of the body resembles the parasitic Hymenoptera. This genus
was placed by Gerstaecker in Ortalidae, but is considered by
later writers to be a member of the Trypetidae. This latter
family is of considerable extent, and is remarkable amongst the
Diptera for the way in which the wings of many of its members
are ornamented by an elaborate system of spots or marks, vary-
ing according to the species.

Fam. 34. Anthomyiidae. — lies similar in appearance to
the House-fly ; the main vein posterior to the middle of the wing
(4th longitudinal) continued straight to the margin, not turned
upwards. Eyes of the male frequently large and contiguous,
bristle of antenna either feathery or bare. This very large family
of flies is one of the most difficult and unattractive of the Order.
Many of its members come close to the Acalyptrate Muscidae
from which they are distinguished by the fact that a well-
developed squama covers the halteres; others come quite as
close to the Tachinidae, Muscidae and Sarcophagidae, but may
readily be separated by the simple, not angulate, main vein
of the wing. The larval habits are varied. Many attack
vegetables, produce disintegration in them, thus facilitating de-
composition. Anthomyia brassicae is renowned amongst market
gardeners on account of its destructive habits. A. cana, on the
contrary, is beneficial by destroying the migratory Locust
Schistocerca peregrina ; and in North America, A. angustifrons
performs a similar office with Caloptenus spretus. One or two
species have been found living in birds; in one case on the head of
a species of Spermophila, in another case on a tumour of the wing
of a Woodpecker. Hylemyia strigosa, a dung-frequenting species,
has the peculiar habit of producing living larvae, one at a time;
these larvae are so large that it would be supposed they are full
grown, but this is not the case, they are really only in the first
stage, an unusual amount of growth being accomplished in this
stadium. Spilogaster angelicae, on the other hand, according to

VII AN THOMYIIDAE——TACHINIDAE 507

Portschinsky, lays a small number of very large eggs, and the result-
ing larvae pass from the first to the third stage of development,
omitting the second stage that is usual in Eumyiid Muscidae.’
Fam. 35. Tachinidae.—J/irst posterior cell of wing nearly or
quite closed. Squamae large, covering the halteres: antennal arista
bare: upper surface of body usually bristly. This isan enormous
family of flies, the larvae of which live parasitically in other living
Insects, Lepidopterous larvae being especially haunted. Many
have been reared from the Insects in which they live, but beyond
this little is known of the life-histories,and still less of the structure
of the larvae of the Tachinidae, although these Insects are of the
very first importance in the economy of Nature. The eggs are
usually deposited by the parent-flies near or on the head of the

Fic. 242.—Ugimyia sericariae. A, The perfect fly, x 3; B, tracheal chamber of a silk-
worm, with body of a larva of Ugimyia projecting ; a, front part of the maggot ;
b, stigmatic orifice of the maggot ; c, stigma of the silkworm. (After Sasaki.)

victim ; Riley supposed that the fly buzzes about the victim and
deposits an egg with rapidity, but a circumstantial account given
by Weeks?” discloses a very different process: the fly he watched
sat on a leaf quietly facing a caterpillar of Datana engaged in
feeding at a distance of rather less than a quarter of an inch.
“Seizing a moment when the head of the larva was likely
to remain stationary, the fly stealthily and rapidly bent her
abdomen downward and extended from the last segment what
proved to be an ovipositor. This passed forward beneath her
body and between the legs until it projected beyond and nearly
on a level with the head of the fly and came in contact with the
eye of the larva upon which an ege was deposited,’ making an
addition to five already there. Ugimyia sericariae does great

' Baron von Osten Sacken informs the writer that this statement has since been

withdrawn by Portschinsky as being erroneous.
2 Ent. Amer. iii. 1887, p. 126.

508 DIPTERA CHAP.

harm in Japan by attacking the silkworm, and in the case of this
tly the eggs are believed to be introduced into the victim by
being laid on mulberry leaves and swallowed with the food;
several observers agree as to the eggs being laid on the leaves, but
the fact that they are swallowed by the silkworm is not so certain.
Sasaki has given an extremely interesting account of the develop-
ment of this larva! According to him, the young larva, after
hatching in the alimentary canal, bores through it, and enters a
nerve-ganglion, feeding there for about a week, after which the
necessity for air becoming greater, as usual with larvae, the
maggot leaves the nervous system and enters the tracheal system,
boring into a tube near a stigmatic orifice of the silkworm, where
it forms a chamber for itself by biting portions of the tissues and
fastening them together with saliva. In this it completes its

Fic, 243.—Diagrammatic section of silkworm to show the habits of Ugimyia. a, Young
larva ; b, egg of Ugimyia in stomach of the silkworm ; ¢, larva in a nerve-ganglion ; _
d, larva entering a ganglion ; e, larva embedded in tracheal chamber, as shown in
Fig. 242, B. (After Sasaki. )

erowth, feeding on the interior of the silkworm with its anterior
part, and breathing through the stigmatic orifice of its host;
after this it makes its exit and buries itself deeply in the ground,
where it pupates. The work of rupturing the puparium by the
use of the ptilinum is fully described by Sasaki, and also the fact
thatthe fly mounts to the surface of the earth by the aid of this same
peculiar air-bladder, which is alternately contracted and distended.
Five, or more, of the Ugimyia-maggots may be found in one
caterpillar, but only one of them reaches maturity, and emerges
from the body. The Tachinid flies appear to waste a large pro-
portion of their eggs by injudicious oviposition ; but they make
up for this by the wide circle of their victims, for a single species
has been known to infest Insects of two or three different Orders.

The species of Jiltogramma—of which there are many in
Europe and two in England—live at the expense of Fossorial

1 J. Coll. Japan, i. 1886, pp. 1-46, plates i.-vi.

vu TACHINIDAE 509

Hymenoptera by a curious sort of indirect parasitism. They are
obscure little flies, somewhat resembling the common House-tfly,
but they are adepts on the wing and have the art of ovipositing
with extreme rapidity ; they follow a Hymenopteron as it is carry-
ing the prey to the nest for its young. When the wasp alights
on the ground at the entrance to the nest, the Miltogramma
swoops down and rapidly deposits one or more eggs on the prey
the wasp designs as food for its own young. Afterwards the larvae
ot the fly eat up the food, and in consequence of the greater
rapidity of their growth, the young of the Hymenopteron perishes.
Some of them are said to deposit living larvae, not eggs. Fabre
has drawn a very interesting picture of the relations that exist
between a species of Miltogramma and a Fossorial Wasp of the
genus Bembex.' We may remind the reader that this Hymenop-
teron has not the art of stinging its victims so as to keep them
alive, and that it accordingly feeds its young by returning to
the nest at proper intervals with a fresh supply of food, instead
of provisioning the nest once and for all and then closing it.
This Hymenopteron has a habit of catching the largest and
most active flies-—especially Tabanidae—for the benefit of its
young, and it would therefore be supposed that it would be safe
from the parasitism of a small and feeble fly. On the contrary,
the IMiltogramma adapts its tactics to the special case, and is in
fact aided in doing so by the wasp itself. As if knowing that the
wasp will return to the carefully-closed nest, the Miltagramma
waits near it, and quietly selects the favourable momfoit, when
the wasp is turning round to enter the nest bachtivurds, and
deposits eggs on the prey. It appears from Fabre’sa, ccount that
the Bembex is well aware of the presence of the ‘ty, and would
seem to entertain a great dread of it, as if conscious that it is a
formidable enemy; nevertheless the wasp never attacks the
little fly, but allows it sooner or later to accomplish its purpose,
and will, it appears, even continue to feed the fly-larvae, though
they are the certain destroyers of its own young, thus repeat
ing the relations between cuckoo and sparrow. Most of us
think the wasp stupid, and find its relations to the fly incredible
or contemptible. Fabre takes a contrary view, and looks on it
as a superior Uncle Toby. We sympathise with the charming
French naturalist, without forming an opinion.

' Souvenirs entomologiques, 1879, pp. 246-254.

510 TOTP AGE VAY CHAP.

Doubtless there are many other interesting features to be found
in the lfe-histories of Tachinidae, for in numbers they are legion.
It is probable that we may have 200 species in Britain, and in
other parts of the world they are even more abundant, about
1000 species beimg known in North America.’ The family
Actiidae is at present somewhat doubtful. According to Karsch,
it is a sub-family of Tachinidae; but the fourth longitudinal
vein, it appears, 1s straight.

Fam. 36. Dexiidae—7Zhese Insects are distinguished from
Tachinidae by the bristle of the antennae being pubescent, and
the legs usually longer. The larvae, so far as known, are found
in various Insects, especially in Coleoptera, and have also been
found in snails. There are eleven British genera, and about a
score of species.

Fam. 37. Sarcophagidae. — Distinguished from Muscidae
and Tachinidae by little more than that the bristle of the
antennae is feathery at the base but hair-like and very fine at
the tip.—Sarcophaga carnaria is one of the commonest British
Insects; it is lke the Blow-fly, though rather longer, con-
spicuously grey and black, with the thorax distinctly striped,
and the pulvilli very conspicuous in the live fly. Cynomyia
mortuorum is a bright blue fly rather larger than the Blow-tly,
of which it is a competitor; but in this country an unsuccessful
one. The larvae of the two Insects are found together, and are
said to "4, pute indistinguishable. Cynomyva is said to lay only
about hag» <e@ number of eggs that the Blow-tly does, but it
appears © if + \in the year, and to this is attributed the fact
that it is ap, altogether crowded out of existence by the more
prolific CallujSora. The species of Sarcophagidae are usually
viviparous, and one of them, Sarcophila magnifica (wohlfahrti),
has the habit of occasionally depositing its progeny in the
nostrils of mammals, and even of human beings, causing horrible
sufferings and occasionally death: it is said to be not uncommon
in Europe, but does not occur in Britain. The genus Swrcophaga
is numerous in species, and many of them are beneficial. Sir
Sidney Saunders found in the Troad that Locusts were destroyed
by the larvae of a Sarcophaga living in their bodies; and

1 A list of the Insects known to be attacked by Dipterous parasites has been
given by Brauer and Bergenstamm, Denk. Ak. Wien, 1xi. 1895.
2 Berlin. ent. Zeit, xxx. 1886, p. 185. .

Vu SARCOPHAGIDAE—MUSCIDAE S511

Kiinckel has recently observed that in Algeria several species
of this genus attack Locusts and destroy large quantities by
depositing living larvae in the Orthoptera. In North America
the Army-worm is decimated by species of Sarcophaga.

Many of these Insects, when food is scarce, eat their own
species with eagerness, and it seems probable that this habit is
beneficial to the species. The parent-fly in such cases usually
deposits more eggs than there is food for, thus ensuring that
every portion of the food will be rapidly consumed, after which
the partially-grown larvae complete their development by the
aid of cannibalism. It is thus ensured that the food will raise
up as many individuals as possible.

Fam. 38. Muscidae——JSristle of antennae feathered. This
family contains many of the most abundant flies, including the
House-fly, Blue-bottles or Blow-flies, Green-bottles, and other
forms which, though very common, are perhaps not dis-
criminated from one another by those who are not entomologists.
The larvae live on carrion and decaying or excrementitious
matters. The common House-fly, J/wsca domestica, runs through
its life-history in a very short time. It lays about 150 very small
eges on dung or any kind of soft damp filth; the larvae hatch
in a day or two and feed on the refuse; they may be full-grown
in five or six days, and, then pupating, may in another week emerge
as perfect flies. Hence it is no wonder that they increase to
enormous numbers in favourable climates. They are thought to
pass the winter chiefly in the pupal state. The House-fly is
now very widely distributed over the world; it sometimes occurs
in large numbers away from the dwellings of man. Of Blow-
flies there are two common species in this country, Calliphora
erythrocephala and C. vomitoria. The Green-bottle flies, of which
there are several species, belonging to the genus Lucilia, have the
same habits as Blow-flies, though they do not commonly enter
houses. The larvae are said to be indistinguishable from those of
Calliphora.

The larvae of Eumyiid Muscidae are, when first hatched,
metapneustic, but subsequently an anterior pair of stigmata
appears, so that the larva becomes amphipneustic. They
usually go through three stages, distinguished by the condition
of the posterior stigmata. In the early instar these have a
single heart-shaped fissure, in the second stage two fissures exist,

512 DIPTERA CHAP.

while in the third instar there is a greater diversity in the
condition of the breathing apertures.

The various forms of Muscidae show considerable distinctions
in the details of their natural history, and these in certain species
vary according to the locality. This subject has been chiefly
studied by Portschinsky, a Russian naturalist, and a very interest-
ing summary of his results has been given by Osten Sacken,'
to which the student interested in the subject will do well
to refer.

A few years ago a great deal of damage was caused in the
Netherlands by Lucilia sericata, a Green-bottle-fly, extremely
similar to our common ZL. caesar, which deposited its eggs in
great quantities on sheep amongst their wool. This epidemic
wes attributed to the importation of sheep from England; but,
according to Karsch, there is reason to suppose that the fly was
really introduced from Southern Europe or Asia Minor.

The larvae of species of the genus Lucilia sometimes attack
man and animals in South America, but fortunately not in this
country, The larva of Lucilia (Compsomyia) macellaria is called
the secrew-worm, and is the best known of the forms that infest
man, the larvae living in the nasal fossae and frontal sinuses,
and causing great suffering. The fly is common in North
America, but is said never to attack man farther north than
in Kansas. A little fly (Stomoxys calcitrans), very like the
common house-fly though rather more distinctly spotted with
grey and black, and with a fine, hard, exserted proboscis,
frequently enters our houses and inflicts a bite or prick on us.
It is commonly mistaken fer an ill-natured house-fly that has
taken to biting. It is frequently a source of irritation to cattle.
A closely allied fly, Haematobia serrata, is very injurious to cattle
in North America, but the same species causes no serious annoy-
ance in England. We may mention that the various attacks of
Dipterous larvae on man have received the general name
“ myiasis.”

The Tse-tse fly (Glossina morsitans), another ally of Stomoxys,
is not very dissimilar in size and shape to the blow-fly.’

1 Berlin. ent. Zeitschr. xxxi. 1887, p. 17.

2 Biol. Centralbl. vii. 1887, p. 521.

3 For an account of the habits of this fly, see Kirk, J. Linn, Soc. viii. 1865, pp.
149-156 ; and for a bibliographic list, Wulp, Tijdschr. Ent. xxvii. 1884, p. xci. and
pp. 143-140.

VII MUSCIDAE—TSE-TSE FLY 513

Tt bites man and animals in South Africa, and if it have
previously bitten an animal
whose blood was charged with
the Haematozoa that really con-
stitute the disease called Nagana
(fly-disease), it inoculates the
healthy animal with the dis-
ease ; fortunately only some
species are susceptible, and man
is not amongst them. It has
recently been shown by Surgeon
Bruce’ that this fly multiples
by producing, one at a time, a
full-grown larva, which imme-

Fic. 244.—The Tse-tse fly (Glossina morsi- :
tans). A, The fly with three divisions iately changes to a pupa, as

of the proboscis projecting; B, adult

Tae (Galan do the members of the series
3 ’ a

Pupipara. There are already
known other Muscid flies with peculiarities in their modes of
reproduction, so that it is far from impossible that the various con-
ditions between ordinary egg-laying and full-grown larva- or pupa-
production may be found to exist. Although it has been supposed
that the Tse-tse fly is a formidable obstacle to the occupation of
Africa by civilised men, there is reason to suppose that this will
not ultimately prove to be the case. It only produces disease when
this pre-exists in animals in the neighbourhood ; only certain
species are liable to it; and there is some evidence to the effect
that even these may in the course of a succession of generations
become capable of resisting the disease inoculated by the fly. As
long ago as 1878 Dr. Drysdale suggested * that this fly only pro-
duces disease by inoculating a blood-parasite, and all the evidence
that has since been received tends to show that his idea is correct.

Although the facts we have mentioned above would lead to
the supposition that Muscidae are unmitigated nuisances, yet it
is probable that such an idea is the reverse of the truth, and
that on the whole their operations are beneficial. It would be
difficult to overestimate their value as scavengers. And in
addition to this they destroy injurious creatures. Thus in
Algeria Idia fasciata, a fly like the House - fly, destroys the

1 Preliminary Report on the Tse-tse Fly Disease, 1895.
2 P. Liverpool Soc. xxxiii. 1878, p. 13, note.
VOL. VI 2 1

514 DIPTERA CHAP.

dreaded migratory Locust Schistocerca peregrina im great quantities,
by the larvae eating the eggs of the Locust. The female of this
fly, in order to reach the desired food, penetrates from one to
three inches below the surface of the ground.

Fam. 39. Oestridae (ot-flies)-—Rather large or very large
flies, with extremely short antennae, bearing a segmented arista, the
front of the head prominent, the posterior part of the wings fre-
quently rough, and with but few veins: the mouth usually atrophied,
the trophi being represented only by tubercles ; larvae living in Verte-
brates, usually Mammals, though it is possible that a few occur in
Birds and even in Reptiles. This is a family of small extent, less
than 100 species being known from all the world, yet it is of
much interest on account of the habits of its members, which,
though of large size, live entirely at the expense of living Verte-
brates, to the viscera or other structures of which they have definite
relations, varying according to the species. Some (Gastrophilus, etc.)
live in the alimentary canal; others (Hypoderma, ete.) are encysted
in or under the skin; while others (Oestrus, etc.) occupy the respir-
atory passages. As many of them attack the animals used by man,
and some of them do not spare man himself, they have attracted
much attention, and there 1s an extensive literature connected
with them; nevertheless the life-histories are still very incom-
pletely known. Indeed, the group is from all points of view a
most difficult one, it being almost impossible to define the family
ewing to the great differences that exist in important points.
Some think the family will ultimately be dismembered; and
Girschner has recently proposed to treat it as a division of
Tachinidae. The chief authority is Brauer, in whose writings
the student will find nearly all that is known about Oestridae.*
Some of them exist in considerable numbers Gt is believed that
they are now not so common as formerly), and yet the flies. are
but rarely met with, their habits being in many respects peculiar.
Some of them, for purposes of repose, frequent the summits of
mountains, or towers, or lofty trees. Some have great powers of
hwnming; none of them are known to bite their victims, indeed
the atrophied mouth of most of the Oestridae forbids such a pro-
ceeding. Some deposit their eges on the hairs of the beasts from

1 We may specially mention the monograph of Oestridae, published in 1863 by
the K. k. Zool.- Bot. Ges. Wien, and supplements in Wien. ent. Zeit. v. vi. 1886,
1887 ; these include copious bibliographic lists.

vi OESTRIDAE—BOT-FLIES BIS

which the larvae are to draw their nutriment, but others place
their larvae, already hatched, in the entrances of the nasal
passages. They do not feed on the blood or tissues of their
victims, but on the secretions, and these are generally altered or
increased by the irritation induced by the presence of the un-
welcome guests. It would appear, on the whole, that their presence
is less injurious than would be expected, and as they always quit
the bodies of their hosts for the purposes of pupation, a natural
end is put to their attacks. We have ten species in Britain, the
animals attacked being the ox, the horse, the ass, the sheep, and
the red deer ; others occasionally occur in connexion with animals

Fig. 245.—Cephalomyia maculata, a Bot-fly of the camel. Arabia. A, The fly with
extended wings ; B, under aspect of the head : a; antenna ; 6, the obsolete mouth-parts.

in menageries. The eggs of Gastrophilus equi are placed by the
fly, when on the wing, on the hair of horses near the front parts
of the body, frequently near the knee, and, after hatching, the
young larvae pass into the stomach of the horse either by being
licked off, or by their own locomotion; in the stomach they be-
come hooked to the walls, and after being full grown pass out
with the excreta: the Bots—as these larvae are called—are some-
times very numerous in the stomach, for a fly will lay as many
as four or five hundred eges on a single horse: in the case of
weakly animals, perforation of the stomach has been known to
occur in consequence of the habit of the Bot of burying itself to
a greater or less extent in the walls of the stomach. Hypoderma
hovis and H. lineata attack the ox, and the larvae cause tumours
in the skin along the middle part of the back. It was formerly

516 DIPTERA CHAP.

inferred from this that the fly places its eggs in this situation,
and as the cattle are known to dread and flee from the fly, it
was supposed to be on account of the pain inflicted when the ege
was thrust through the skin. Recent observations have shown
that these views are erroneous, but much still remains to be ascer-
tained. The details of oviposition are not yet fully known,
but it appears that the eggs are laid on the lower parts of the
body, especially near the heels, and that they hatch very speedily."
As the imago of Hypoderma appears for only a very short period in
the summer, the time of the oviposition is certain. The newly-
disclosed larva is considerably different from the more advanced
‘instar found in the skin of the back; moreover, a long period of
many months intervenes between the hatching of the larva and
its appearance in the part mentioned. Brauer has shown that-
when the grub is first found in that situation it is entirely sub-
cutaneous. Hence it would be inferred that the newly-hatched
larva penetrated the skin probably near the spot it was deposited
on, and passed a period in subcutaneous wandering, on the whole
going upwards till it arrived at the uppermost part: that after
moulting, and in consequence of greater need for air, it then
pierced the skin, and brought its breathing organs into contact
with the external air; that the irritation caused by the admission
of air induced a purulent secretion, and caused the larva to be
enclosed in a capsule. Dr. Cooper Curtice has however found,
in the oesophagus of cattle, larvae that he considers to be quite
the same as those known to be the young of Hypoderma ; and
if this prove to be correct, his inference that the young larvae
are licked up by the cattle and taken into the mouth becomes
probable. The larva, according to this view, subsequently pierces
the oesophagus and becomes subcutaneous by passing through the
intervening tissues. The later history of the grub is briefly, that
when full grown it somewhat enlarges the external orifice of its
cyst, and by contractions and expansions of the body, passes to
the surface, falls to the ground, buries itself and becomes a pupa.
If Dr. Curtice be correct, there should, of course, be as many,
if not more, larvae found in the oesophagus as in the back
of the animal; but, so far as is known, this is not the case, and
we shall not be surprised if the normal course of development be
found different from what Dr. Curtice supposes it to be. His

1 Riley, Insect Life, iv. 1892, p. 302.

VII OESTRIDAE—PUPIPARA 517

observations relate to Hypoderma lineata. Our common British
species is usually supposed to be A. bovis ; but from recent ob-
servations it seems probable that most of the “Ox-warbles” of this
country are really due to the larvae of HZ. lineata.

The history of Oestrus ovis, which attacks the sheep, is also
incompletely known, but appears to be much simpler. This fly
is viviparous, and deposits its young larvae at the entrance of the
nasal passages of the sheep, thereby causing extreme annoyance
to the animal. The larvae penetrate to the frontal sinuses to
complete their growth. The duration of their lives is unknown,
for it is commonly the case that larvae of various sizes are found
together. Cephenomyia rufibarbis has recently been found in
Scotland. It attacks the Red deer, and its life-history is similar
to that of Oestrus ovis, though the larvae apparently prefer to
attain their full growth in the pharynx of the deer.

In reference to the Oestridae that attack man, we may merely
mention that the larva of the Hypoderma of the ox is occasionally
found in Europe infesting human beings, but only as an extremely
rare and exceptional event; and that only those engaged in
attending on cattle are attacked; from which it is inferred that

_ the flies are deceived by an odour emanating from the garments.

Im America numerous cases are known of Oestrid larvae being
taken from the body of man, but information about them is very
scanty. It appears, however, that there are at least four species,
one of which, Dermatobia noxialis, is known as a fly as well as a
larva. Whether any of these are peculiar to man is uncertain.'
There are several larvae of Muscidae that have similar habits to the
Oestridae ; hence the statements that exist as to larvae being found
in birds and reptiles cannot be considered to apply to members of
the latter family until the larvae have been studied by an expert.

The family Ctenostylidae has been established by Bigot for a
South American Insect, of which only a single individual exists
in collections. It is doubtful whether it can be referred to
Oestridae.”

Series V. Pupipara

The four families included in this Series are, with the
exception of the Hippoboscidae, very little known. Mest of
1 See Blanchard, Ann. Soc. ent. France (7) ii. 1892, pp. 109, 154.

2 See Bigot, Ann. Soc. ent. France (6) ii. 1882, p. 21, Brauer, Monograph, 1863,
p- 51, and Wien. ent. Zeit. vi. 1887, p. 75.

518 DIPTERA CHAP.

them live by sucking blood from Mammals and Birds, and some-
times they are wingless parasites. The single member of the
family Braulidae lives on bees. The term Pupipara is erronecus,
and it would be better to revert to Réaumur’s prior appellation
Nymphipara. Miiggenburg has suggested that the division is
not a natural one, the points of resemblance that exist between
its members being probably the results of convergence. Recent
discoveries as to the modes of bringing forth of Muscidae give
additional force to this suggestion. A satisfactory definition of
the group in its present extent seems impossible.

Fam. 40. Hippoboscidae.— Wings very variable, sometimes
present and large, then with waved surface and thick nervures
confined to the anterior and basal part ; sometimes mere strips,
sometimes entirely absent. Certain members of this family are well
known, the Forest-fly, or Horse-fly, and the Sheep-tick belonging to
it. The proboscis is of peculiar formation, and not lke that of other
flies. Seen externally it consists of two elongate, closely adapted,
hard flaps ; these are capable of diverging laterally to allow an inner
tube to be exserted from the head. The details and morphology
of the structure have recently been discussed by Miiggenburg.!
Melophagus ovinus, commonly called the Sheep-tick, is formed for
creeping about on the skin of the sheep beneath the wool, and
may consequently be procured with ease at the period of sheep-
shearing: it has no resemblance to a fly, and it is difficult to
persuade the uninitiated that it is such. Mippobosca equina (called
in this country the Forest-fly, perhaps because it 1s better known
in the New Forest than elsewhere), looks like a fly, but will be
readily recognised by. the two little cavities on the head, one
close to each eye, in which the antennae are concealed, only the
fine bristle projecting. Very little seeras to be known as to
the Natural History of this fly. Lipoptena cervi lives on the
Red deer; the perfect Insect has apparently a long life, and both
sexes may be found in a wingless state on the deer all through
the winter. When first disclosed in the summer they are how-
ever provided with wings, but when they have found a suitable
host they bite off, or cast, the wings. The female, it appears, does
this more promptly than the male, so that it is difficult to get
winged individuals of the former sex.” Most of the known

~

1 Arch. Naturgesch. lviii. 1.1892, pp. 287-322, pls. xv. xvi.
2 Stein, Deutsche ent. Zeit. xxi. 1877, p. 297.

Vu HIPPOBOSCIDAE 519

Hippoboscidae live on birds, and are apparently specially fond of
the Swallow tribe. They are all winged, though in some species
the wings are very small. The bird-
infesting Hippoboscidae have been very
little studied, and will probably form a
distinct family; the antennae of Sten-
opteryx hirundinis are quite different
from those of Hippobosca. The devel-
opment is remarkable, and has been
studied by Leuckart! and by Pratt? in
the case of Melophagus ovinus. The
ovaries are peculiarly formed, and pro-
duce one large egg at a time; this
passes into’ the dilated oviduct, and there
goes through its full growth and a cer- Ee ety eS
: arene Fia. 246.—Diagrammatic section
tain amount of development; it is then of the larva of Melophagus
exuded sand undergome; littlesor ino, -07rus. (After Exatt.) 2,
: : mouth ; 4, suctorial pouch ;
change of form becomes externally — ¢, imaginal disc for adult
hedenediaby wihes excretion jaf chitin,» Ded: meso, and meta-
; , eu notal dises, e, anterior trach-
passing thus into the condition of the — eal anastomosis; f, _ first
este : z % muscular belt ; g, transverse
Euny uid pupa. Dufour thought DIA i Seem pein othe dorsal
there is no larval stage in this Insect, tracheal tube ; 7, sex-organ ;
“7 08 : Beg. : py eet = k, Malpighian tube; J, ter-
but it is quite clear from later researches inal part of intestine ; m,
that he was wrong, and that a larval terminal chamber of tracheal
eae oy pei nck uate Py tube ; 2, stigmatic fossa; 0,
stage of a peculiar kind, but in some re- terminal part of intestine ; p,
spects resembling that of the Eumyiid — anus; g, anal disc ; 7, ventral
tracheal tube; s, stomach ;
Muscidae, occurs. The larva has no t, nervous system; w, discs
true head, but the anterior part.of the for the three pairs of legs of
: ; : : the imago ; v, ventral pouch ;
body is invaginated, and the most w, pharynx ; ~, suctorial lip.
anterior part again protrudes in the
invagination, so that two little passages appear on section (Fig.
246); the upper one leads to the stomach, which is of very
large size. The tracheal system is peculiar; it is metapneustic,
there being neither anterior nor lateral spiracles. Pratt says
that there is at first a single pair of terminal spiracles, and sub-
sequently three pairs, hence he considers that the terminal part
of the body corresponds to three segments. This is however
probably a mistaken view; it appears more probable that the so-
called three pairs of stigmata really correspond with the complex

1 Abh, Ges. Halle, iv. 1858, p. 145. 2 Arch. Naturgesch. lix. 1. 1898, p. 151.

520 DIPTERA CHAP.

condition of the stigmata in the later instars of certain- other
Dipterous larvae. The MJelophagus-larva is nourished by secre-
tion from certain glands of the mother-fly; this is swallowed
and the stomach is greatly distended by this milky fluid.
Probably it was this condition that induced Dufour to suppose
the larva to be only an embryo.

Some of the Hippoboscidae that lve on birds take to the
wing with great readiness, and it is probable that these bird-
parasites will prove more numerous than is at present suspected.

We may here notice an animal recently described by Dr.
Adensamer and called Ascodipteron.' He treats it as the female
imago of a Pupiparous Dipteron. It was found buried in the
skin of the wing of a bat of the genus Phyllorhina, in the
Dutch East Indies, only one individual bemg known. It is
entirely unsegmented, and externally without head. If Dr.
Adensamer should prove to be correct in his surmise the creature
can scarcely be inferior in interest to the Strepsiptera.

Fam. 41. Braulidae.—This consists only of a minute Insect
that lives on bees. The antennae are somewhat lke those of
the sheep-tick, though
they are not so com-
pletely concealed in
the cavities in which
they are inserted. Ac-
cording to Miiggen-
burg? a ptilinum
exists, and he is also
of opinion that al-
though the parts of
the mouth differ very
much from those of
Hippoboscidae — they
are essentially similar. Lucas says that Braula specially affects
the thorax of the bee: Miiggenburg, that it is fond of the
queen-bee because of the exposed membranes between the body-
segments that exist in that sex. Whether this Insect is truly
Pupiparous is unknown, though Boise states that a pupa is
deposited in the cell of the bee by the side of the young larva of

Fie, 247.—Braula coeca. x

1.§B. Ak, Wien. ev. 1896, Abtheil. i. p. 400.
2 Arch. Naturges. lviil. i. 1892, p. 287.

VIL BRAULIDAE—STREBLIDAE—NYCTERIBIIDAE 521

the bee, and appears as the perfect Insect in about twenty-one
days. Miiggenburg suggests that Braula may be oviparous, as he
has never found a larvain the abdomen. Packard says that on the
day the larva hatches from the egg it sheds its skin and turns to an
oval puparium of a dark brown colour. The Insect is frequently
though inappropriately called bee-louse ; notwithstanding its name
it is not quite blind, though the eyes are very imperfect.

Fam. 42. Streblidae.— Winged ; possessing halteres; the
head small, narrow and free. These very rare Diptera are
altogether problematic. According to Kolenati the larvae live in
bats’ excrement and the perfect Insects on the bats.’ If the
former statement be correct the Insects can scarcely prove to be
Pupipara. The wing-nervuration is, in the figures of the
Russian author, quite different from that of Hippoboscidae. The
Streblidae have been associated by some entomologists with
Nycteribiidae, and by Williston with Hippoboscidae.

Family 43. Nycteribiidae.—The species of this family are

frees yy) GS
Dotan

ae
TATTINTT NE

We APA NNN

Fic. 248.—Nycteribia sp., from Xantharpyia straminea. Aden. A, Upper surface of
female, with head in the position of repose ; B, under surface of male. x 42.

found on bats; they are apparently rare, and we have been able
to examine only one species. The form is very peculiar, the
! Horae Soc. ent. Ross. ii. 1863, p. 90.

522 DIPTERA—APHANIPTERA CHAP.

Insects looking as if the upper were the under surface. They
are wingless, with a narrow head, which reposes on the back of the
thorax. The prothorax appears to be seated on the dorsum of the
mesothorax. According to Miiggenburg there is no trace of a
ptilinum. A brief note on the metamorphosis’ by Baron Osten
Sacken indicates that the mature larva differs from that of

Mra. 249.—Anterior part of the body of Nycteribia sp., found on Xantharpyia straminea
by Colonel Yerbury at Aden. A, Upper surface of female, with head extended ;
B, under surface of male, with head extended ; C, claws of a foot.

Melophagus in the arrangement of the stigmata; they appear to
be dorsal instead of terminal. There are apparently no characters
of sufficient importance to justify the association of these
Insects with the other divisions of Pupipara; the sole ground
for this connection being the. supposed nature of the life-history
of the larva.

Sub-Order Aphaniptera or Siphonaptera (//cas)

Fam. Pulicidae.— Wingless, with the body laterally compressed,
so that the transverse diameter is small, the vertical one great. The
head indistinctly separated from the body, small, with short thick
antennae placed in depressions somewhat behind and above the un-
Jaceted eyes. These are always minute, and sometimes wanting.—

1 Tr. ent. Soc. London, 1881, p. 360.

VII APHANIPTERA——FLEAS 523

We all know that the Flea is so flat, or compressed sideways, that it
does not mind the most severe
squeeze. This condition is
almost peculiar to it; a great
flattening of the body is com-
mon in Insects—as is seen in
another annoying Insect, the
bed-bug—but the compression,
in the flea, is in the reverse
direction. In other respects the Fie. 250.— Hystrichopsylla talpae. Lritain.
(After Ritsema. )
external anatomy of the flea
shows several peculiarities, the morphological import of which has
not yet been elucidated. The head is of very
peculiar shape, small, with the antennae placed
in an unusual position; the clypeus is said to
be entirely absent, the front legs are articulated
in such a manner that they have a large addi-
tional basal piece—called by some anatomists
the ischium—and in consequence appear to be
placed far forwards, looking as if they were
attached to the head; the meso- and meta-
thorax have certain flaps that have been con-
sidered to be homologues of wings; and the
maxillary palpi are attached to the head in
: such a way that they appear to play the part
F | | of the antennae of other Insects (Fig. 250),
/ mi. tp and were actually considered to be the antennae
by Linnaeus, as well as others; the mouth-
Fic. 251. — Mouth- * 00
parts of a flea, Ver. Parts themselves are differently constructed
mipsylla alakurt from those of any other Insects.!. The maxillae
6. H. Unpaired : o
pricking organ; and labium are considered to be not only present,
oe peers but well developed, the former possessing palpi
Mv. maxilla; Mxp. Moderately well developed, while the labial
ee palps are very large and of highly peculiar
form, being imperfectly transversely jointed
and acting as sheaths; the mandibles are present in the form of

' The best general description of the external anatomy of the flea is to be found
in Taschenberg, Die Fléhe, 1880. The morphology is better elucidated, though still
incompletely, in Wagner’s valuable ‘‘ Aphanipterologische Studien,” Horae Soc. ent.
Ross. xxiil. 1889, pp. 199-260, 5 plates, and op. cit. xxxi. 1897, pp. 555-594, 3
plates. Cf. also N. C. Rothschild, Nov. Zool. v. 1898, pp. 533-544, 3 plates.

524 APHANIPTERA CHAP.

a pair of elongate, slender organs, with serrated edges; and there
is an unpaired, elongate pricking-organ, thought by some to be a
‘nypopharynx, and by others a labrum. The antennae are of
unusual form, consisting of two basal joints, and, loosely con-
nected therewith, a terminal mass of diverse form and more or
less distinctly, though irregularly, segmented. The full number
of ten stigmata exists, Wagner giving three thoracic, with seven
abdominal, placed on segments 2-8 of the abde-
at men; but Packard thinks the supposed meta-
= thoracic stigma is really the first abdominal.
Fleas undergo a very complete metamorphosis ;
the larvae are wormlike, resembling those of
y Mycetophilid Diptera (Fig. 252). The egg of
the cat’s flea is deposited among the fur of the
animal, but (unlike the eggs of other parasites)
apparently is not fastened to the hair, for the
egos fall freely to the ground from infested
animals; the young larva when hatched bears
on the head a curious structure for breaking
the egg-shell. It has the mouth-parts of a
mandibulate Insect and is peripneustic, having
ten pairs of stigmata. It subsequently becomes
of less elongate form. Flea-larvae are able to
nourish themselves on almost any kind of refuse
animal matter, Laboulbene having reared them
. on the sweepings of apartments; they may

Fic. 252.—Larva of : p
Pulex serraticeps, Perhaps sometimes feed on blood; at any rate
Haeca See the contents of the alimentary canal appear red
Kiinckel.) through the transparent integuments. When
full grown the larva makes a cocoon, and
frequently covers it with pieces of dust. The perfect flea
appears in a week or two thereafter; the pupa has the members
free. The food of the larvae of fleas has been much discussed
and a variety of statements made on the subject. It has been
stated that the mother-flea after being gorged with blood carries
some of it to the young, but Kiinckel has shown that there is
very little foundation for this tale. Enormous numbers of fleas
are sometimes found in uninhabited apartments to which animals
have previously had access, and these fleas will attack in numbers
and with great eagerness any unfortunate person who may eiite

go

VII FLEAS 52

Ut

the apartment. The cat-flea can pass through its growth and
metamorphosis with excessive rapidity, the entire development of
a generation in favourable conditions extending but little beyond
a fortnight.’

About a hundred kinds of fleas are known, all of which live
on mammals or birds. Hystrichopsylla talpae (Fig. 250) is one
of the largest, it occurs on the Mole. It was found by Ritsema
in the nests of Bombus subterraneus (and was described under
the name of Pulex obtusiceps). As these nests are known to be
harried by Voles, and as this flea has also been found on Field-
mice,.it is probable that the parasites are carried to the nests
by the Voles. The species that chiefly infests man is Pulex
irritans, an Insect that is nearly cosmopolitan, though arid desert
regions are apparently unsuitable to it. Pulex avium occurs on
a great variety of birds. P. serraticeps infests the dog and the
cat, as well as a variety of other Mammals. It is a common
opinion that each species of Mammal has its own peculiar flea,
but this is far from correct. Fleas pass readily from one species
of animal to another; the writer formerly possessed a cat that
was a most determined and successful hunter of rabbits, and she
frequently returned from her excursions swarming with fleas that
she had become infested with when in the rabbits’ burrows; her
ears were on some occasions very sore from the flea-bites. Some
of the fleas of other animals undoubtedly bite man. There
appears, however, to be much difference in the lability of different
individuals of our own species to the bites of fleas. Sarcopsylla
penetrans differs in habits from other fleas, as the female
buries the anterior parts of her body in the flesh of man or
other Vertebrates, and the abdomen then becomes enormously
enlarged and distended and undergoes a series of changes that
are of much interest.” While in this position the Insect dis-
charges a number of eggs. This species multiples sufficiently
to become a serious pest in certain regions, the body of one man
having been known to be affording hospitality to 300 of these
fleas. Sarcopsylla penetrans 1s known as the Sand-fiea, or
chigger, and by numerous other names. Originally a native of
tropical America it has been carried to other parts of the world.
Another Sarcopsylla, S. gallinacea, attaches itself to the eyelids

1 Howard, Bull. Dep. Agric. Ent. N.S. No. 4, 1896.
* Schimkewitsch, Zool. Anz. vii. 1884, p. 673.

526 APHANIPTERA—THYSANOPTERA CHAP.

of the domestic fowl in Ceylon, and an allied form, Rhynchopsylla
pulex, fastens itself to the eyelids and other parts of the body of
birds and bats in South America. In Turkestan Vermipsylla
alakurt attacks cattle—ox, horse, camel, sheep—fastening itself to
the body of the animal after the fashion of a tick. Retaining
this position all through the winter, it becomes distended some-
what after the manner of the Sand-flea, though it never forms
a spherical body. The parts of the mouth in this Insect (Fig. 251)
are unusually long, correlative with the thickness of the skins of
the animals on which it lives. Grassi considers that the dog’s
flea, Pulex serraticeps, acts as the intermediate host of Taenia.

Great difference of opinion has for long prevailed as to
whether fleas should be treated as a Sub-Order of Diptera or as
a separate Order of Insects. Wagner and Kiinckel, who have
recently discussed the question, think they may pass as aberrant
Diptera, while Packard,’ the last writer on the subject, prefers
to consider them a separate Order more closely allied to Diptera
than to any other Insects. Although widely known as Aphan-
iptera, several writers call them Siphonaptera, because Latreille
proposed that name for them some years before Kirby called
them Aphaniptera. Meinert considers them a separate Order
and calls it Suctoria, a most unfortunate name.

Order VIII. Thysanoptera.

Small Insects, with a palpigerous mouth placed on the under side of
the head and apposed to the sternwm so as to be concealed.
With four slender wings, fringed with long hairs on one or
both margins, or with rudiments of wings, or entirely apterous.
Tarsi of one or two joints, terminated by a vesicular structure.
The young resemble the adult in general form, but there is a
pupal stadium in which the Insect 1s quiescent and takes no
food.

The tiny Insects called Thrips are extremely abundant and
may often be found in profusion in flowers. Their size is only
from =, to + of an inch in length; those of the latter magni-
tude are in fact giant species, and so far as we know at present
are found only in Australia (Fig. 253). As regards the extent

1 P. Boston Soc. xxvi. 1894, pp. 312-355.

VIt THRIPS 527

of the Order it would appear that Thysanoptera are insignificant,
as less than 150 species are known. Thrips have been, how-
ever, very much neglected by entomologists, so it will not be a
matter for surprise if there should prove to be several thousand
species. These Insects

present several points >_> Z :

of . interest ; their :

mouth - organs ATE aaah |
=) Oy AN . hi)

: : ss MQ. SR Se a
unique in structure; By» Weer KI
besides this, they ex- L prrmnm
hibit so many points eS
of dissimilarity from = we

other Insects that it 7 a" ie
is impossible to treat
them as subdivisions
of any other Order.
They have, however,
been considered by
some to be aberrant
Pseudoneuroptera (cf.
Vol. V.), while others
have associated them
with Hemiptera. Both
3rauer and Packard
have treated® Thysa-
noptera as a separate
Order, and there can
be no doubt that this Fie. 253.—Idolothrips spectrum. Australia.
is correct. Thysano-
ptera have recently been monographed by Uzel in a work that 1s,
unfortunately for most of us, in the Bohemian language.’
The antennae are never very long, and are 6 to 9-jointed.
The head varies much, being sometimes elongate and tubular,
but sometimes short ; it has, however, always the peculiarity that
the antennae are placed quite on its front part, and that the
mouth appears to be absent, owing to its parts being thrust
against the under side of the thorax and concealed. Their most
remarkable peculiarity is that some of them are asymmetrical :
Uzel looks on the peculiar structure, the “ Mundstachel,’ m, m

1 Monographie der Ordnungy Thysanoptera, Koniggratz, 4to, 1895.

528 THYSANOPTERA

CHAP.

(Fig. 254) found on the left side of the body, as probably an enor-
mous development of the epipharynx. Previous to the appearance
of Uzel’s work, Garman had, however,
correctly described the structure of the
mouth;' he puts a different interpretation
on the parts; he points out that the
mandibles (7), so-called by Uzel, are at-
tached to the maxillae, and he considers
that they are really jointed, and that
they are lobes thereof; while the Mund-
stachel or piercer is, he considers, the
left mandible ; the corresponding struc-
ture of the other side being nearly
entirely absent. He points out that
the labrum and endocranium are also
asymmetrical. We think Garman’s
view a reasonable one, and may re-
mark that dissimilarity of the man-
dibles of the two sides is usual in

Fia. 254.—Face (with base of

a a et ee -

the antennae) of Aeolothrips
fasciata. (After Uzel.) a,
Labrum ; 6, maxilla with its
palp (c); 6l, terminal part of
vertex near attachment of
mouth-parts ; d, membrane
between maxilla and mentum ;
é, mentum ending in a point
near f/f; g, membrane of
attachment of the labial palp
h; 1, ligula ; 7, 7 the bristle-
like mandibles ; 4, the thicker
base of mandible ; /, chitin-
ous lever; mm, mouth-spine,
with its thick basal part n,
and o, its connection with
the forehead, 7, 7; p, for-
amen of muscle; s and ¢,
points of infolding of vertex ;
u, a prolongation of the
gena.

Insects, and that the mandibles may
be hollow for sucking, as is shown by
the larvae of Hemerobiides. There
are usually three ocelli, but they are
absent in the entirely apterous forms.
The wings appear to "spring from
the dorsal surface of the body, not
from the sides; the anterior pair is
always quite separated from the pos-
terior ; the wings are always slender,
sometimes very slender; in other re-
spects they exhibit considerable variety ;
sometimes the front pair are different
in colour and consistence. from the
other pair. The abdomen has ten seg-

ments, the last of which is often tubular in form. The peculiar
vesicular structures by which the feet are terminated are, during
movement, alternately distended and emptied, and have two hooks
or claws on the sides. The stigmata are extremely peculiar,
there being four pairs, the first being the mesothoracic, 2nd

1 Bull. Essex Inst. xxii. 1890, p. 24; also Amer. Natural. xxx. 1896, p. 591.

vil THRIPS 529

metathoracic, 3rd on the second abdominal segment, 4th on the
eighth abdominal segment.’ There are four Malpighian tubes,
and two or three pairs of salivary glands. The dorsal vessel
is said to be a short sack placed in the 7th and 8th abdominal
segments. The abdominal gangha of the ventral chain are con-
centrated in a single mass, placed in, or close to, the thorax ; the
thorax has two other approximated ganglia, as well as an anterior
one that appears to be the infra-oesophageal.

The metamorphosis is also pecuhar; the larva does not differ
greatly in appearance from the adult, and has similar mouth-
organs and food-habits. The wings are developed outside the
body at the sides, and appear first, according to Heeger, after
the third moult. The nymph-condition is like that of a pupa
inasmuch as no nourishment is taken, and the parts of the body
are enclosed in a skin: in some species there is power of movement
to a slight degree, but other species are quite motionless. In
some cases the body is entirely bright red, though subsequently
there is no trace of this colour. Jordan distinguishes two
nymphal periods, the first of which he calls the pronymphal; in
it the Insect appears to be in a condition intermediate between
that of the larva and that of the true nymph; the old cuticle
being retained, though the hypodermis is detached from it
and forms a fresh cuticle beneath it. This condition, as Jordan
remarks, seems parallel to that of the male Coccid, and ap-
proaches closely to complete metamorphosis; indeed the only
characters by which the two can be distinguished appear to
be (1) that the young has not a special form; (2) that the
wings are developed outside the body.

Thrips take their food, it is believed, in the same manner
as Aphidae, by suction; but the details of the process are not by
any means certain, and examination of the stomach is said to have
resulted in finding pollen therein. Walsh thought that Thy-
sanoptera pierce and suck Aphidae. An _ elaborate inquiry
by Osborn ® failed to elicit satisfactory confirmation of Walsh’s
idea, though Riley and Pergande support it to some extent ;
Osborn concludes that the ordinary food is not drawn directly
from sap, but consists of exudation or pollen, the tissues

1 Jordan in an interesting paper, Zeitschr. wiss. Zool. xlvii. 1888, p. 573, says
that in the division ‘‘ Terebrantia”’ there are only three pairs of stigmata.

2 Insect Life, i. 1888, p. 138.

VOL. VI 2

ad
onl

530 THYSANOPTERA CHAP.

of the plant being pierced only when a supply of food from
the usual sources falls short. Members of this family have
been reputed as being very injurious to cultivated plants, especially
‘to cereals, and it is said that as a result the harvests in Europe
have been seriously diminished. Several species may take part:
in the attacks. These appear to be directed chiefly against
the inflorescence. Lindeman thought that Limothrips denticornis
(= Thrips secalina), and Anthothrips aculeata (= Phloeothrips
frumentarius), were the most destructive species in an attack of
Thrips on corn that he investigated in Russia. Uzel suggests
that injuries due to other causes are sometimes ascribed to Thrips."
In hot-houses these Insects are well known, and sometimes
occasion considerable damage to foliage. The German _horti-
culturalists call them black-fly, in distinction from Aphidae or
ereen-fly. Some Thysanoptera live under bark, and even in
fungi, and in Australia they form galls on the leaves of trees.
This observation is due to Mr Froggatt, and is confirmed by
specimens he sent to the writer. Vesicular bodies in the leaves
of Acacia saligna were traversed on one side by a longitudinal
sht, and on a section being made, nothing but Thrips, in various
stages of growth, was found inside them. A second kind of gall,
forming masses of considerable size on the twigs of Callistemon,
is said by Mr Froggatt to be also due to Thrips, as is a third
kind on Bursaria spinosa. It is curious that Thrips’ galls have
not been observed in other parts of the world.

Thysanoptera are devoured by small bugs of the genus
Triphleps, as well as by beetles; a small Acarid attacks them by
fixing itself to the body of the Thrips. Nematode worms and their
eges were found by Uzel in the body-cavity. He found no less
than 200 Nematodes in one Thrips, and noticed that they had
entirely destroyed the ovaries. Woodpeckers, according to him,
tear off the bark of trees and eat the Thysanoptera that are
concealed thereunder, though one would have surmised that
these minute Insects are too small to be game for such birds.
They have, it appears, no special protection, except that one
species (a larva of Phloeothrips sp.) is said to emit a protective
fluid.

Parthenogenesis seems to be frequent amongst Thysanoptera,

1 See Lindemann, Buil. Soc. Moscow, 1xii. 1886, No. 2, p. 296, and Uzel, Mon.
1895, pp. 397, 398.

VII THRIPS 531

and is found in concurrence with diversity as to winged and wing-
less females of the same species, so as to have given rise to the idea
that the phenomena in this respect are parallel with those that
are more widely known as occurring in Aphidae. Under certain
circumstances few or no males are produced (one of the cir-
cumstances, according to Jordan, being season of the year), and
the females continue the species parthenogenetically. In other
cases, though males are produced they are in very small numbers.
Some species of Thysanoptera are never winged; in others the
individuals are winged or wingless according to sex. But there
are other cases in which the female is usually wingless, and
is exceptionally winged. The winged specimens in this case
are, it is thought, of special use in disseminating the species.
Jordan has suggested that these phenomena may be of a
recular nature, but Uzel does not take this view. Another
condition may be mentioned, in which the species is usually wing-
less, but winged individuals of the male as well as of the female
sex oceasionally appear. Zhrips lini apparently makes regular
migrations, feeding at one time underground on the roots
of flax, and then changing to a life in the open air on other
plants.

Numerous forms of Thysanoptera, belonging to both of the
great divisions of the Order, have been found fossil in Europe
and North America, but all are confined to deposits of the Tertiary
epoch.

Of the 135 species known to Uzel, 117 are European; they
are divided into two Sub-Orders. 1, Terebrantia, in which the
females are provided with an external toothed ovipositor, of two
valves; 2, Tubulifera, in which there is no ovipositor, and the
extremity of the body is tubular in both sexes. The British
species are about 50 in number, and were described by
Haliday about 60 years ago; of late they have been very
little studied.

The name Physopoda or Physapoda is used for this Order,
instead of Thysanoptera, by several naturalists.

1 Entomological Magazine, ili. 1836, p. 489, and iv. 1837, p. 144.

CHAPTER VIII

HEMIPTERA——_OR BUGS

Order IX. Hemiptera.

Mouth consisting of a proboscis or mobile beak (usually concealed
by being bent under the body), appearing as a transversly-
jointed vod or grooved sheath, in which are enclosed long
slender setae (like horse-hairs). Wings (nearly always) four ;
the anterior frequently more horny than the posterior pair,
and folding flat on the back, their apical portions usually
more membranous than the base (Heteroptera); or the four
wings may cover the abdomen in a vroof-like manner, and
those of the anterior pair may not have the basal and apical
parts of different consistences (Homoptera); sometimes all
four of the wings are transparent. The young resembles the
adult in general form ; the wings are developed outside the
body, by growth, at the moults, of the sides of the hinder por-
tions of the meso- and meta-notum ; the metanotal prolonga-
tions being more or less concealed by the mesonotal.

THE Hemiptera or Bugs are perhaps more widely known as
Rhynchota. In deciding whether an Insect belongs to this
Order the student will do well to examine in the first place
the beak, treating the wings as subordinate in importance, their
condition being much more variable than that of the beak. The
above definition includes no reference to the degraded Anoplura
or Lice. These are separately dealt with on p. 599; they are
absolutely wingless, and have an unjointed proboscis not placed
beneath the body, the greater part of it being usually withdrawn
inside the body of the Insect.

The Hemiptera are without exception sucking Insects, and

q
x

CHAP. VIII BUGS S16)

the mouth-organs of the individual are of one form throughout
its life. In this latter fact, coupled with another, that the
young are not definitely different in form from the adult,
Bugs differ widely from all other Insects with sucking-mouth.
They agree with the Orthoptera in the facts that the mouth does
not change its structure during the individual life, and that the
development of the individual is gradual, its form, as a rule,
changing but little. In respect of the structure of the mouth,
Orthoptera and Hemiptera are the most different of all the
Orders. Hence, Hemiptera is really the most isolated of all the
Orders of Insects. We shall subsequently see that, like Orthoptera,

Fic. 255.—Husthenes
pratti (Pentato-
midae). China. A,
Nymph : a, case of
anterior, b, of pos-
terior wing ; ¢, ori-
fices of stink-
glands; B, the
adult Insect.

the Order appeared in the Palaeozoic epoch. Although a very
extensive Order, Hemiptera have for some incomprehensible
reason never been favourite objects of study. Sixty years ago
Dufour pointed out that they were the most neglected of all the
ereat Orders.of Insects, and this is still true; our acquaintance
with their lfe-histories and morphology especially being very -
limited.

There is probably no Order of Insects that is so directly con-
nected with the welfare of the human race as the Hemiptera ;
indeed, if anything were to exterminate the enemies of Hemiptera,
we ourselves should probably be starved in the course of a few
months. _The operations of Hemiptera, however, to a large ex-
tent escape observation, as their mouth-setae make merely pricks
that do not attract notice in plants: hence, it is probable that

534 HEMIPTERA CHAP.

injuries really due to Hemiptera are frequently attributed to
other causes.

In the course of the following brief sketch of the anatomy
and development of Hemiptera, we shall frequently have to use
the terms Heteroptera and Homoptera; we may therefore here
mention that there are two great divisions of Hemiptera having
but little connection, and known by the above names: the
members of these two Sub-Orders may in most cases be dis-
tinguished by the condition of the wings, as mentioned in the
definition at the commencement of this chapter.

External structure.—The mouth-parts consist of an anterior
or upper and a posterior or lower enwrapping part, and of the
organs proper, Which are four hair-hke bodies, dilated at their
bases and resting on a complex chitinous framework. The
lower part forms by far the larger portion of the sheath and is of
very diverse lengths, and from one to four-jointed: it is as it were
an enwrapping organ, and a groove may be seen running along it,
in addition to the evident cross-segmentation. The upper
covering part is much smaller, and only fills a gap at the base
of the sheath ; it can readily be lifted so as to disclose the setae ;
these latter organs are fine, flexible, closely connected, rods, four
in number, though often seeming to be only three, owing to the
intimate union of the components of one of the two pairs; at
their base the setae become broader, and are closely connected
with some of the loops of the chitinous framework that is con-
tained within the head. Sometimes the setae are much longer
than the sheath; they ‘are capable of protrusion. Although
varying considerably in minor points, such as the lengths of the
sheath and setae, and the number of cross-joints of the sheath,
these structures are so far as 1s known constant throughout the
Order. There are no palpi, and the only additions exceptionally
present are a pair of small plates that im certain forms (aquatic
family Belostomidae) lie on the front of the proboscis near the
tip, overlapping, in fact, the last of the cross-articulations.

Simple as is this system of trophi its morphology is uncertain,
and has given rise to much difference of interpretation. It may
be granted that the two portions of the sheath are respectively
upper lip, and labium; but as to the other parts wide difference
of opinion still prevails. On the whole the view most generally
accepted, to the effect that the inner pair of the setae correspond

VII STRUCTURE 535

in a broad sense with maxillae of mandibulate Insects, and the
outer pair with mandibles, is probably correct. Mecznikow, who
studied the embryology,! supports this view for Heteroptera,
but he says (4c. p. 462), that in Homoptera the parts of the
embryo corresponding with rudimentary maxillae and mandibles
disappear, and that the setae are subsequently produced from
peculiar special bodies that are at first of a retort-shaped form ;
the neck of the retort becoming afterwards more elongate to form
the seta; also that in the Heteropterous genus Gerris the
embryology in general resembles that of Homoptera, but the

Fic. 256.—Mouth - parts of

cat Hemiptera. (After
Pigs Wedde.) A, Section of

the head and proboscis of
Pyrrhocoris apterus : dr,
gland ; 7.g, infra - oeso-
phageal ganglion; /b,
labium ; 77, labrum; m,
muscles ; m1, muscle (de-
pressor of labium); m7,
muscle of syringe; ph,
pharynx; s, setae; sg,
supra - oesophageal gang-
lion; sp. dr, salivary
gland; spr, syringe: B,
transverse section of pro-
boscis of Pentatoma ruji-
pes, at third joint of
sheath: m, m, muscles ;
md, mandibular seta ; ma,
maxillary setae ; 7, nerve ;
p, the sheath or labium ;
tr, trachea.

KW!

development of the setae is like that of other Heteroptera (¢.c. p.
478). This discontinuity in the development of the Homopterous
mouth has since been refuted by Witlaczil,? who found that the
retort-shaped bodies really arise from the primary segmental
appendages after they have sunk into the head. We are there-
fore justified in concluding that the mouth-parts are at first
similarly developed in all Hemiptera, and that this development
is of a very peculiar nature.

Smith is convinced that there are no traces of mandibular
structure in any Hemiptera.” On the other hand, numerous
entomologists have supposed they could honiologise satisfactorily
various parts of the Hemipterous trophi with special parts of the

1 Zeitschr. wiss. Zool. xvi. 1866, p. 389. 2 Arb. Inst. Wien. iv. 1882, p. 415.
3 Tr. Amer. Phil. Soc. xix. 1896, p. 176.

536 HEMIPTERA CHAP.

maxillae and labium of mandibulate Insects. This point has
recently been discussed by Marlatt’ and by Heymons.” From
the latter we gather that the mode of growth is peculiar by the
extension backwards of some of the sclerites, and their becoming
confounded with parts of the wall of the head. From all this it
appears that at present we cannot correctly go farther than
saying that the trophi of Hemiptera are the appendages of three
head-segments, like those of other Insects. The views of Savigny,
Léon,’ and others to the effect that labial palpi, and even other
parts of the labium of Mandibulata can be satisfactorily identified
are not confirmed by Heymons.

Underneath the pharynx, in the head, there is a peculiar
structure for which we have as yet no English term. It was
apparently discovered by Landois and Paul Mayer,’ and has been
called “ Wanzenspritze,’ which we translate as syringe. It may be
briefly described as a chamber, into which the salivary ducts open,
prolonged in front to the neighbourhood of the grooves of the
setae in the rostrum; behind, it is connected with muscles ;
it has no direct connection with the pharynx, and though it was
formerly supposed to be an organ of suction, it seems more prob-
able that it is of the nature of a force-pump, to propel the pro-
ducts of some of the bug’s glands towards the tips of the setae.

The rostrum being extended from its position of repose, the
tip of the sheath is brought into contact with the object to be
pierced, the surface of which is probably examined by means of
sensitive hairs at the extremity of the sheath; these therefore
functionally replace to some extent the palpi of other Insects.
As a rule the sheath does not penetrate (though there is reason
for believing that in various of the animal-feeding bugs it does
so), but the setae are brought into action for piercing the skin
of the plant; they are extremely sharp, and the outer pair are
usually barbed, so that when once introduced a hold is easily
maintained. This being established it is thought that the salivary
pump comes into play, and that a fluid is injected into the object
pierced so as to give rise to irritation or congestion, and thus
keep up a supply of fluid at the point operated on: this fluid
extends along the grooved setae by capillary attraction, and the
1 P. ent. soc. Washington, iii. 1895, p. 241. 2 Ent. Nachr. xxii. 1896, p. 178.

3 Zool. Anz. 1897, No. 527, p. 73.
4 Arch. Anat. Physiol. 1874, p. 318, and 1875, p. 309.

vu STRUCTURE Sey

rapidity of the current is increased by a pumping action of the
pharynx, and possibly by movements of the setae themselves.
Though the setae are often extremely elongate—sometimes several
times the length of the body—they are nearly always slender, and
there is no reason to suppose that a perfect, or air-tight, tube is
formed; hence it is probable that capillary attraction is really
the chief agent in the ingestion of the fluid. The slight diversity
of structure of the Hemipterous trophi is in very striking con-
trast with what we find in mandibulate Insects, and in the less
purely suctorial Insects, such as Diptera and some divisions of
Hymenoptera. Schiodte im com-
menting on this has suggested that
it is probably due to the small variety
of actions the rostrum is put to.!

The head exhibits great variety
of form; in the Homoptera the
front part is deflexed and inflexed,
so that it is placed on the under
surface, and its anterior margin
is directed backwards; it is often
peculiarly inflated ; in the Lantern-
flies or Fulgoridae (Fig. 282) toan f{
incomprehensible extent. In the great
Water-bugs, Belostomidae, there ison |
the under surface a deep pocket for
each antenna, beautifully adapted to f
the shape of the curious.;-.ormed ap-
pendage (Fig. 279). The prothorax is
always very distinct, frequently large,
and in many of the Heteroptera
(Fig, 207), as well as in the Homop- M6, 27 5s, Ratan
terous family, Membracidae (Fig. (Antennae absent in the specimen
283), assumes the most extraordinary Eepeceeater)
shapes. Both meso- and meta-thorax are well developed. The
former is remarkable for the great size of the scutellum ; in
some cases (Plataspides, Scutellerides) this forms a large process,

1 For the structure and development of the Hemipterous trophi, see Mayer, Arch.
Anat. Physiol. 1874 and 1875 ; Mecznikow, Zeitschr. wiss. Zool. xvi. 1866, p. 389 ;
Geise, Arch. Naturgesch. xlix. 1, 1883, p. 315; Wedde, op. cit. li. 1, 1885, p. 113 ;
Mark, Arch. mikr. Anat. xiii. 1877, p. 31: Smith, Zr. Amer. Phil. Soc. xix. 1896,
p- 176.

538 HEMIPTERA CHAP.

that entirely covers and conceals the alar organs, so that the
Insect has all the appearance of being apterous. The exact com-
position of the abdomen has not been satisfactorily determined,
opinions varying as to whether the segments are nine, ten, or
eleven in number. The difficulty of determining the point
is due to two facts: viz. the extreme modification of the terminal
segments in connection with the genital appendages, and the
prominence of the extremity of the alimentary canal. If this
terminal projection is to be treated as a segment, it would appear
that eleven segments exist, at any rate in some cases; as the
writer has counted ten distinct segments in a young Coreid bug,
in addition to the terminal tube. This tube in some of the male
Heteroptera is very subject to curious modifications, and has
been called the rectal cauda. Verhoeff considers that ten seg-
ments were invariably present in the females examined by him
in various families of Heteroptera and Homoptera.’ In Aphidae
(a division of Homoptera), Balbiani considers there are eleven
abdominal segments present; but he treats as a segment a pro-
jection, called the cauda, situate over the anus; this structure does
not appear to be homologous with the rectal cauda we have just
mentioned. In Coccidae the number of abdominal segments is
apparently reduced. Schiodte states” that the older authorities
are correct in respect of the stigmata; there are, he says, in
Heteroptera invariably ten pairs; one for each thoracic segment ;
and seven abdominal, placed on the ventral face of the pleural
fold of the abdomen. In some cases there are additional orifices
on the external surface that have been taken for stigmata, though
they are really orifices of odoriferous glands; these openings may
exist on the metasterna or on the dorsal surface of the abdomen.
The lateral margins of the abdomen are frequently greatly de-
veloped in Heteroptera, and are called “connexivum ;” the upper
and lower surfaces of the body meeting together far within the
marginal outline. Dr Anton Dohrn many years ago® called atten-
tion to the extremely remarkable structure of the terminal segments
in many male Hemiptera; and the subject has been subsequently
very umpertectly treated by the present writer and other ento-
mologists, but it has never received the attention it deserves.

Ent. Nachr. xix. 1893, p. 369.
2 Naturhist. Tidskr. (8) vi. 1896; translated in Ann. N. Hist. (4), vi. 1870,
10 2S. 3 Ent. Zeit. Stettin, xxvii. 1866, p. 321.

Vil STRUCTURE 539

In the females of numerous Heteroptera and Homoptera
(Capsidae, Cicadidae, etc.) there is a well-developed ovipositor, that
serves both as a cutting instrument to make slits in the stems
of plants, and as a director to introduce the eggs therein. Verhoeff
considers that it always consists of two pairs of processes (though
one pair may be very small), one from the eighth abdominal
segment, the other from the ninth."

The antennae usually have very few joints, often as few as
four or five, their maximum number of about twenty-five being
attained in the males of some Coccidae, this condition being,
however, present in but few of even this family. In Selostoma
(Fig. 279) they assume extremely curious forms, analogous to
what we find in the Coleopterous genus Hydrophilus. In addi-
tion to the compound eyes, there are usually ocelli, either two or
three in number, but wanting in many cases. The usual number
of joints of the tarsi is three, but in Coccidae there is only one
joint.

The wings (Fig. 258) exhibit much diversity. The anterior
pair usually differ greatly from the pos-
terior ; they are called elytra, hemi-elytra
or tegmina. This difference in the two
pairs is the rule in the first of the great
divisions of the Order, and the name
Heteroptera is derived from the fact. In
this Sub-Order the front wings close over
the back, and are more or less horny, the
apical part being, however, membranous.
Systematists make use of the wings for
the purpose of classification in Heterop-
tera, and distinguish the following parts,

“ clavus,’ “corium,’ “membrane,” the B
corium being the larger horny division, Fie. 258.—Alar organs of a
; 7 7 Capsid bug (Capsus lani-

avus ying ne3 = g
the clavus the part lying next the scu sR Mivtron :
tellum and frequently very sharply dis- clavus; 3, corium; ©,
: ‘ : cuneus ; D, membrane ;
: : an 0; :
tinguished from the corium; the mem aie cell Gf tha with.

brane is the apical part. The outer brane ; B, hind-wing.

or costal part of the wing is also often
sharply delimited, and is called the “embolium;” in the great
family Capsidae and a few others, the outer apical part of the

1 Ent. Nachr. xix. 1893, p. 375.

540 HEMIPTERA CHAP.

corium is differentiated from the rest of the surface, and is termed
the “cuneus.” In Plataspides, one of the divisions in which
the alar organs are entirely covered by the scutellum, they are
modified in a very remarkable manner. In the Homoptera the
divisions named above do not exist, and the wings in repose are
placed in a different position, as stated in our definition of the
Order. It is said to be very difficult to homologise the wing-
nervures of Hemiptera, and nothing appears to be known as to
the mode of their development.

The alar organs in Hemiptera exhibit a very frequent form
of variation within the limits of the same species; this has not
yet been elucidated.’ In some cases in the Heteroptera nearly
all the individuals of a generation may have the wings aborted ;
sometimes this occurs as a local variation. In Aphidae the
occurrence of winged and wingless individuals is very common,
and has even become an important factor in their extraordinary
life cycles. (See Chermes, etc., subsequently.)

Internal anatomy.—The alimentary canal presents consider-
able diversity and some remarkable features. There is a slender
tube-like oesophagus and a large crop. It is difficult to assign
any of the parts posterior to this to the divisions usual in
other Insects, and it is said that the distinction of parts histo-
logically is as vague as it is anatomically. In the Heteroptera
the Malpighian tubes open into two (or one) vesicular dilatations
seated immediately in front of the short rectum: between this
point and the crop there may be a very elongate, slender portion
with one or more dilatations, these parts apparently replacing
the true or chylific stomach. There is no gizzard. In the
Homoptera the relations of the divisions of the alimentary canal
are even more puzzling; the canal is elongated and forms coils,
and these are connected with tissues and tunics so as to make
their dissection extremely difficult. List says that there are
great differences in the alimentary canal among the members
of the one family Coccidae. There are usually four Malpighian
tubes, but in Coccidae there is only one pair, and in Aphidae
none. The excretory cells of these tubes are in Hemiptera of
remarkably large size. There is a large development of salivary
glands, at least two pairs existing. There can be little doubt
that some of their products are used for purposes of injection, as

' On this subject, see Reuter, Ann. Soc. ent. France (5) v. 1875, p. 225.

VIII STRUCTURE 541

already described, though Kiinckel came to the conclusion that
the saliva when placed in living plants is totally innocuous.’

The ganglia of the nervous system are all concentrated in
the thorax and head. In some cases (in various Homoptera) the
infra-oesophageal ganglion is placed at a distance from the supra-
oesophageal ganglion, and may even be united with the thoracic
mass of ganglia (Orthezia, etc.); in this case the chitinous frame-
work of the mouth-parts is interposed between the supra-.and
the infra-oesophageal ganglia. In Pentatoma all the three gan-
glionic masses are brought into close proximity, but in Nepa the
thoracic mass of ganglia and the infra-oesophageal ganglion are
widely separated.

The ovarian tubes vary greatly in number: according to List
in Orthezia cataphracta the number differs considerably in dif-
ferent individuals, and even in the two ovaries of the same
individual, the number being usually ten. The testes are not
placed in a common tunic, though they are frequently approxi-
mated or even contiguous.”

The smell of bugs is notorious. In many species it is not
unpleasant, though as a rule it is decidedly offensive. It is a
remarkable fact that the structures connected with the production
of this odour are different in many cases in the young and in
the adult. The odour emitted by the latter proceeds from a sac
seated at the base of the abdomen, and opening exteriorly by
means of an orifice on each side of the metasternum; while in
the young there are two glands situated more dorsally and a
little more backwards, and opening on two of the dorsal plates
of the abdomen (Fig. 255, A). In the young the dorsum of the
abdomen, where the stink-glands open, is exposed, but this part
in the adult is covered by the wings. The odorific apparatus is
specially characteristic of Heteroptera, and Kiinckel states that
there is so much variety that generic and even specific characters
might be drawn from conditions of the stink-glands. As a rule
they are most constantly present in the plant-feeding forms; in
some essentially carnivorous forms (Reduviidae, Nepidae, Noto-

1 Ann. Soc. ent. France (4) vii. 1867, p. 45.

* The chief work on the internal anatomy of Hemiptera is still Dufour’s Ze-
cherches anatomiques et physiologiques sur les Hémiptéres, Mem. Savans Etrungers,
Paris, iv. 1833, p. 129.

3 Kunckel, Ann. Soc. ent. France (4) vil. 1867, p, 45, and C.&. Ac. Paris, cxx.
1895, p. 1002.

542 HEMIPTERA CHAP.

nectidae) they are entirely absent. The offensive matter emitted
by Notonecta. is of a different nature, and is probably anal in
origin.

Metamorphosis or postembryonic development.—In the
language of the systematists of metamorphosis, Hemiptera are
said to be Homomorpha Paurometabola—that is, the young
differ but little from the adult. According to Brauer’s general-
isations they are Menorhynchous, Oligonephrous, Pterygogenea,
i.e. they have a sucking mouth that does not change during life,
few Malpighian tubes, and are winged in the adult state. It is
generally admitted that the Homoptera do not completely agree
with Heteroptera in respect of the metamorphosis, it being
more marked in the former, and in Coccidae attaining (as we
shall mention when discussing that family) nearly if not quite
the condition of complete metamorphosis of a pecuhar kind.
Unfortunately we are in almost complete ignorance as to the
details of the life-histories and development of Heteroptera, so
that we can form no generalised opinion as to what the post-
embryonic development really is in them, but there are grounds
for supposing that considerable changes take place, and that
these are chiefly concentrated on the last ecdysis. The young of
some bugs bear but httle resemblance to the adult; the magnifi-
cently-coloured species of Husthenes (Fig. 255), before they attain
the adult condition are flat, colourless objects, almost as thin as a
playing-card ; it is well known that the extraordinary structures
that cover and conceal the body in Plataspides, Scutellerides,
Membracides, ete., are developed almost entirely at the last moult :
it is not so well known that some of these changes occur with much
rapidity. A very interesting account of the processes of colour-
change, as occurring in Poecilocapsus lineatus at the last ecdysis,
Ras been given by Lintner,’ and from this it appears that the
characteristic coloration of the imago is entirely developed in
the course of about two hours, forming a parallel in this respect
with Odonata. When we come to deal with Aphidae we shall
describe the most complex examples of cycles of generations that
exist in the whole of the animal kingdom.

Fossil Hemiptera.—Hemiptera are believed to have existed
in the Palaeozoic epoch, but the fossils are not numerous, and
opinions differ concerning them. Hugereon hockingi, a Per-—

1 In Slingerland’s Cornell Univ. Bull. No. 58, 1893, p. 222.

VI FOSSIL FORMS—CLASSIFICATION 543

mian fossil, was formerly supposed to be a Homopterous Insect,
but it is very anomalous, and its claim to a position in Hemip-
tera is denied by Brauer,’ who considers it to be Orthopterous.
It is now generally recognised that this fossil requires complete
reconsideration. Another Permian fossil, Fulgorina, is admitted
to be Homopterous by Scudder, Brauer and Brongniart. Scudder
thinks the Carboniferous Phthanocoris was an Archaic Hetero-
pterous Insect, and if correct this would demonstrate that both of
the two great Sub-Orders of Hemiptera existed in Palaeozoic times.
Brauer, however, is inclined to refer this fossil to Homoptera, and
Brongniart > speaks of it as being without doubt a Fulgorid.
Dictyocicada, Rhipidioptera and Meganostoma, from the Carboni-
ferous shales of Commentry, have also been referred to Fulgoridae
by Brongniart, but the evidence of their alliance with this group
is far from satisfactory. In the Secondary epoch numerous
Hemiptera existed, and are referred to several of the existing
families. They come chiefly from the Oolite. In the Eocene of
the Isle of Wight a fossil has been discovered that is referred to
the existing Homopterous genus 7'riecphora.

We are not entitled to conclude more from these facts than
that Homoptera probably appeared before Heteroptera, and date
back as far as the Carboniferous epoch.

Classification and families—No complete Laney of
Hemiptera exists, but one by M. Severin is in course of publica-
tion. It is probable that there are about 18,000 species at
present described, two-thirds of this number being Heteroptera.
In Britain we have about 430 species of Heteroptera and 600
of Homoptera. The classification of the Order is not in a very
advanced condition. The following table exhibits the views of
Schiédte® in a modified form :—

Front of head not touching the coxae. I, HETEROPTERA.
Front of head much inflexed so as to be in contact with the coxae.
Il. Homoptera.

Sub-Order I. HrTEROPTERA.

Posterior coxae nearly globose, partiy embedded in cavities, and having
a rotatory movement. Mostly terrestrial forms. 1. Trochalopoda.

Posterior coxae not globose, larger, and not embedded ; their articulation
with sternum almost hinge-like. Posterior aspect of hind femur usually

1 SB. Ak. Wien. xci. 1 Abth., 1885, p. 275.
° Les Insectes fossiles, etc., 1894, p. 452. 3 Ann. Nat. Hist. (4) vi. 1870, p. 225.

544 HEMIPTERA CHAP.

more or less modified for the reception of the tibia when closed on it: mostly
aquatic forms, 2. Pagiopoda.

Division 1, TROCHALOPODA.
This division includes the majority of the families of Heteroptera—viz.
the whole of the terrestrial families except Saldidae, and it also includes
Nepidae, a family of water-bugs.

Division 2. PaGropopa.

This includes the six purely aquatic families of Heteroptera, except
Nepidae, which appear to have very little connection with the other aquatic
bugs. The only terrestrial Insects included in the family are the Saldidae ;
in these the femora are not modified as they are in the aquatic forms.
Hemiptera that live on the surface of water, not in the water, are classed
with the terrestrial species. With these exceptions this arrangement agrees
with that of Gymnocerata and Cryptocerata as usually adopted,! and
therefore followed in the following pages. Schiddte’s characters, moreover,
do not divide his two divisions at all sharply.

Sub-Order IT. Homoprera.
Tarsi usually three-jointed : . Series Trimera.
s », two-jointed », Dimera.
‘ » Of one joint ; : », Monomera.

The classification of Homoptera is in a most unsatisfactory state ;2 no two
authors are agreed as to the families to be adopted in the series 'Trimera.
We have recognised only five—viz. Cicadidae, Fulgoridae, Membracidae,
Cercopidae, and Jassidae. The Dimera consists of Psyllidae, Aphidae,
Aleurodidae ; and the Monomera of Coccidae only. It is usual to associate the
Dimera and Monomera together under the name of either Phytophthires or
Sternorhyncha, but no satisfactory definition can be given of these larger
groups, though it seems probable that the families of which they are com-
posed are natural and distinct.

Sub-Order I. HETEROPTERA.

Series 1. Gymnocerata.

The majority of the terrestrial families of Heteroptera form
the series Gymnocerata, in which the antennae are conspicuous,
and can be moved about freely in front of the head, while in

1 A table of the families is given by Ashmead, but does not work out quite
satisfactorily, Hntom. Americana, iv. 1888, p. 65; a brief table of the characters
of the British families is given by Saunders, Hemiptera-Heteroptera of the British
Islands, 1892, p. 12.

2? Those who wish to see tables of the families are referred to Ashmead, Joc.
cit. ; to Pascoe, Ann. Nat. Hist. (5) ix. 1882, p. 424; to Stal’s Hemiptera Afri-
cana, vol. iv. 1866; and for the families found in Britain to Edwards, Hemiptera-
Homoptera of the British Islands. For a discussion in Danish on the value of the
characters used, cf. Hansen, Ent. Tidskr. xi. 1890, pp. 19-76.

Sad saan

musitciliaiines

VIII HETEROPTERA—PENTATOMIDAE 545

Cryptocerata they are hidden. The series Gymnocerata includes
all the terrestrial Heteroptera, and the two families, Hebridae
and Hydrometridae, which live on the surface of the water or in
very damp places; while Cryptocerata includes all the forms that
live under water.

Fam. 1. Pentatomidae.—Scutellum very large, at least half
as long us the abdomen, often covering the whole of the after-body
and alar appendages. Antennae often five-jointed. Proboscis-
sheath four-jointed. Ocelli two. Hach tarsal claw with an
appendage.-—This, the largest and most important family of the
Heteroptera, includes upwards of 4000 species, and an immense
variety of forms. It is divided into no less than fourteen sub-
families. The species of one of these, Plataspides, are remarkable
for their short, broad forms, and the peculiar condition of the
alar organs, which are so completely
concealed by the great scutellum that
it is difficult to believe the Insects are
not entirely apterous. The head is usually
inconspicuous though broad, but in a
few forms it is armed with horns.
Though this sub-family includes upwards
of 200 species, and is very widely dis-
tributed in the Old World, it has no
representatives in America. The Scutel-
lerides also have the body covered by
the scutellum, but their organs of flight
are less peculiar than they are in the |
Plataspides ;< the: Insects: of this’ sub- ~ Ps/209:— Phdeen corticata.

South America.
family are highly remarkable on account
of their varied and frequently vivid coloration; some of them
are metallic, and the colour of their integuments differs greatly
in some cases, according to whether the specimen is wet or
dry ; hence the appearance after death is often very different
from that of the living specimen. These Insects are extremely
numerous in species. The sub-family Phloeides (Fig. 259), on
the contrary, includes only three or four South American
species: they have no resemblance at all to other Pentatomidae ;
they are flat, about an inch long, and look like scales of bark,
in this respect agreeing with Ledra and some other Homoptera.
The South American sub-family Cyrtocorides (Fig. 260) is of
VOL. VI 2N

546 HEMIPTERA-HETEROPTERA CHAP.

equally small extent ; the species are of strange irregular shapes,
for which we can find no reason. The Tessaratomides includes
many of the largest Hemiptera-Hetero-
ptera, some of its members attaining
two inches in length.

The great family Pentatomidae, con-
taining about 400 species, is represented
in Britain by about 36 native species,
the most interesting of which are perhaps
those of the genus Acanthosoma. De
Geer noticed long ago that the female of
A, grisewm exhibits great solicitude for
its young, and his statement has since
Fie. 260. Meprssomia ne been confirmed by Mr. Parfitt and the

strosus. South America. Rey. J. Hellins, who found that the

ares mother not only protects the eggs but
also the young, and that for a considerable time after hatching.’

Very little is known as to the life-histories of Pentatomidae.
In some cases the young are very different in appearance
from the adults. The peculiar great scutellum is not developed
till the mature condition is reached. But httle attention has
been given to the habits of Pentatomidae; it is generally con-
sidered that they draw their nutriment from plants; the
American Huthyrhynchus floridanus has, however, been noticed
to suck the honey-bee, and we think it probable that a good
many Pentatomids will be found to attack Insects.

The term Pentatomidae as applied to this family is of modern
origin: in most books the equivalent group is called Scutata, or
Seutati,and the term Pentatomidae is restricted in these works
to the sub-family called Pentatomides in the system we adopt.

Fam. 2. Coreidae.—Scutellum not reaching to the middle of
the body ; proboscis-sheath four-jointed ; ocelli present ; antennae
yenerally elongate and four-jointed, inserted on the upper parts
of the sides of the head; femora not knobbed at the tip.—The
members of this great family are easily recognised by the above
characters ; formerly it was called Supericornia in connection with
the characteristic position of the antennae. About 1500 species
are known, and they are arranged in no less than twenty-nine
sub-families. Many of them are Insects of large size, and they

1 Ent. Mag. vii. 1870, p. 53.

i a, a ma

eee or)

VIII HETEROPTERA—COREIDAE 547

frequently have a conspicuous disc, or dilatation, on one of the
joints of the antennae. Another very curious and, as yet, inex-
plicable peculiarity very commonly met with among them, is
that the hind legs may be of great size and deformed ; either the
femora or the tibiae, or both, being very much distorted or
armed with projections. Brilliant colour is here comparatively
rare, the general tone being indefinite tints of browns, greys, or
smoky colours. The South American genus Holymenia (Copius

Fic. 261.—Diactor bilineatus. Fra. 262.—Phyllomorpha laciniata, carry-
South America. x 4. ing some of its eggs. Spain.

of older authors) consists of slender forms, having the elytra
transparent even on the basal part like Homoptera; this and
some other peculiarities give the species of this genus a certain
resemblance to Insects of other Orders; Westwood says that
Diateina holymenoides (Diptera) greatly resembles a bug of the
genus Holymenia. The tropical American genus Diactor consists
of a few species of elegant colour having the hind legs very
peculiarly shaped, the tibiae being flattened and expanded in a
sail-like manner, and ornamented with agreeable colours different

548 HEMIPTERA-HETEROPTERA CHAP.

from those on the rest of the body; they are made more con-
spicuous by the femora being remarkably long and thin; it is
probable that they are used as ornaments. The sub-family
PhyHNomorphides consists of about a dozen species, and is found
in several of the western parts of the Eastern hemisphere, one
species, P. laciniata, occurring in Southern Europe. This Insect is
of very delicate texture, and the sides of the body are directed
upwards and deeply divided so that a sort of basin is formed, of
which the dorsum of the body is the floor; the Insect is very
spinose, and is thus enabled to carry its eggs, the spines helping
to retain them in position on the back. It is said to be the male
that thus carries the eggs. This species is able to stridulate,
and when doing so vibrates its antennae with excessive rapidity.
We have only about a score of species of Coreidae in Britain,
and none of the remarkable forms of the family are among them.

Fam. 3. Berytidae.—Very slender Insects with the first
joint of the antennae and the femora thickened at the tips.—
This small family was not distinguished from Coreidae by the
older authors. It consists of about fifty species, eight of which
are found in Britain.

Fam. 4. Lygaeidae.— Zhe characters are the same as those
mentioned for Coreidae, except as regards the insertion of the
antennae ; the upper surface or face of the head is not so flat, but
is transversely convex, so that seen in profile the antennae appear
to be inserted well down on the sides of the head—The name
Infericornia was formerly apphed to these Insects. They
are on the average of smaller size than the members of the
Coreidae or Pentatomidae, and are much less conspicuous in colour
and form; a good many of the larger Lygaeids are, however,
variegate with black, yellow, and red. The family is very numerous
in species, about 1400 being known; they are arranged in
thirteen sub-families; we have about sixty species in Britain,
nearly all small. Hvemocoris lives, when immature, in the nests
of the wood-ant, according to Wasmann. The family includes
some notorious Insect-pests. The Chinch-bug, blissus leuco-
pterus, commits very serious ravages on corn and grasses in North
America. The Cotton-stainer, Dysdercus suturellus is also very
injurious to cotton in certain parts of the New World: its growth
has been described by Riley,’ who thinks a dye valuable for

1 Insect Life, i. 1889, p. 234.

VIII LYGAEIDAE—PYRRHOCORIDAE—TINGIDAE 549

commercial purposes might be procured from the Insect. This
bug has recently developed the habit of sucking oranges, and has
thus become injurious in Florida, as the fruit readily decays
after it has been punctured by these Insects. The phenomenon
of “micropterism ” is exhibited by numerous Lygaeids, as well as
by Pyrrhocoridae.

Fam. 5. Pyrrhocoridae.—Distinguished from Lygaeidae only
by the absence of ocelli, and not recognised as a distinct family
by all Hemipterists. About 300 species are included. Our
only British member is the notorious Pyrrhocoris apterus ; it 1s,
however, very rare in this country, though it abounds on the
Continent, and has been the object of investigation by embryo-
logists and others. It displays in a most marked manner the
curious dimorphism as to the alar organs that is so common in
certain divisions of Hemiptera; the elytra and wings being some-
timés normally developed, while in other cases the wings are
entirely absent, and the horny, basal part of the elytra only is
present. In some localities, and in some years, only the micro-
pterous form is found, while on other occasions there may be a
large percentage of the macropterous form. The abundance of
this Insect has enabled the French chemist Physahx to obtain
an amount of its colouring matter sufficient for analysis; as
the result he procured a substance, insoluble in water, very
closely allied to carotine.’ The Oriental Insect Lohita grandis is
one of the most remarkable of Bugs, the male of the Sumatran
variety being over two inches in length, having enormously long
antennae, and the abdomen extended to about twice the normal
length, while the other sex is in the usual condition in these
respects. The species is said to be injurious to the cotton-plant
in India.

Fam. 6. Tingidae.—7arsi two-jointed. Elytra more or less
reticulate, consisting of strong, irregular, thick lines forming a frame-
work of cells, the enclosed part of the cell being of different texture
and frequently transparent ; antennae with terminal joint more or
less knob-like, the preceding joint very long ; ocelli wanting ; pro-
notum prolonged behind, covering the scutellum ; front coxae placed at
the back of the thorax.—This 1s the first of a series of families
with only two joints to the feet. These little bugs are very
remarkable objects, and exhibit much variety in their peculiar

1 C.R. Ac. Sct. Paris, exviil. 1894, p. 1282.

550 HEMIPTERA-HETEROPTERA CHAP.

sculpture, which in numerous forms attains a condition of
elegance well worthy of attention. There are nearly 300 species
known, and in Britain we have about a score. The characters
we have given above do not apply to the genus Piesma, though
it is usually placed in
this family; its scutel-
lum is not covered, and
ocelli are present. Al-
though but little is
known as to the nature
of the lives of Tingidae,
yet it was pointed out
long ago by Reéaumur
that a species of the
family (probably C. elavi-
corne, Fig. 263), lives in
Fie. 263. —Copium clavicorne. Europe. deformations of the

(After Riibsaamen.)
flowers of the Labiate
plant now called Teueriwm chamaedrys ; Frauenfeld has more
recently confirmed this observation, and shown that the closely allied
C. teucrii affects the flowers of 7. montanwm in a similar manner.'
Fam. 7. Aradidae.—Very jlat, broad; scutellum exposed,
large or moderate ; abdomen broader than the alar organs, which it
Frequently encases like a broad frame. Front coxae placed in the
middle of the prosternum.—These very flat Insects, of obscure colour,
have frequently very peculiar sculpture. They live under bark, or
on fungi growing from bark,and
are supposed to draw their nut-
riment from the fungi, though
but little is actually known as
to their natural history. The
family is almost cosmopolitan,
and includes about 500 species,
of which five occur in Eng-
land. The small sub-family
Isoderminae consists of a few
species that are placed only
provisionally in Aradidae; they
differ from the normal members by there being no groove on the

Fic. 264.—Aradus orientalis. Siam.

1 Verh. Ges. Wien. iii. 1858, p. 157.

hal Ae $i

EE EEEEeEeaEaeE

VIII ARADIDAE—HEBRIDAE—HYDROMETRIDAE iS Syl

breast, so that the rostrum is free. Of the five species, three
oceur in Chili and Patagonia, two in Tasmania, and one in
Australia.

Fam. 8. Hebridae.—JJinute bugs, of semiaquatic habits,
clothed beneath with a dense, minute, silvery pubescence ; antennae
five-jointed ; legs of not more than average length ; elytra in larger
part membranous.—This small family consists altogether of only
about a dozen species; we have two species of the genus /ebrus
in Britain; they are usually found in very wet moss.

Fam. 9. Hydrometridae.—Morm very diverse; antennae
four-jointed, tarsi two-jointed. Coxae usually widely separated.
Either wingless or with elytra of one texture throughout, having
no membranous part. Under surface with a minute velvet-like
pubescence. In many forms the legs are of great length—Although
of comparatively small extent—scarcely 200 species being at
present known—this family is of great
interest from the habit possessed by its
members of living on the surface of
water. In the case of the notorious
genus Halobates (Fig. 265) the Insects
can even successfully defy the terrors
of Neptune and live on the ocean
many hundreds of miles from land.
There is great variety of form among
Hydrometridae. The European and
British genus JMesovelia is of short
form, and but little dissimilar from
ordinary land-bugs, with which, indeed,
it is connected by means of the genus
Hebrus, already noticed. Aesovelia
represents the sub-family Mesoveliides,
which, though consisting of only four
species, occurs in both hemispheres, and
in the su giaics as well ap in the tem- Fic. 265.— Halobates sobrinus.
perate regions. Our species, M. furcata, Under surface of a female
walks on the surface of the water, the Ta dee D228
movements of its legs and the posi- ;
tion of its coxae being those of land-bugs. Another British
Insect—the highly remarkable Hydrometra stagnorum—is ot
excessively slender form, with long thin legs, by aid of which it

SZ HEMIPTERA-HETEROPTERA CHAP.

walks on the surface-film of water, above which its body is held
well separated. It is easily drowned, and if submerged it has
great difficulty in escaping from the water. This genus repre-
sents the sub-family Hydrometrides, and is apparently almost
cosmopolitan. Velia currens is another common British Insect ;
it loves the eddies and currents of backwaters on burns and
streams, and is very abundant in Scotland. An American ally,
Rhagovelia plumbea, appears to be not uncommon on the surface
of the ocean in the Gulf of Mexico, near the shores. The great
majority of the family belong to the division Gerrides, of which
the curious, long Insects that float so lazily and skim so easily
on the surface of quiet streams are typical. The species of the
genus now called Gerris, but formerly known as Hydrometra are
apparently distributed all over the world; we have ten in
Britain. They have very long legs, and on being alarmed move
away with the greatest ease.

The genus Ha/obates includes at present fifteen species. They
are found on the ocean, where the surface-water is warm, in
various parts of the world. They are destitute of any trace of
alar organs, the meso- and meta-thorax are closely united and
large, while the abdomen is very small, so that the body is of
oval form; the middle legs are thrown so far back that they are
placed immediately over the posterior pair. When the sea is
calm these Insects skim over the surface with rapidity, but
disappear as soon as it becomes agitated. They are believed to
feed on small animals recently deceased; Witlaczil says on
the juices of jelly-fish, The young are frequently met with,
and there can be no doubt that the whole life-cycle may be
passed through by the Insect far away from land. The Italian
ship Vettor Pisani met with a bird’s feather floating on the
ocean off the Galapagos Islands, covered with eggs which proved
to be those of Halobates in an advanced stage of development.
It was formerly believed that the female carries the eggs for
some time after their exclusion, and although this has since
been denied, it is nevertheless an undoubted fact, for it was
observed by Mr. J. J. Walker,’ to whom we are indebted for
a specimen having the eggs still attached to the body, as shown
in Fig 265. Mr. Walker believes the bugs shelter themselves
when the sea is at all rough by keeping at a sufficient distance

1 Ent. Mag. xxix. 1898, p. 227.

ar
y

vu HYDROMETRIDAE 553

below the surface; they can dive with facility, and are gregarious.
They are frequently found close to the shore, and Mr. Walker
has even met with them on land. The stink-glands of other
Hemiptera are said
by Nassonoff to be
replaced in Halobates
by peculiar ventral
glands. An allied
genus, Halobatodes,
was supposed to be
oceanic, but this 1s
not the case, some of
the species having
been found recently in
fresh water in India,
and others in estu-
aries at Port Darwin.
A remarkable allied
form, Hermatobates
haddoni, was recently
discovered by Pro-
fessor Haddon in
Torres Straits. Apart
from the oceanic life, | ]
Halobates is by no \
means the most ex-
traordinary of the
Hydrometridae. The

Javanese Ptilomera

laticaudata repeats Fic, 266.—Rheumatobates bergrothi. x 10.
West Indies. (After Meinert.)

some of its peculi-
arities, and is of larger size, with the sexes very different. The
most remarkable of the family is perhaps the fresh-water genus
Rheumatobates (Fig. 266), in which the males have peculiar
prehensile antennae that look like legs. These curious Insects
inhabit North America and the West Indies.

We may here notice an enigmatic Insect called Hemidiptera
haeckeli by Léon. From the single specimen known it is con-
cluded that the Insect has only one pair of wings, and that they
are attached to the metathorax. It is, however, possible, as

554 HEMIPTERA-HETEROPTERA CHAP.

suggested by Bergroth,’ that the anterior pair have been
detached by some accident.

Fam. 10. Henicocephalidae.—Head swollen behind the

eyes so as to form a sort of globe, on the anterior part of
which the ocelli are placed. Rostrum extremely short. Elytra
rather large, of one consistence throughout ; conspicuously veined.—
There is only one genus; it is very widely distributed, about a
dozen species being known; one of these occurs in the south of
Europe. These curious little bugs appear to. be most nearly
allied to the Reduviidae. According to Westwood and others
they are somewhat gregarious; a Tasmanian species dances in
the air after the fashion of midges or May-flies, and dispenses an
agreeable, musk-like odour.

Fam. 11. Phymatidae.—Front legs of peculiar structure, short
and stout, with long coxae, short thick femora, and tibiae curvate,
povnted ; frequently without tarsii—The Insects of this family are

Fig. 267.—Carcinocoris binghami (Phymatidae). Burma.
J

believed to be predaceous, the structure of the legs being such as
is called raptorial, and one species, Phymata erosa, being known
to capture and suck honey-bees in North America. There are
only about seventy species of Phymatidae known. We have

1 Wien. ent. Zeit. xi. 1892, p. 169.

eer egg

VIII

PHYMATIDAE—REDUVIIDAE 55

Ut

none mm Britain, though there are a few in Southern Europe;
one of these, P. crassipes, extends as far north as Paris. The
distinction of the family from Reduviidae is doubtful There
are a few very rare forms (Fig. 267) in which the front tibia is
articulated to the femur in

such a way that a pair of

pincers is formed : the tarsus
is in this form, as well as
in some other Phymatidae,

absent.

Fam. 12. Reduviidae.
—Head more or less elon-
gate, very movable, eyes placed
much in front of the thorax,
ocelli, when present, behind
the eyes. Proboscis short, or
moderately short, not ex-
tending on to the breast, in
repose curved under the head
so as to form a loop there-

with.

Elytra, when present,

consisting of three divisions.
Tarsi three-jointed—This is

one of

the largest and most

important families of Hemi-

ptera.
species

Upwards of 2000
are already known ;

the habits seem to be chiefly
of a predaceous nature, the
creatures drawing their
nutriment from the animal

rather

than from the vege-

table kingdom, and their
chief: prey being in all

probability other kinds of
Insects.

There is, perhaps,

no family of Insects exhibit-

ing a greater variety of form and colour.

X

SS

=e

Fic. 268.—Ghilianella filiventris. Brazil. A,
the female Insect. B, extremity of the body

The Emesids are amongst

the most delicate of Insects, equalling in this respect the daddy-

1 Monograph of Phymatidae: Handlirsch, Ann. Hofmus. Wien, xii. 1897, p. 127.

556 HEMIPTERA-HETEROPTERA CHAP.

long-leg flies; they are, however, highly predaceous; their front
legs are peculiarly formed for capturing and holding their prey, and
have long coxae, like Mantis, so that these Insects are commonly
mistaken for small or young Mantises, from which their sucking
proboscis at once distinguishes them. This curious starved-look-
ing form of bug reaches its maximum of peculiarity in the South
American genus (hili-
anella (Fig. 268). Ac-
cording to Pascoe the
linear form enables the
young larva to becarried
about by the mother,
the long slender abdo-
men of the larva being
eurled around the
thorax of the parent.
Ploiaria pallida, from
Woodlark Island, is
an Insect of excessive
fragility and elegance,
with the long thin legs
coloured with alternate
patches of black on a
white ground, giving
rise to a very curious
appearance remarkably
analogous to what we
Fic. 269.—Nabis lativentris, young. Cambridge. anes te cea Ore ee
A, Insect seen from above; B, profile. equally delicate daddy-
long-leg flies.

We have three species of Emesides in Britain, but most of our
Reduviidae belong to the sub-family Nabides. These approxi-
mate to ordinary bugs in appearance and characters more than
do-any other of the Reduviidae. One of cur indigenous Nabides
is of great interest from the curious resemblance it has to an ant
(Fig. 269). The likeness is brought about by the sides of the
base of the abdomen being very pallid in colour, except a dark
mark in the middle; this mark is in shape like the pedicel of an
ant. Viewed in profile it is found that on the base of the abdo-
men. there is an elevation like the “scale” in this position in

VIII REDUVIIDAE

ul
ut
N

ants, and that the abdomen is extremely ant-like in form. This
resemblance is quite parallel with that of an Orthopteron to an
ant (see Vol. V. p. 323); the Insect is by no means uncommon,
and it is strange that this curious case
of resemblance should hitherto have
escaped notice. The bug runs about on
plants and flowers, and is frequently in
company with ants, but we do not know
whether it preys on them. Not the

Fic. 270.—Ptilocnemus sidnicus.
Australia. (After Mayr.)

least remarkable of the
facts connected with
this Insect is that the
resemblance is confined
to the earlier instars ;
the adult bug not being
like an ant. We may
here mention that there
: are numerous bugs that
S =a “ik closely resemble ants,
~ and that on the whole
there is reason to be-
lieve that the resem-
bling forms are actually
associated during life,
though we really know
very little as to this last
point.

The little sub-family
Holoptilides, with
twenty-five species, but
Fic. 271.—Myiodocha tipulina. China. widely distributed in

the Eastern hemisphere, is remarkable on account of the feathered

558 HEMIPTERA-HETEROPTERA CHAP.

antennae and legs of its members (Fig. 270). Altogether four-
teen sub-families are recognised, the most extensive one being
Harpactorides, including a great variety of remarkable forms; in
the South American genus Notocyrtus (better known as Saccoderes,
Fig. 257), the prothorax is swollen and covers the body to a
greater or less extent in the fashion of a hood. In Yolinus
and Hulyes the coloration is the most conspicuous system that
could be devised, the sides of the abdomen (connexivum) being
expanded into bright-red lobes on which are placed patches
of polished-black. The most remarkable form of Reduviid
is, perhaps, one from China (Fig. 271) of considerable size,
of great fragility, and greatly resembling, like some Emesides,
a daddy-long-legs fly, though it does not belong to the Emes-
ides. It is an altogether anomalous form. According to
Seitz there is found on the Corcovado in Brazil a Reduviid
that exactly resembles one of the
dark stinging-wasps of the genus
Pepsis, and the bug makes the same
sort of movements as the wasp does,
though these are of a kind quite
different from those of ordinary bugs.”
Although the attacks of Redu-
viidae on animals are usually con-
Fic. 272.—Eggs of Endochus cinga- are a pallor gad ee
lensis. “The eggs are attachea Gefenceless kinds, yet this is by no
to a leaf and to each other bya means invariably the case; there
viscid substance; eggs red, the : 5
cover pale yellow, with the club are in fact numerous species that do
a een tip." —MS. note of not hesitate to attack man himself.
Several species of Reduvius do this
in Southern Europe, and are frequently met with in houses. 2.
personatus is the only species of the genus in England; though
far from common anywhere, it is sometimes found in houses, and
is said to destroy the common bed-bug’; it is able to pass its
whole existence in our habitations, for the young are found as
frequently as the adult, and are usually concealed by a quantity
of dusty matter, or refuse, adhering to the body. This habit of
covering the body with some foreign substance is natural to the
Insect, the young that are found on trees being covered with
matter derived therefrom. Darwin has given us an account of

4

1 Ent. Zeit, Stettin, li. 1890, p. 281.

vill AEPOPHILIDAE—CERATOCOMBIDAE—CIMICIDAE 559

the Benchucha,’ a bug an inch long, which in South America
attacks human beings after the fashion of the common bed-bug.
In this case no ill-effects follow the attack, but in the case of
Conorhinus sanguisuga in Arizona, great pain and inflammation
ensue and may end in the gathering and discharge of pus.

Not the least remarkable of characters of Reduviidae is the
form’ of the eggs of some of the species (Fig. 272, and Vol. V.
Fig. 78, C); the egg bearing a peculiar operculum, the purpose of
which is at present quite mysterious.

Fam. 13. Aépophilidae.—A single species forms this family.
It is of considerable interest, as it is incapable of flight, passing a
large part of its life covered by the sea. Aépophilus bonnairer
is a small Insect with quite short head, without ocelli, and with
the organs of flight represented by a pair of very short elytra,
with rounded hind-margins. It is found on the shores of
Western France, and, as a great rarity, on our own south coast.
It no doubt sucks small soft animals. In the Channel Islands
it occurs in spots where it is nearly always covered by a con-
siderable depth of water.

Fam. 14. Ceratocombidae.—Jinute bugs with ocelli and
elytra. Rostrum free. Head not broad, somewhat prolonged in
front; eyes close to the thorax. Hlytra usually without a dis-
tinetly separated membrane. Tarsi three-jointed—This family
includes at present only a few, minute, fragile bugs, that have
often been classified with Cimicidae or Anthocoridae. We have
only two British species, one of which, Dipsocoris alienus, is
common amongst the damp shingle at the margins of the burns
and waters of Scotland.

Fam. 15. Cimicidae—Ocelli absent; elytra very short and
broad, so that the broad abdomen is left uncovered. Head short
and broad. Rostrum received in a groove beneath the head. Tarsi
ithree-jointed—Although this family .consists of only a dozen
species, it is the most notorious of all the Order, as it includes
the detestable Cimez lectularius or common Bed-bug. This Insect
is now peculiar to the habitations of man, and is said not to
trouble savage races; or rather it is supposed to be present only
when the habitations have a certain degree of comfort and per-
manence. It has no fixed period of the year for its development,
but the generations succeed one another so long as the temperature

1 Naturalist’s Voyage, ed. 1884, p. 330 ; chap. xv.

560 HEMIPTERA-HETEROPTERA CHAP.

is sufficiently elevated; during too cold weather the Insects
merely become stupefied, their hves being as it were interrupted
till warmth returns. It is a favourite food with other Insects,
and is destroyed by cockroaches and ants as well as by Reduvius ;
the small black ant Monomoriwm will, it 1s said, clear a house of the
bed-bug in a few days. Nothing is really known as to the origin
of this Insect; it is now very widely distributed. The other
species of the family frequent birds and bats, and are very
similar to the common bug. The genus to which the bed-bug
belongs is in many works called Acanthia instead of Cimex.
Other authors apply the term Acanthia to Salda, but it is better
to allow the name Acanthia to fall into disuse.

Fam. 16. Anthocoridae.—JJinute bugs, usually with ocelli
and with elytra; the latter occasionally abbreviated, but usually
fully developed, with membranous tip. Head prolonged in the
middle in front much beyond the insertion of the antennae; eyes
not far from the thorax. Rostrum free—These small and obscure
Insects appear to be rather numerous in species, and to be chiefly
connected with woods and forests. Some of the species live
in ants’ nests. We have 27
British species belonging
to 11 genera. About 200
species of the family are
known. The members of
the sub-family Microphy-
sides are remarkable from
the great dissimilarity of
the sexes, for which it is
not possible to assign any
reason.

Fam. 17. Polyctenidae.
—Proboscis - sheath three -
jointed, tarsi four -jointed,
antennae four-jointed. Teq-
f _ mina quite short, of one con-

Fic. 273.—Polyctenes fumarius. Sees ie on ae

(After Westwood.) anomalous species forming

this family are parasites on

bats of the genus J/olossus, and have been found in both the
Eastern and Western hemispheres. Westwood, who first described

VIII POLYCTENIDAE—CAPSIDAE 561

them,' treated them as aberrant Anoplura or Lice, but there do
not appear to be any sufficient grounds for removing these para-
sites from Hemiptera-Heteroptera. The condition of their alar
organs reminds one of what exists in Cimex and Aépophilus, and
the mouth is not known to possess any very peculiar structure.
We have had no opportunity of making a thorough examination
of Polyctenes, and therefore speak with some diftidence.

Fam. 18. Capsidae.— Moderate-sized or small bugs, of delicate
consistence, without ocelli ; the elytra and wings usually large in
proportion to the body, the former with two cells (occasionally

Fic. 274.—Helopeltis sp. East India.

only one) in the membrane. Antennae four-jointed, the second joint
usually very long, the terminal two more slender than the others.
The proboscis not received in a groove. Scutellum exposed, mode-
rately large. Tarsi three-jointed. Female with an ovipositor
capable of exsertion—This family is one of the most extensive of
the Hemiptera; we have about 170 species in Britain, where
they are most abundant in the south. The exotic species have
‘been but little collected. Their colours
are usually delicate rather than vivid,
and are never metallic. They frequent
plants of all kinds, and many of them
skip by the aid of their wings with great
agility in the sunshine. The majority
probably suck the juices of the plants, but
some are known to prey on other Insects.
The species of the Indian genus Helo- ao 46: Ue
peltis (Fig. 274) are remarkable by — egg of a Capsid bug allied
possessing a knobbed spine projecting eS ene,
straight up from the scutellum, making

the individual look as if it were a specimen with a pin through

1 Thesaurus ent. Oxoniensis, 1874, p. 197.
VOL. VI 20

562 HEMIPTERA-HETEROPTERA CHAP.

it: they attack the tea-plant and do considerable damage. They
are known as Mosquito-blight. The egg is of comparatively large
size, and is placed by the bug in the stems of the tea-plant,
but attached to one end of the egg are two long slender threads
that project externally. A similar egg (Fig. 275) and method
of oviposition have been described by Mr. Dudgeon as occurring
in another species of Capsidae, called Moesa-blight, in India.'

Fam. 19. Saldidae.—Head short and broad, with large, pro-
minent eyes. Ocelli present. Proboscis not applied to under surface
of head or breast in repose. Scutellum large, not covered. Elytra
covering the upper surface of the abdomen, formed of three distinct
parts. Tarsi three-jointed—These little bugs run with velocity
over mud in damp places, or live in wet moss; some of them can
jump; they are all of dark or obscure colour. There,are only
three genera: Salda, of which we have numerous British species,
being the principal one.

Series 2. Cryptocerata.

The remaining families of Heteroptera are of aquatic habits,
and form in nearly all works a separate division called Hemiptera
Cryptocerata (or Hydrocorisae, or Hydrocores), distinguished by
the antennae being apparently absent; they are, however, really
present, being situate on the under side of the head, to which
they are closely pressed, or in some cases placed in a pocket in
front of each eye. There are six of these families. Schiddte
is doubtless correct in treating this division as an unnatural one ;
it 1s, however, generally adopted, and is convenient for the pur-
poses of nomenclature and arrangement. :

Fam. 20. Galgulidae or Pelogonidae—/orm short and
broad ; head very broad, with prominent eyes, ocelli present. Hind
legs thin, formed for running—The Insects of this family are
but little known; they are only sub-aquatic in habits, frequent-
ing damp places at the margins of streams and waters. The
presence of ocelli distinguishes them from other water-bugs, with
which indeed the Galguldae appear to be but little related.
There are only about twenty species of the family known. We
possess none in Britain; but one, Pelogonus marginatus, occurs

1 Ind. Mus. Notes, iii. No. 5, 1894, p. 53.

VIII CRYPTOCERATA—NEPIDAE 563

in South Europe. The other members of the family are very
widely scattered over the surface of the earth.

Fam, 21. Nepidae.— Abdomen furnished behind with a long
slender siphon; front legs more or less elongate for capturing
prey, placed quite at the front
edge of the prothorax.—This
family consists of two interest-
ing but very dissimilar genera,
Nepa and Ranatra. Both are
widely distributed over the
earth, and are rather numerous
in species’ We have one
species of each genus in Britain.
Nepa cinerea, the common
“water-scorpion,” 1s one of
the commonest of Insects in
Southern Britain, living con-
cealed in shallow waters when
nearly or quite stagnant.
Ranatra linearis (Fig. 276) is
much less common, and appears
to be getting rarer ; it is not re-
corded from farther north than
Cambridge.

The nature of the respir-
atory arrangements in these
Insects is of considerable in-
terest; the long tube at the
extremity of the body consists
of two parts (as shown in Fig.
276) brought together in the
middle, one from each side.
Lacaze-Duthiers states that the Fie. 276—Ranatra linearis, with the two
processes are elongated pleurae, See ie AU ae see

ge.
but in the young it is tar
from clear that this is the case. However that may be, they
seem to convey air to the true breathing organs, situate inside
the cleft on the apical part of the abdomen itself; but details
as to the way in which transfer of air is effected along this

1 Ferrari, Monograph of Nepa, Ann. Hofmus. Wien, iii. 1888, p. 171.

564 HEMIPTERA-HETEROPTERA CHAP.

very protracted passage are not forthcoming. The develop-
ment in Nepa has been studied to a certain extent. The
apical stigmata are the only pair of the abdominal stigmata
that exist in the imago of Nepa, the other six pairs being
obliterated ; the third, fourth, and fifth, according to Schicdte, in
a very peculiar manner: hence, as Martin says,’ the respiratory
system is metapneustic. In an earlier stage of the life, however,
these six pairs of stigmata exist in functional activity placed in
a groove on the under surface of the body; so that the condition
is that termed peripneustic, and remains so till the final moult,
when the long siphon appears. In the early life there is a
short prolongation from the end of the body in connection with
the pair of grooves alluded to, but it is a single unpaired organ,
and does little therefore to explain the appear-
ance of the siphon, which must, at present, be
considered as being suddenly developed at the
last moult.

The eggs of Nepidae are remarkable objects ;
that of the common water-scorpion bears seven
filaments at one end (Fig. 277); while thet of
Ranatra is more elongate, and bears only two,
very elongate, threads. These eggs are deposited
in the stems of water-plants, being introduced
therein, so that the body of the egg is concealed
while the threads project: those of Ranatra are
placed in stems floating on the water, and in
consequence of the threads the stems look as if
they were infested by some fungus. The struc-
Fig. 277.—Egg of ture and formation of the eggs have been

te Korschelt,) WvVestigated with considerable detail by Kor-
schelt.". He looks on the filaments as pneu-

matic, and considers that they supply a coating of air to the
body of the egg; they consist of a spongy mass encircled by two
layers of egg-shell, both of these latter being peculiar in struc-
ture ; the spongy mass is continuous with a layer of the same
kind of substance placed on the interior of the shell of the body

' Bull. Soc. Philomat, (8) vy. 1893, p. 57. There is some diversity of opinion
as to the respiratory orifices, and some authorities say that thoracic stigmata exist
even in the imago.

* Acta Ac. German. li. 1887, p. 224, and Zeitschr. wiss. Zool. xliii. 1886, p. 537.

Vill NEPIDAE—N AUCORIDAE—BELOSTOMIDAE 565

of the egg. It will be recollected that we have described (p.
562) an egg, apparently of the same nature, deposited by Capsids
in the stems of land plants, so that it is very doubtful whether
the threads are really connected with the aquatic development
of the embryo in Nepidae. But the most interesting feature
connected with these eggs is, according to Korschelt, the mode of
development of the filaments, which is sui generis; the shell of
the egg is developed in the ordinary manner as an exudation or
excretion from epithelial cells; but the shell of the filament is
formed as an intracellular product; a mode of chitin-formation
that appears to be peculiar to this structure. Korschelt remarks
that “it is in the highest degree worthy of attention how by
any process of development through a large number of successive
generations so complex a condition could be established as the
result of adaptation to external conditions; and this becomes
even more interesting when we remember that highly peculiar
special processes and departures from the usual modes of tissue-
formation are necessary to permit the development of this
apparatus.” |

Fam. 22. Naucoridae.—No ocelli, and no terminal process
to the body; front legs inserted on or near the front of the
prosternum. Anterior femora usually broad and flat——The
members of this family are truly aquatic, and swim readily in
the water. The family is small, including about nine genera
and thirty species, but, like many water-Insects, the genera are
widely distributed. We have two in Britain—one of them,
Naucoris, common; the other, Aphelocheirus, rare.

Fam. 23. Belostomidae.—No ocelli, and no long terminal
tube to the body; front legs inserted near the front of the pro-
sternum. Posterior tibiae not spiny ; flattened and provided with
swimming hairs.—Although these Insects have been classified
with Nepidae they have but little relation therewith; on the
other hand, the distinctions from Naucoridae are far less im-
portant. The family includes some of the largest Insects. The
South American Lelostoma grande attains a length of four or four
and a half inches. Notwithstanding their considerable size
Belostomidae exist in very large numbers in some localities, and

frequently destroy young fish by aid of the powerful though

! Korschelt, Acta. t.c. p. 245. Compare the remarks we have made on p. 559 as .
to the peculiarities of eggs of many other Hemiptera.

566 HEMIPTERA-HETEROPTERA CHAP.

short rostrum. They appear to be unable to resist the attrac-
tion of artificial light, and are consequently sometimes destroyed
in large numbers. It has long been known that species of the
genera Diplonychus and Zaitha carry
| their eggs on their backs. There is no
special receptacle for the purpose, but the
eges are kept in their pecuhar position
by means of a cement insoluble in water.
It has been stated by Dimmock that
they are placed in position by means of
a long, flexible ovipositor. Schmidt, how-
ever, found that a specimen of Dzplo-
nychus, bearing eggs and examined by
him, was a male, and he subsequently
found that this was the case with other
Fie. 278.—Zaitha anura, egg-bearing individuals of other species,
carrying eggs on its back. go that the mode in which the eggs are
West Indies. 22)
placed in this position and the object of
so curious a habit, remain uncertain. The species of Belostoma
are highly remarkable on account of the curious and complex
structure of their antennae, in respect of which the nearest
analogy is to be found in the large Coleoptera of the genus
Hydrophilus, A very deep, ear-like pocket, exactly suited
to the form of the antennae,
exists on the under side of the
head; hence in repose no sign
of the pecuhar shape of the
antennae exists. When the
antennae are placed in this ear-
like pocket only the one side
of the basal joints is exposed,
the long processes being re- ;
ceived into the deep pocket. !,272z-Aniama of Eeociona, sp, A
In Hydrophilus the antenna is with antenna, 6, extended ; B, with the
antenna retracted. «a, Side of head ; ¢,
used as an accessory organ of pocket for antenna ; @, position of the eyes.
respiration, and it will be in- The corresponding joints of the antenna
: : are numbered 1, 2, 3, 4 in each figure.
teresting to learn whether this
is also the case in Belostoma. Belostomidae have patches of air-
carrying pubescence, analogous with those of Hydrophilus, on
the under sides of the body, elytra and wings, but we do not

VII BELOSTOMIDAE—NOTONECTIDAE—CORIXIDAE 567

know how they are charged. Another extremely interesting
analogy is found in the manner in which the elytra are locked
to the body; a projection from the thoracic side-pieces, forming
a long pouch, into which a fold on the inner side of the elytra
fits, the two being subsequently locked by the action of some
special projections. This arrangement is similar to that which
exists in the anomalous family of water-beetles “Pelobiidae. In
order to make this mechanism more perfect the side-pieces in
Belostoma form free processes. Martin has informed us that the
young have the metasternal episternum prolonged to form a
lamella that he thinks*may be for respiratory purposes.’ About
twelve genera and upwards of fifty species of Belostomidae are
known. None exist in our isles, but several species extend their
range to Southern Europe. In the waters of the warm regions of
the continents of both the Old and New Worlds they are common
Insects, but as yet they have not been found in Australia.

Fam. 24. Notonectidae.—Prosternum short, so that the legs
are placed near the back part of it as well as near the front ; back
of the head overlapped by the front of the pronotum.—The water-
boatmen are extremely common in our ponds, where they may
be seen rising to the surface and raising the posterior extremity
of the body for breathing. They swim on their backs instead of
in the usual position, and have an elaborate arrangement of long
hairs on the body to assist them to carry about an air-supply.
They are said to be lighter than the water, and to have some
difficulty in keeping away from the surface. Notonecta glauca
is the only British species, but we have a second minute Insect,
Plea minutissima, belonging to the family. It les in the mud at
the bottom of shallow waters, and may sometimes be fished up in
great numbers. It is considered by some authors to form a
distinct family. The oviposition of Motonecta has been observed
by Regimbart ; the eggs are inserted into the stems of aquatic
plants.

Fam. 25. Corixidae—Prosternum short, as in Notonectidae ;
summit of the head free from the thorav—We have numerous
species of the genus Coriza in Britain; and others extremely
similar in appearance occur in various parts of the world. The
head is remarkably free, and capable of great rotation. On
dissection it is found to be attached to the thorax only by a

1 Bull. Mus. Paris, 1896, p. 238.

568 HEMIPTERA CHAP.

narrow area; in this respect it differs widely from NVotonecta,
which possesses an extremely large occipital foramen, and the
head of which possesses but little freedom of movement. The
extremely short proboscis is more or less retractile, and therefore
frequently appears absent. A second British genus consists of a
single species, Sigara minutissima. These Insects, unlike oto-
necta, are quite at home beneath the water, where they scurry
about with extreme rapidity, and occur sometimes in enormous
numbers. In Mexico the eggs of Coriaa americana and of C.
Jemorata are used as food, and are said to be very nice. The
Insects themselves are used as food in both Mexico and Egypt.
The species of this family can make a noise beneath the water by
rubbing the front feet against the proboscis." The males have a
very complex asymmetry of the terminal segments, and in some
Species possess on one side of the dorsal surface a curious asym-
metrical organ consisting of rows of very closely-packed, in-
tensely black, comb-like plates, called by Buchanan White a
strigil. This organ seems to be similar to the peculiar structures
found on the terminal segments of certain species of Scutellerides.

Sub-Order II. Homoptera.”

Fam. 1. Cicadidae.— Head with three ocelli, placed triangularly
on the summit between the compound eyes; antennae consisting
of a short basal joint, surmounted by a hair-like process divided into
about five segments. Front femora more or less thick, armed with
teeth. Peduncle (or basal joints) of antennae without sensitive
organs.—This important family consists chiefly of large Insects,
few being as small as one inch across the expanded wings, while
in some the expanse is as much as seven inches. As a rule
the four wings are transparent and shining, with the nervures
remarkably distinct and dark coloured; but there are numerous
forms where the whole creature, including the wings, is highly
pigmented in a showy manner; frequently in black and yellow.
Cicadas are said to be without any special protection, and to be
destroyed in considerable numbers by birds and other animals.
The body is broad and robust, and is never shaped into the
extravagant forms we meet with in some of the other families of
Homoptera. Cicadidae are almost confined to the warmer regions

1 See Carpenter, Trish Naturalist, iv. 1895, p. 59.
* See remarks on pp. 543, 544.

a

Vul HOMOPTERA—CICADIDAE 569

of the earth, but we have one species, a great rarity, in the ex-
treme south of England; altogether there are about 800 species
known. These Insects are seen above ground—so far as the
life-histories are at present known—only in the perfect condi-
tion, the creatures in their earlier stages being subterranean and
living on roots. As soon as the individual comes out of the
ground it splits open the nymph-skin, and the perfect Cicada
emerges. One species—the North American Cicada septendecim
—is a most notorious Insect owing to its life-cycle of seventeen

Fic. 280.—Cicada septendecim. North America. (After Riley.) A, Larva; B,
nymph ; C, nymph skin after emergence of the imago, D; E, section of twig with
series of eggs ; F, two eggs magnified.

years. It is considered that the individual, after nearly seven-

teen years of underground existence, comes to the surface and

lives for a brief period the life of a noisy Insect. This is the
only Insect at present known having so considerable a longevity.

This fact, and several other peculiarities, have attracted much

attention, so that there is an extensive literature connected with

the seventeen-year Cicada. It has a wide distribution over the

United States, but does not confine its appearance to every

seventeenth year, being found somewhere or other—frequently

in numerous localities—almost every year. The evidence as to
its periodicity has been obtained by taking the locality and other
points into consideration as well as the year of appearance.

570 HEMIPTERA-HOMOPTERA CHAP.

By so doing it has been found possible to establish the existence
of twenty-two broods which are distinguished by consecutive
numeration. This being done, the evidence as to the years
during which Cicadas have appeared in any given locality is
examined, and the result is believed to bear out the view that the
life-cycle of the individual Insect is really one of seventeen years.
According to this view there are, underground, in certain localities
individuals of different ages that will appear on the surface as
mature individuals in different years. Thus in 1885 it was
understood that there were underground in Alabama two broods,
viz. brood xvii. that would appear on the surface in 1894, and
brood iv. that would appear on the surface in 1896. The pre-
dictions made as to the years in which Cicadas would appear in
some given locality are considered to have proved correct. More-
ever, particular entomologists have in certain localities verified
by personal examination the appearance of the Insects for several
consecutive periods of seventeen years. These facts appear fairly
conclusive, but they are much complicated by another point, viz.
that in certain localities the period is one of thirteen, not of
seventeen, years. This is to some extent a question of climate,
the thirteen-year interval being chiefly characteristic of the
Southern States. It is not, however, entirely so, for there
are localities in which the broods have an interval of either
thirteen years or seventeen years. Another fact should be
remembered, viz. that it is admitted that not quite all the
individuals of a particular brood are true to their proper time of
appearance ; In other words, a few specimens may appear pre-
cociously a year or two before their comrades, while some may
lag behind to a considerable extent. It is therefore a matter
for great surprise that, under these circumstances, the broods
should keep distinct at all, for one would suppose that time-
variation of this kind would lead to completely obscuring the
distinctness of the broods. We must also call attention to the
fact that both the seventeen-year and the thirteen-year broods
have a dimorphic form, or sub-species, called C. cassinit which
accompanies the ordinary form, with which it is apparently as a
rule not connected by intermediates.'

1 We must refer those who may wish for further information as to this complex
and difficult question to the writings of the late Professor Riley, especially to
Bulletin No. 8, 1885, U.S. Department of Agriculture, division of entomology ;
and to the more recent report by Marlatt, Bull. Dep. Agric. Ent., N.S. No. 14, 1898.

VIII CICADIDAE WA

Cicadidae are provided with powerful ovipositors. The eggs
of CO. septendecim are deposited in the woody stems of bushes ;
after remaining there a few weeks the young hatch out, drop
to the ground, and, as previously stated, disappear for nearly
seventeen years, nearly the whole of which time is passed
in the larval state, the nymph-condition existing for only
a few days. They feed on the roots of various trees; it has
been said that they are injurious in this way, but other
authorities maintain that they suck only a moist exudation
from the roots. It is very difficult to obtain information as to
their strange, prolonged, subterranean life ; it said that the Insects
sometimes penetrate to a great depth—ten feet, even twenty feet
are mentioned ;—and as great changes may take place on the surface
during their long lives, these Insect Rip Van Winkles some-
times emerge in very strange conditions, and may appear even in
deep cellars. When the pupa comes to the surface it hooks
itself on to the stem of some plant or other object, the skin of
the back splits, and the Cicada emerges. Among the inexplicable
peculiarities of this Insect must be mentioned the fact that when
emerging it sometimes constructs chimneys, or flues, extending
several inches above the surface of the ground. The reason for
this is much disputed; it was said that they are for refuge
against inundations, but this appears to be very doubtful. — Certain
of the broods consist of an almost incalculable number of indi-
viduals, and it is very strange to hear woods, or other localities,
that have been for many years free from these Insects, all at
once resounding with their noisy song. The seventeen-year
Cicada is considered to be doomed to a speedy extinction; the
extension of cultivation and building, and the introduction to
America of the English sparrow, are likely to prove too much
for the Insect.

Although Hemiptera are classified by many among the
Ametabola or Insects without metamorphosis, it is impossible
to deny that the Cicadidae exhibit a considerable amount of
metamorphosis, and they are usually mentioned as exceptional.
The young (Fig. 280, A) is totally unlike the adult in form
and colour, and maintains, to a certain extent, its existence
by the aid of a different set of implements. The larva of the
Cicada is colourless, with an integument of very feeble consistence,
rather large antennae, and a remarkable pair of fossorial legs;

572 HEMIPTERA-HOMOPTERA CHAP.

the wings are totally wanting. The mode of passage from the
larval to the pupal state has not been recorded. The pupa, or nymph,
differs from the larva by its much shorter, compressed form; by
being encased in a remarkably hard shell; and by the antennae
approximating in form to those of the adult. It has short wing-
pads at the sides of the body; the front legs are remarkably
powerful, and the creature is capable of moving about; the imago
escapes from the pupa by the splitting dorsally of the middle of
the thoracic segments. The empty pupa-skin does not shrivel,
but retains its form, and in countries where Cicadas occur, fre-
quently attracts attention by the strange form it presents, being
often placed in a conspicuous position.

Song.—Cicadas are the most noisy of the Insect world; the
shrilling of grasshoppers and even of crickets being insignificant
in comparison with the voice of Cicada. Darwin heard them
in South America when the Beagle was anchored a quarter
of a mile from the shore; and Z'ympanoterpes gigas, from the
same region, 1s said to make a noise equal to the whistle of a
locomotive A curious difference of opinion prevails as to
whether their song is agreeable or not; in some countries they
are kept in cages, while in others they are considered a nuisance.
The Greeks are said to have decided in favour of their per-
formances, the Latins against them. Only the males sing, the
females being completely dumb; this has given rise to a saying
by a Greek poet (so often repeated that it bids fair to become
immortal) “Happy the Cicadas’ lives, for they all have voice-
less wives.” * The writer considers the songs of the European
species he has heard far from unpleasant, but he is an entomologist,
and therefore favourably prepossessed; and he admits that
Riley’s description of the performances of the seventeen-year
Cicada is far from a satisfactory testimonial to the good taste of
that Insect; Riley says, “The general noise, on approaching the
infested woods, is a combination of that of a distant threshing-
machine and a distant frog-pond. That which they make when
disturbed, mimics a nest of young snakes or young birds under
similar circumstances—a sort of scream. They can also produce

' Some entomologists consider that this ‘‘railway-whistle” note is the result
of the combined efforts of several individuals. Cf. Mathew, Ent. Mag. xi. 1875,
joy AB

? It is unnecessary to say that the poet was not Sappho, but one of the baser sex,
named Xenarchus.

‘+ ae etl eo

VIII CICADIDAE 575

a chirp somewhat like that of a cricket and a very loud, shrill
screech prolonged for fifteen or twenty seconds, and gradually
increasing in force and then decreasing.” The object, or use of
the noise is very doubtful; it is said that it attracts the
females to the males. “De gustibus non est disputandum !” per-
haps, however, there may be some tender notes that we fail to
perceive; and it may be that the absence of any definite organs
of hearing reduces the result of a steam-engine whistle to the
equivalent of an agreeable whisper. No special auditory organs
have been detected! as we have already intimated; and certain
naturalists, amongst whom we may mention Giard, think that
the Insects do not hear in our sense of the word, but feel
rhythmical vibrations ; it is also recorded that though very shy
the Insects may be induced to approach any one who will stand
still and clap his hands—in good measure—within the range of
their sensibilities. There is a good deal of support to the idea
that the males sing in rivalry.

Vocal structures.—Although we may not be able to pro-
nounce a final opinion as to the value to the Insect of the sounds,
yet we cannot withhold our admiration from the structures from
which they proceed. These are indeed so complex that they
must be ranked as amongst the most remarkable voice-organs in
the animal kingdom. They are totally different from the
stridulating organs that are found in many other Insects, and are
indeed quite peculiar to the Cicadidae. Some difference of
opinion has existed as to the manner in which the structures
act, but the account given by Carlet, some of whose figures we
reproduce, will, we believe, be found to be essentially correct. The
structures are partly thoracic and partly abdominal. On ex-
amining a male Cicada there will be seen on the under surface
two plates—the opercula—usually meeting in the middle line of
the body and overlapping the base of the abdomen to a greater
or less extent according to the species, sometimes nearly covering
this part of the body ; these are enlargements of the metathoracic
epimera; they can be slightly moved away from the abdomen,
and, as the latter part is capable of a still greater extent of
movement, a wide fissure may be produced, displaying the complex

1 Swinton claims that one of the membranes in the vocal apparatus is an auditory
organ ; if so, the male would be deafened by his own noise, while the females, not
possessing the organ, should not hear the song.

574 HEMIPTERA-HOMOPTERA CHAP.

structures. In order to see the parts it is better to cut away
an operculum ; underneath it three membranes can be seen, an
external, the timbal; an anterior, the folded or soft membrane ;
and a posterior, the mirror. This last is a most beautiful object,
tensely stretched and pellucid, yet reflecting light so as to be
of varied colours; there are also three stigmata, and some
chambers connected with the apparatus. The sound is primarily
produced by the vibrations of the timbal, to which a muscle is
attached ; the other membranes are probably also thrown into a
condition of vibration, and the whole skeleton of the Insect helps
to increase or modify the sound, which is probably also influenced

A

Fic. 281.—Musical apparatus of Cicada plebeia. (After Carlet.) A, Ventral view (Oper-
culum on right side is removed) ; ap, apophysis ; c, cavern ; ¢, trochantin (cheville
of Réaumur) ; exf, part of internal skeleton of abdomen ; mi, specular membrane ;
m.pl, soft or folded membrane; P, base of leg; st, st’, st’, stigmata; ¢, drum
“timbale” ; v, operculum ; 1a, first, 2a, second abdominal segment: B, same seen
laterally, portion of abdominal wall as well as operculum removed ; A, point of
insertion of hind wing; Jes, mesothorax ; sc, scutum of metathorax ; 3a, third

g
abdominal segment ; rest as in A.

by the position of the opercula. The stigmata probably play an
important part by regulating the tension of the air in the
chambers. In the female some of the structures are present in
a rudimentary form, but there are no muscles, and this sex
appears to be really quite voiceless.

Fam. 2. Fulgoridae.—Ocelli two (rarely three, or entirely
obsolete), placed beneath the eyes or near the eyes, usually in
cavities of the cheeks, antennae placed beneath the eyes, very variable
in form; usually of two joints terminated by a very fine hair,
the second joint with a peculiar texture of the surface, owing to the
existence of sensitive structures (Hansen). Form of head very
diverse ; vertex and face forming either a continuous curve, or the
planes of the vertex and face forming an acute angle, or both pro-

ee ee ee

Se “2 el tn mt

= FULGORIDAE ey

longed so as to form a projection or growth that may be monstrous.
Frothorax neither armed nor unusually developed.

This family is of large extent, and includes at present so great
a variety of forms that it is really almost impossible to frame a
definition that will apply to all. The unusual situation of the
ocelli and the peculiar second joint of the antennae must at
present be taken as the best diagnostic characters: occasionally a
third ocellus is present. Some of the Fulgoridae are amongst the
largest Insects, others are quite small, The family includes the
so-called Lantern-flies, in which the front of the head forms a huge
misshapen proboscis that was formerly believed to be luminous.

Fie. 282.—Fulgora candelaria. x1. China.

Many of the species are of brilliant or beautiful coloration. A
great many—and of very different kinds—have the curious power
of excreting large quantities of a white, flocculent wax. This is
exhibited by our little British Insects of the genus Cizius,and in
some of the exotic forms is carried to an extent that becomes a
biological puzzle. The Tropical American genus Phenaz may be
cited as an example; being about an inch long it flies about with
a large mass of this waxy substance twice as long as itself;
indeed, in the Mexican P. auricoma, the waxy processes are four
or five inches long. This wax forms a favourite food of certain
kinds of Lepidoptera, and two or three larvae of a maggot-like
nature may frequently be found concealed in the wax of the live
Fulgorids ; this has been recorded by Westwood as occurring
in India; and Champion has observed it in the New World.

1 P. ent. Soc. London, 1883, p. 20.

576 HEMIPTERA-HOMOPTERA CHAP.

The wax of Fulgorids is used by the Chinese for candles and
other purposes; and this white Insect-wax is said to be much
esteemed in India. Very curious chemical substances have been
obtained from it, but its importance in the economy of the
Insects that produce it is quite obscure. We have about seventy
species of Fulgoridae in Britain. . They belong to the sub-families
Tettigometrides, Issides, Cixiides, and Delphacides, which by
many authors are treated as separate families. The exotic sub-
family Flatides is highly peculiar. In some of its members the
head is very different from that of the ordinary forms, being
narrow, and the vertex and front forming a continuous curve.
Some of these Insects are remarkably like butterflies or moths
(e.g. the African Ityraea nigrocincta and the species of the genus
Pochazia), but the young are totally unlike the old, the posterior
part of the body bearing a large bush of curled, waxy projections,
several times the size of the rest of the body.

Fam. 3. Membracidae.—Prothorax prolonged backwards into
a hood or processes of diverse forms ; antennae inserted in front of
ihe eyes ; ocelli two, placed between the two eyes.—This family 1s of

Fie. 283.—A, B, Heteronotus trinodosus. A, Male seen from above; B, profile of
female ; a, terminal part of pronotum ; 4, terminal part of abdomen: C, front
view of head and pronotum of Cyphonia clavata. Both species from Central
America, (From Biol. Centr. Amer. Rhynch. Homopt. 11.)

large extent but its members are chiefly tropical, and are specially

abundant in America. Although not of large size the Membracidae

are unsurpassed for the variety and grotesqueness of their shapes,
due to the unusual development of the pronotum. We figure two
ot these forms (Fig. 283).' Very little is known about their

1 A considerable variety of these extraordinary creatures are figured in Biol.
Centr. Amer. Rhynch. Homopt. ii.

Vill MEMBRACIDAE——CERCOPIDAE 577

habits and life-histories. We have only two species of the family
in Britain, and these do not afford any ground for supposing that
there are any peculiarities in their lives at all commensurate
with the oddness of the Insect’s structures. Belt has recorded
the fact that in Nicaragua the larvae of certain Homoptera were
assiduously attended by ants for the sake of a sweet juice
excreted by the bugs, but it is by no means clear that these larvae
were really those of Membracidae. In North America Ceresa
bubalus and C. taurina place their eggs in an extremely neat’
manner in the woody twigs of trees. The young have but little
resemblance to the adults, the great thoracic hood being absent,
while on the back there is on each segment a pair of long, sub-
erect processes having fringed, or minutely spiny, margins.’
Fam. 4. Cercopidae.—(celli two (occasionally absent) placed
on the vertex; antennae placed between the eyes. Thorax not
peculiarly formed.—In the characteristic forms of this family
the front of the vertex bears a suture, touched on each side
by one at right angles to it, or converging to it so as to form a
triangle or a sort of embrasure ; the hind tibiae have only one to
three strong spines. The Cercopidae are much less extraordinary
than many of the previously considered families. But some of
them have the habit cf secreting a large quantity of fluid; and
when in the immature stages, certain of them have the art of
emitting the liquid in the form of bubbles which accumulate
round the Insect and conceal it. These accumulations of fluid
are called cuckoo-spits or frog-spits; and the perfect Insects are
known as frog-hoppers, their power of leaping being very great.
The most abundant of the frog-hoppers in our gardens is
Philaenus spumarius, a little Insect of about a quarter of an inch
long, obscurely coloured, with more or less definite pale spots;
it is so variable in colour that it has received scores of names.
Some of the Insects do not use their fluid in this manner, but
eject it in the form of drops, and sometimes cast them to a con-
siderable distance. The phenomena known as weeping-trees are
due to Cercopidae; some of the species make such copious exuda-
tions of this kind that the drops have been compared to a shower
of rain. In Madagascar it is said that Ptyelus goudoti exudes
so much fluid that five or six dozen larvae would about fill a
1 Riley, P. ent. Soc. Washington, iii. 1895, p. 88. For the younger stages of
Membracis foliata, see Tijdschr. Ent. (2) iv. 1869, pl. viii.
VOL. VI 2P
|

578 HEMIPTERA-HOMOPTERA CHAP.

quart vessel in an hour anda half. The frog-spit is considered
by some naturalists to be a protective device; the larvae are,
however, a favourite food with certain. Hymenoptera, which pick
out the larvae from the spits and carry them off to be used as
stores of provision for their larvae. In Ceylon the larva of
Machaerota guttigera constructs tubes fixed to the twigs of the
tulip-tree, and from the tube water is exuded drop by drop.
According to Westwood, this Insect is intermediate between
‘Cercopidae and Membracidae.'

Fam. 5. Jassidae.—Ocelli two, placed just on the front
margin of the head (almost in a line with the front of the eyes
or more to the front) or on the deflexed frons. Hind tibiae
usually with many spines. This vaguely limited family includes
a very large number of small or minute Insects, usually of narrow,
parallel form, and frequently excessively delicate and fragile.
They are often mentioned under- the name of Cicadellinae.
Ashmead distinguishes two families, Bythoscopidae, in which the
ocelli are clearly on the frons or front, and Jassidae, in which
they are on the upper edge thereof. Ulopa, Ledra, and a few
other exceptional forms, are also by many distinguished as
representatives of distinct families. Very little is actually known
as to the life-histories of these small and fragile Insects, but it is
believed that the eggs are usually deposited in the leaves or
stems of plants, and more particularly of grasses. In North
America the development of Deltocephalus inimicus, from hatching
to assumption of the adult form, has been observed by Webster
to occupy about six weeks. As Jassidae are numerous both in
species and individuals it is believed that they consume a con-
siderable part of the vegetation , of pastures. Osborn has
valculated that on an acre of pasture there exist, as a rule, about
one million of these hoppers, and he considers they obtain quite
as large a share of the food as the Vertebrates feeding with
them.

Fam. 6. Psyllidae. — Minute Insects with wings usually
transparent, placed in a roof-like manner over the body ; with three
ocelli, and rather long, thin antennae of eight to ten joints. Tarsi
two-jointed—These small Insects have been studied chiefly in
Europe and North America, very little information having yet
been obtained as to the exotic forms. They are about the

1 Tr, ent. Soc. London, 1886, p. 329.

VIII PSYLLIDAE 579

size of Aphidae, but in form and general appearance remind
ene rather of Cica- |
didae. The wings
are in many cases
even more perfectly
transparent than
they are in many
Cicadidae. They are
sometimes called
springing plant-lice,
as their habit of
jumping distin-
gushes them from
the Aphidae. Low
has called attention
to the remarkable
variation in colour
they present in con-
formity with either
the age of the indi-
vidual, the food-
plant, the climate, Fic. 284.—Psylla succincta. x 15. Europe. (After
and, more particu- Peroea Dart Cte first moult. B, larva after
larly, the season of

the year." Réaumur long since pointed out that at their ecdyses
these Insects go through a remarkable series of changes of colour,
and Low found that this did not take place in the normal
manner in the winter generation that hibernates. This has
been confirmed by Slingerland in North America in the case
of Psylla pyricola, which has been introduced there. He finds
that there are several generations in the year, and that the
hibernating adults differ from the summer adults in size, being
nearly one-third larger; in their much darker colouring; and
especially in the coloration of the front wings.

In the earlier stages, Psyllidae differ greatly in appearance from
the adult forms; the legs and antennae in the newly hatched larvae
are short, and have a less number of joints. In the nymph the
shape is very peculiar,the large wing-pads standing out horizontally

1 Verh. z.-b. Ges. Wien, xxvi. 1876, p. 167.
2 Cornell Univ. Agric. exp. station Bulletin, 44, 1892, and Bull. 108, 1896.

580 HEMIPTERA-HOMOPTERA CHAP.

from the sides of the body, so that the width of the creature is about
as great as the length. The period occupied by the development
apparently varies according to season. Witlaczil, who has given an
account of many details of the anatomy and histology of various
Psyllidae,’ considers that there are four larval stages; Heeger’s
account of Psylla succincta is not quite clear on this point, and
Slingerland indicates a stage more than this, the perfect Insect
being disclosed as the result of a fifth moult ; it is probable that he is
correct. In these earlier stages the body bears long hairs called
wax-hairs; according to Witlaczil in the young larvae of certain
species—T7rioza rhamni, e.g.—these are broad and flat, so as to
make the body appear studded with oval processes; he states
that these hairs change their form during the growth of the
individual. Nothing is more remarkable in Psyllidae than the
amount of matter they secrete or exude from their bodies; in
some species the substance is a “ honey-dew,’ and the nymph
may keep itself covered with a drop of it: in other cases it is
solid, as shown in Réaumuv’s figures of P. bua, where this exuda-
tion forms a string several times longer than the body, and attached
to it. Another form of exudation is a light downy or waxy
matter. Slingerland says that honey-dew was exuded by LP.
pyricola in such quantities that it “literally rained from the trees
upon the vegetation beneath ; in cultivating the orchard the back
of the horse and the harness often became covered with the
sticky substance dropping from the trees. It attracts thousands
of ants, bees, and wasps, which feed upon it.” The writer last
year observed in the New Forest a stunted sloe-bush, about which
a large number of Bombi were busily occupied ; and examination
showed that they were thrusting their proboscides into the curled
and deformed leaves, in which were secreted nymphs of a Psy//la
exuding honey-dew. It must not be assumed that this honey-
dew is the excrement of the Insect; this also is known, and is a
different substance. Those who have tasted it say that the
honey-dew has a clean, good flavour. The source of the honey-
dew is not quite certain, but it seems probable that it comes, like
the solid matter figured by Réaumur, directly from the alimentary
canal, and not from hairs or pores on the body. Psyllidae give
rise to definite formations or galls on certain plants; sometimes
these Psyllid galls are mere changes in form of a limited part, or

1 Zeitschr. wiss. Zool. xlii. 1885, pp. 569-638.

VII PSYLLIDAE—APHIDAE 581

parts, of a leaf, giving rise either to crumpling or to growth of a
porticn in one direction only, so that on one surface of the leaf
a swelling is formed,and on the opposite side a more or less deep
cavity in which the Insect dwells. A formation of this kind on
the leaves of Aegopodium podagraria is described by Thomas ' who
states that the growth is due to the deposition of an egg of the
Psylla, and is independent of the after life of the Insect ; a fungus
—Puecinia aegopodii—torms similar structures on the leaves.
Structures much more definite than this may be the result of the
attacks of Psyllidae; for an example the reader may refer to
Réaumur’s account of Psylla buxi2 In Australia and Tasmania
there are Psyllidae known as Laap or Lerp Insects, the products
of which are called leaf-manna or Lerp, and are used as food.
This manna is a scale produced by the young Insect on the leaves
of Eucalyptus as a covering or protection. The scale is
fastened to the leaf by a hinge, and is somewhat lke the shell
of a cockle. Although the scales are said to be in some cases
objects of great beauty, very little is known about these Australian
Psyllidae, one of which has, however, been referred by Schwarz
to the genus Spondyliaspis, Signoret.? About 160 species of
Psyllidae are known to occur in the Palaearctic region, and about
fifty of them have been found in Britain.*

Fam. 7. Aphidae (Pant-lice or Green-fly.)— Minute Insects ;
as usually met with destitute of wings, though many individuals
have two pairs of transparent wings. Antennae long, or moder-
ately long, three= to seven-jointed ; abdomen frequently with a parr
of tubes (siphons), or short processes on the upper side of the fifth
abdominal segment. Tarsi two-jointed, first joint sometimes
excessively short.—These soft-skinned Insects are frequently called
blight, and are so abundant in temperate climates that a garden,
however small, is sure to afford abundance of specimens during
the warm months of the year. This great abundance is due to
peculiarities in the physiological processes that render these
obscure little animals highly important creatures; the individual
life for several generations is restricted to constant, or at any rate
copious, imbibition of food,accompanied by an almost uninterrupted

1 Zeitschr. Naturw. (2) xii. 1875, p. 438.

* Réeaumur, Mém. ili. 1737, Diviéme Mémoire.

2 P. ent. soc. Washington, iv. 1897, p. 66.

4 For list see Scott, Hnt. Mag. xviii. 1882, p. 253.

582 HEMIPTERA-HOMOPTERA CHAP.

production of young by parthenogenetic females, the young so
produced becoming rapidly (sometimes in the course of eight or
ten days, but more usually in about twenty days) themselves
devoted to a similar process; so that in the comparatively short
period of a few months the progeny resulting from a single
individual is almost innumerable. This remarkable state of
affairs is accompanied by other peculiarities of physiology, with
the result that the life-histories of successive generations become
very diverse, and complex cycles of series of generations differing
more or less from one another are passed through, the species
finally returning to bi-sexual reproduction, and thus inaugurat-
ing another cycle of generations. The surprising nature of
these facts has in the last 150 years caused an immense
amount of discussion, but no satisfactory hght has yet been
thrown on the conditions that really give rise to the exceptional
phenomena. These phenomena are (1) parthenogenesis; (2)
oviparous and viviparous reproduction; (3) the production of
generations of individuals in which the sexes are very unequally
represented, males being frequently entirely absent ; (4) the pro-
duction of individuals differing as to the acquirement of wings,
some remaining entirely apterous, while others go on to the
winged form; (5) the production of individuals of the same sex
with different sexual organs, and distinctions in the very early
(but not the earliest) stages of the formation of the individual ;
(6) differences in the life-habits of successive generations; (7)
differences in the habits of individuals of one generation, giving
rise to the phenomenon of parallel series. All these phenomena
may occur in the case of a single species, though in a very
variable extent.

The simple form of Aphid life may be described as follows :—
egos are laid in the autumn, and hatch in the spring, giving
rise to females of an imperfect character having no wings; these
produce living young parthenogenetically, and this process may be
repeated for a few or for many generations, and there may be in
these generations a greater or less number of winged individuals,
and perhaps a few males.’ After a time when temperature falls,

1 There is some doubt on this point, as the earlier observers seem to have supposed
that a winged individual appearing in a generation chiefly apterous was ipso facto,
a male ; it seems, however, to be certain that perfect winged males appear in some
species in generations producing no perfect sexual females. Speaking generally,

VUl APHIDAE 583

or when the supply of food is less in quantity, or after a period
of deliberate abstention from food, sexual individuals are pro-
duced and fertilised eggs are laid which hatch in the spring, and
the phenomena are repeated. In other cases these phenomena
are added to or rendered more complicated by the intercalated
parthenogenetic generations exhibiting well-marked metamorphosis,
of kinds such as occur in apterous or in winged Insects; while
again the habits of successive generations may differ greatly, the
individuals of some generations dwelling in galls, while those of
other generations live underground on roots.

Parthenogenesis. — Returning to the various kinds of
peculiarities we have enumerated on the preceding page, we may
remark that the phenomena of parthenogenesis have been
thoroughly established as occurring in Aphidae since Bonnet dis-
covered the fact 150 years ago; and though they have not been
investigated in much detail it is known that the parthenogenesis
is usually accompanied by the production of young all of the
female sex. In other cases males are parthenogenetically produced ;
but whether these males come from a female that produces only
that sex is not yet, so far as the writer knows, established. A
note by Lichtenstein’ suggests that usually only one sex is pro-
duced by a parthenogenetic female, but that both sexes are some-
times so produced. There is not at present any species of Aphid
known to be perpetuated by an uninterrupted series of partheno-
genetic generations. It was formerly supposed that there are no
males at all in Chermes, but, as we shall subsequently show, this
was erroneous. It has, however, been observed that a series of
such generations may be continued without interruption for a
period of four years, and we have no reason to suppose that even
this could not be much exceeded under favourable conditions.
The parthenogenetic young may be produced either viviparously
or oviparously, according to species.

Oviparous and viviparous reproduction —The distinction
between these two processes has been extensively discussed, some
naturalists maintaining that they are thoroughly distinct ab
initio. This view, however, cannot be sustained. The best

the course of events seems to be that in summer there exist only wingless and
winged parthenogenetic females, and that the sexually perfect forms appear for the
first time in autumn.

1 Mitt. Schweiz. ent. Ges. iv. 1876, p. 529.

584 HEMIPTERA-HOMOPTERA CHAP,

authorities are agreed that in the earliest processes of individual-
isation the ovum, and the pseudovum' giving rise to a viviparous
individual, are indistinguishable. Leydig, Huxley, Balbiani, and
Lemoine are agreed as to this. Nevertheless, differences in the
development occur extremely early. The nature of these differ-
ences may be briefly described by saying that in the viviparous
forms the embryonic development sets in before the formation of
the ege is properly completed. Balbiani says, “In fact at this
moment [when the viviparous development is cammencing] the
germ [pseudovuim] is far from having obtained the development
it is capable of, and from having accumulated all the matter
necessary for the increase of the embryo, so that the evolution
of the former coincides, so to speak, with that of the latter. On
the other hand, in the true ovum the two processes are chrono-
logically separate, for the rudiment of the new individual never
appears before the egg has completed the growth of its constituent
parts.”* As regards the difference in structure of the organs of
viviparously and oviparously producing individuals, it is sufficient
to remark that they are not of great importance, being apparently
confined to certain parts remaining rudimentary in the former.
Leydig, indeed, found an Aphis in which certain of the egg-tubes
contained eggs in various stages of development, and others
embryos in all stages.’

As regards the physiology of production of winged and wing-
less individuals there has been but little exact inquiry. Vast
numbers of individuals may be produced without any winged
forms occurring, while on the other hand these latter are occa-
sionally so abundant as to float about in swarms that darken
the air; the two forms are probably, however, determined by the
supply of food. The winged forms are less prolific than the
apterous forms; and Forbes has noticed in Aphis maidi-radicis,
where the generations consist partly of apterous and partly of
winged individuals, that when the corn begins to flag in conse-
quence of the attacks of the Aphis, then the proportion of

1 The term pseudovum is applied, as a matter of convenience, to the earlier condi-
tion of the viviparously-produced form, and the term pseudovarium to the ovary
producing it.

? Balbiani, Ann. Sei. Nat. Zool. (5) xi. 1869, p- 29. For concise recent re-
marks on the early embryonic states, see Lemoine, Lull. Soc. ent. France, 1893,
1h Ibex.

3 Acta Ac. German. xxxiii. 1869, No. 2, p. 81.

VII APHIDAE 585

winged individuals becomes large.’ The appearance of winged
individuals is frequently accompanied by a peculiar change of
habit ; the winged individuals migrating to another plant, which
in many cases is of a totally different botanical nature from that
on which the apterous broods were reared: for instance Aphis
mali, atter producing several apterous generations on apple, gives
rise to winged individuals that migrate to the stems of corn or
grass, and feeding thereon commence another cycle of generations.
The study of this sort of Aphis-migration is chiefly modern, but
many very curious facts have already been brought to light ;
thus Drepanosiphum platanoides, after producing a certain number
of viviparous generations on maple (dee), quits this food-plant for
another, but after two or three months returns again to the
maple, and produces sexual young that lay eggs.” Histories such
as this are rather common. Even more interesting are the cases
of those species that, after some weeks of physiological activity
ona plant, pass into a state of repose on the same plant, and
then after some weeks produce sexual young. On the whole, it
would appear that the appearance of winged forms is a con-
comitant of decreasing nutrition. It is a very remarkable fact
that the sexually perfect females are invariably apterous, and this
is frequently also the case with the males. It is also highly
remarkable that the sexually perfect individuals are of com-
paratively small size. There are at least three kinds of males
in Aphidae—1, winged males; 2, wingless males with mouth
well developed; 3, wingless small males with mouth absent.
As regards some of these points the conditions usual in Insect
life are reversed? Huxley inclined to treat all these products of
a fertilised ege, that are antecedent to another process of gamo-
genesis (7.e. production with fertilisation), as one zoological indi-
vidual: in that case the Aphis zoological individual is winged
before attaining the mature state, and is wingless and smaller
when mature. Some species may have as a rule two, others
three, winged generations in a year.

Parallel series.—In certain cases individuals of one genera-
tion assume different habits, and so set up the phenomenon

1 Seventeenth Rep. Insects Illinois, 1891, p. 66.

2 Kessler, Acta Ac. German. li. 1887, pp. 152, 153.
% Tn connection with this the absence of a functional mouth in the imago state
of numerous Lepidoptera, and of Oestrid Diptera, should not be forgotten.

586 HEMIPTERA-HOMOPTERA CHAP.

known as parallel series. This has been recently investigated in
the genus Chermes by Blochmann, Dreyfus, and Cholodkovsky.
This latter savant informs us! that a wingless parthenogenetic
female of Chermes hibernates on a fir-tree—Picea excelsa—and
in the spring lays numerous eggs; these hatch, and by the effects
of suction of the Chermes on the young shoots, galls are formed
(Fig. 286), in which the Insects are
found in large numbers; when they
have grown the galls open, and allow-
ing the Insects to escape these moult
and become winged females. They
now take on different habits; some of
them remain on the Picea, lay their
egos thereon, and out of these there
are produced young that grow into
hibernating females, which next spring
produce galls as their grandmothers
did; but another portion migrates to
the Larch (Larix); here eges are laid,
from which proceed wingless partheno-
genetic females, that hibernate on their
new or secondary plant, and in the
following spring lay their eggs and

©

Fic. 285.—Chermes abietis; hiber- — , ‘ ‘ } ;
nating female or ‘‘winter- give rise to a dimorphic generation,

ae ( Ree fenaeee part of them becoming nymphs and
going on to the winged condition,

while the other part remain wingless and lay eggs, that give rise
to yet another wingless generation ; in fact, a second pair of parallel
series is formed on the new plant, of which one is wingless, and
exclusively parthenogenetic, and contimues to live in this fashion
for an indefinite period on the secondary plant, while the other
part becomes winged; these latter are called sexuparous, and go
back to the Picea, and there lay eggs, that give rise to the
sexual forms. If we would summarise these facts with a view
to remembering them, we may say that a migration of a part of
a generation from the Picea was made with a view of producing
a sexual generation, but that only a portion of the migrants suc-
ceeded in effecting the object of the migration, and this only in
their third generation. Thus portions remained on the Picea,

1 Tiorae Soc. ent. Ross. xxiv. 1890. p. 386.

,
he
QC

VIII APHIDAE 587

producing unisexual (female) individuals, and a portion of those
that emigrated to the Larix remained thereon, producing also uni-
sexual (female) individuals, while the others returned to the Picea
and produced a sexual generation. How long the production of
the unisexual generations may continue has not been determined.
Phylloxera—The Phyllowera, that has caused such an enor-
mous amount of damage in the Old World during the last thirty
years, is a smal] Aphid that was introduced from North America
into Europe. In North America it is not so injurious as it 1s
in Europe, owing, no doubt, to shght distinctions in the condi-
tions of life in the two hemispheres, as one of which may be
mentioned that in Europe a larger proportion of the imdividuals
produced appear to confine themselves to feeding on the roots, P.
vastatriz being one of the species that lives both in galls on
leaves, and underground on the roots. The species is one that
exhibits in their most complex form the peculiar phenomena of
Aphid life we have already mentioned. It has probably only
one congener, Phylloxera quercus, and of this Lichtenstein says
that in its cycle, from the starting-point of the winter-egg to
the assumption of the sexual condition,
it exhibits a series of no less than
twenty-one forms.’ The life of Phyl-
loxera vastatriz apparently differs essen-
tially from what we have described in
Chermes, masmuch as the migrations are
only between leaf and root of the same
plant—the vine—and not from one
species of plant to another. Some autho-
rities treat Phylloxera and Chermes as a
separate family under the name of Phyl-
loxeridae.

Galls— Like Phylloxera, many species
of Aphidae live partially, others wholly, Fis. 286.—Gall, or false cone
in galls that are produced by plants as hee eis acl ae
the result of one or more Aphids inter- ie Jet, is 8 pec
fering with a delicate part of the plant the pom line a, and
when it is in a young and growing Se eee conta
state. The usual position of Aphid galls eee OF
is on a leaf or leaf-stalk. But in the case of the genus Chermes,

1 Ent. Zeit. Stettin, xxxvi. 1875, p. 368.

5388 HEMIPTERA-HOMOPTERA CHAP.

a bud or some growing part of the spruce-fir is affected in such a
way that it gives rise to an object having externally the appear-
ance of a little fir-cone, while inside it consists of chambers in which
the Aphids reside. The forms of Aphid-galls are very diverse, but
this is probably due to the plant rather than to the Insect, for the
same species of Aphis may give rise to different forms of galls.
Réaumur thought that each Aphid-gall was due to a single indi-
vidual that irritated the tissue of the plant, so that-the latter
grew up at the point of irritation and enclosed the Insect.

A few points as to the anatomy of Aphids should be
noticed. It is doubtful whether the antennae have ever really
inore than six joints, the apparent seventh joint being actually a sort
of appendage of the sixth. The rostrum is externally three-jointed,
and is remarkable for the great diversity in its length, sometimes
it 1s quite short, at others several times longer than the body
(Fig. 285); the setae are often very much longer than the
sheath ; in cases where this great length of rostrum exists, the
individual may often be found with the tip firmly fixed in the
bark, and, as it were, tethered by means of the rostrum, the
length of which allows, nevertheless, considerable locomotion.
Suction is performed by contractions of the pharynx. There
has been much difference of opinion as to whether there is a
salivary syringe, and Witlaczil failed to find it. Kyrassilstschik is,
however, positive that it exists,’ and that it is analogous to that
described by Mayer in Pyrrhocoris, but there are great differences
of structure between the two. It is very difficult to determine
the number of segments at the extremity of the body; this is
terminated dorsally by a median organ placed above the anus,
and known as the cauda. Balbiani apparently considers that
there are ten abdominal segments and the cauda. The alimentary
canal has a small stomach, and an elongate intestine, the
terminal division of which is capacious and remarkably long.
There are no Malpighian tubes; according to Kowalevsky, their
function is discharged by the posterior part of the alimentary
canal. There exists, however, a peculiar structure, the pseudo-
vitellus, a sort of cellular, double string; and Witlaczil, in his
valuable paper” on the anatomy of Aphidae, suggests that this

1 Zool. Anz. xv. 1892, p. 220.
2 Arb. Inst. Wien, iv. 1882, Heft iii. p- 397 ; see on this organ also Mordwiiko,
7

VIII APHIDAE 5389

organ may in some way replace the missing Malpighian tubes.
Another highly pecular structure is the siphons, frequently
ealled nectaries, honey-tubes, or siphuncles. They are situated
on the dorsal aspect of the fifth abdominal segment, but exist
only in certain of the sub-families; they are of very different
lengths according to the species, and are capable of movement ;
they open directly into the body cavity, though exceptional
openings into the body cavity are extremely rare in Insects.
They excrete a waxy matter, which first appears as oil-like
globules. It was formerly supposed that they were the means
of secreting the sugary matter, called honey-dew, so much prized
by ants and some other Insects; but this is now ascertained
to be erroneous. This matter comes from the alimentary canal,
and is secreted in large quantities by some species, Biisgen having
observed that forty-eight drops, each about 1 mm. in diameter,
were emitted by a single individual in twenty-four hours.’ Certain
gall-dwelling Aphidae—Pemphigus, Chermes (Fig. 285), Schizo-
neura—possess numerous wax glands; these seem to replace the
siphons, and excrete the peculiar, whitish flocculent matter that
is so conspicuous in some of these Aphids.

Earlier anatomists failed to find any dorsal vessel, and it is
consequently reported in books to be absent. It has been, how-
ever, recently detected by Witlaczil, and Mordwilko states that
it does not differ from that of other Insects.

We have already alluded to the fact that the mode of repro-
duction of Aphids leads to an unrivalled increase. This, however,
is not due to the prolificness of the individual, which, in point of
fact, appears to be considerably below the average in Insects, but
rather to the rapidity with which the young begin to reproduce.
This has been discussed by Huxley, Buckton, and others. The
first-named naturalist calculated that the produce of a single
Aphis would, in the course of ten generations, supposing all the
individuals to survive, “contain more ponderable substance than
five hundred millions of stout men; that is, more than the whole
population of China.”* It has since been contended that
Professor Huxley’s calculation was much below the mark.
Although it is somewhat difficult to make a calculation dealing
adequately with the actual facts, yet it is clear that the increase

1 Biol. Centralbl. xi. 1891, p. 193.
2 See, inter alia, Webster, J New York ent. Soc. i. 1893, p. 119.

590 ITEMIPTERA-HOMOPTERA CHAP.

of Aphids is such that, drawing as they do their nutriment
directly from the plant in its growing state, in the course of
two or three years there would be no nutriment available for
other animals, except such as might be derived from plants not
attacked by Aphids. The numbers of Aphidae would be so great
that they could not be expressed by ordinary numerical methods,
and their increase would be actually limited only by the relations
existing between different kinds of plants, and between plants
and Aphids. This result is avoided by the fact that Aphids are
themselves the victims of a whole army of Insect enemies. They
have the numerous members of a special group (Braconidae,
Aphidudes) of minute Hymenoptera to live inside their bodies, and
many Aculeate Hymenoptera depend entirely on the Aphidae as
the source of food for their own progeny. The Lady-birds-——
Coccinelidae—live on Aphids and Coccids, and themselves in-
crease to such an extent as to be in many years a conspicuous
part of the Insect world. Crowds of the larvae of Hemerobiids
and Syrphids are constantly engaged in spearimg and sucking
the Aphides. Hence the old naturalist Bonnet said that, just
as we sow grain for our benefit, Nature has sown Aphids for the
benefit of multitudes of different Insects. He might have added
that these different Insects are for the benefit of man, it being
clear that without them the population of the world must
rapidly decrease.

Ants treat Aphidae more intelligently than most other Insects
do, for they do not destroy the helpless creatures, but utilise
their products in the way man does those of the cows he keeps.
The relations between ants and Aphids is itself an extensive
chapter in Natural History; many facts have been brought to
light showing that the ants manage the Aphids in a prudent or
intelligent manner, distributing them when too numerous in one
place, keeping guard over them, even building shelters for them,
and in some cases keeping them in direct association, by retaining
the Aphids in their own dwellings. The further investigation of
these points goes, the more it tends to raise the actions of the
ants to the level we call in ourselves intelligent. It would even
appear that the ants are acquainted with the migrations of the
Aphids from one species of plant to another, Webster informing
us that as the Aphis-population on an apple tree multiplied the
ants in attendance anticipated their migration to wheat and grass

VIII ALEURODIDAE 59I

by carrying them to those plants.’ We have nearly 200 species
of Aphidae in Britain,” and there may perhaps be 800 known
altogether. To what extent they may occur in the tropics is
undetermined. There are said to be no native species in New
Zealand. ;

Fam. 8. Aleurodidae.—J/inute Insects, with four mealy wings,
seven-jointed antennae, two-jointed feet, terminated by two claws
and a third process. These minute Insects are at present a source
of considerable perplexity, owing to the curious nature of their
metamorphosis, and
the — contradictory
accounts given of
them. In the earlier
stages they are
scale-like and qui-
escent, being fixed
to the under side of
a leaf. The French
authors Signoret and
Girard state that the
young are hatched
having visible ap-
pendages and seg-
mentation, but that
amveG st (bhey axe
attached to the leaf
the organs gradu-
ally suffer atrophy.
Maskell states the Fic. 287.—Instars of <Alewrodes immaculata. Europe.

7 : ; os Jace ps above : =
opposite, saying that (After Heeger.) A, Nymph, from above; B, nymph,
‘2 under surface ; C, imago.

TU (G
Nu

Nu

the organs in the

earliest stages are not usually recognisable, but become faintly
visible with the growth of the Insect. Heeger states that
the larva undergoes three ecdyses, and he gives the figures
we reproduce; if he be correct it would appear that the
nymph undergoes a great development. Réaumur, on account
apparently of their great metamorphosis, treated the species

1 J. New York Ent. Soc. i. 1893, p. 120. See also as to knowledge on the part
of ants, Forbes, Highteenth Rep. Insects Lilinois, 1894, pp. 66, ete.
2 Monograph by Buckton, Ray Society, 4 vols. 1879-1883.

592 HEMIPTERA-HOMOPTERA CHAP.

known to him as being Lepidopterous, though he correctly
pointed out their distinctions. At present we can only con-
clude that the Aleurodidae undergo a metamorphosis of a kind
peculiar to themselves, and requiring renewed investigation. The
family has been monographed by Signoret, and more recently by
Maskell, who has increased the number of species to about sixty.’
We have three or four in Britain, one of which, A. brassicae, is
extremely abundant on various kinds of cabbage in certain years.
Fam. 9. Coccidae (Scale - Insects, Mealy - bugs). — Insects,
usually minute, with only a single claw to the foot; the male with
one pair of wings, but without mouth-parts ; the female wingless
and usually so degraded in form that most of the external organs
and appendages cannot be distinguished. The form in which these
Insects are most generally known
is that of a small scale or shell-like
body closely adhering to leaves,
fruits, or bark. The scales are of
the most varied form, so that no
general description can be given of
them. The scale may be defined
as an accumulation of excreted
matter, combined with the cast
skin or skins of the Insect, cover-
ing the body either totally or
partially, and thus acting as a shield

ee emcee nae A, Aspidiotus yyder which the subsequent devel-

vmelliae, on the stem of a plant; B, i

a female scale magnified. (After opment takes place. All Coccidae
ree) do not form scales; but the habit of
excreting a large quantity of peculiar matters to the outside of
the body is universal ; this excreted substance is frequently white,
and of a powdery nature, and Coccids of this kind are known as
mealy-bugs. In other cases the exudation is like shell or glass,
and the creature may become quite encysted therein. In this way
the forms of Cocidae known as “ground-pearls” are formed. When
first hatched from the egg Coccidae are mite-like creatures, and it
is only subsequently that the females lose the power of loco-
motion. The females of numerous forms of Coccidae—more
particularly the mealy-bugs—do not lose the antennae and legs.
There is also a group (Brachyscelides) of Coccids that live in

1 Tr. New Zealand Inst. xxviii. 1895.

VII ; COCEIDAE——SCALE-INSECLS 593

galls. This highly aberrant group is, however, peculiar to
Australia; elsewhere very few gall-making Coccids have been
discovered.

There are upwards of 800 species of Coccidae at present
known.' The family was monographed by Signoret about twenty-
five years ago, and since then there has been
very much matter concerning them published
in a scattered manner.” No general work
has been published on the British species,
but Mr. Newstead is preparing one. The
classification of Insects so minute as Coc-
cidae, and with such extreme difference in
the sexes, is, of course, a matter of great
difficulty ; the best divisions are those given
by Green in his Coccidae of Ceylon.’

The fact that there is only one pair of
wings in the perfect male Coccid would appear
to ally these Insects with the Diptera; these
Coecidae have, too, like the Diptera, a small
appendage on each side of the metathorax.
Witlaczil shows that these little processes may
really represent a pair of wings, inasmuch as
they are developed from imperfect folds of Tig, 989. Dachilohius
hypodermis, 7.e. imaginal dises. Beyond these —— Jonyispinus. Female
facts and the occurrence in certain females on portion of a fig-

; : é leaf. (After Berlese.)
(Margarodes) of a great histolysis during the
post-embryonic development, there is nothing to indicate any rela-
tionship between Coccidae and Diptera. It has been shown by
Riley that these little processes, in some forms, serve as hooks to
attach or control the true wings, and this function is never assumed
by the halteres of Diptera. Although Coccidae are placed next
Aphidae, yet the two families appear to be really very different.
The modes of reproduction so peculiar in Aphidae reappear to a
certain extent in Coccidae, but are associated with profound

1 A catalogue of Coccidae has recently been published by Mr. T. D. A. Cockerell
in Bull. Illinois Lab. iv. 1896, pp. 318-339.

2 Signoret’s papers are to be found in eighteen parts in Ann. Soc. ent. France,
1868 to 1876: the most considerable subsequent systematic papers are those by
Maskell in the Transactions of the New Zealand Institute from 1878 to the present
time.

3 Coccidae of Ceylon, pt. 1, 1896, p. 16.

VOL. VI : 2Q

594 HEMIPTERA-HOMOPTERA CHAP.

distinctions. Though the viviparous method of reproduction
and parthenogenesis occur in Coccidae, yet they are only ex-
ceptional, and they are not put to the same uses by the species
that exhibit the phenomena. Thus we have seen that in Aphidae
generations of imperfect individuals are produced with rapidity,
while the individual is not directly very prolific. In Coccidae
the reverse is the case—the generations are usually similar to
one another; they do not, as a rule, follow with rapidity, and the
female is usually very prolific, thousands of young being some-
times produced by a single individual. The extraordinary poly-
morphism of the species of Aphidae is not exhibited by Coccidae,
though, contrary to what we find in Aphidae, the males and
females are usually excessively different. The two families ap-
parently also differ in that Coccidae are specially characteristic
of warm climates, Aphidae of the temperate regions.
Parthenogenesis. —Owing to the fact that the males are very
minute creatures, totally different from the females, and living
but a very short time, they were but little known to the earlier
observers. It was therefore only natural to suppose that par-
thenogenesis was very common. Of late years the males of a great
many species have become known, so that ordinary sexual repro-
duction must be considered as the normal method in Coccidae,
although, in the great majority of cases, the male is still unknown.
It has, however, been shown in numerous cases that parthenogenesis
may occur even when males exist; and there are some abundant
species of which it has not been possible to find a male. In
1887 Moniez’ announced that he had discovered the male of
Lecanium hesperidum (one of the notoriously parthenogenetic
species) in an ovarian cul-de-sac in the body of the female, and
he therefore considers that sexual reproduction occurs. He does
not say how pairing takes place, and we are not aware that his
observation has been confirmed. If correct it will be necessary
to reconsider the whole question as to parthenogenesis in
Coccidae. Apterous males are known in two or three species.
The post-embryonic development of Coccidae is of the most
unusual character. It is quite different in the two sexes, and in
each of them it presents features not found elsewhere. It has,
however, as yet been studied in only a few forms, and even in
them is incompletely known. When hatched from the egg

LC) Rt Ac. Sti. Paris, cive 1887, p. 449°

ee ee ee ee

VIII COGCIDAE—SCALE-INSECTS 595

the young Coccids are all similar, male and female being indistin-
guishable. A difference
soon appears, with the
result that the male, after
passing through more
than one pupal condi-
tion, appears as a winged
Insect. The female never
becomes winged, but, if
we may judge from the
incomplete accounts we at
present possess, her de-
velopment varies much
according to species. In
some she retains the legs,
antennae, and mouth-
organs; in others she
loses these parts, though
retaining the original
form ina general manner ;

while in a third (Mar- Fic. 290.—Instars of Dactylopius citri. (After Ber-

garodes) she becomes en- lese.) A, Egg; B, young larva; C, first male
sted d ee tl nymph ; D, second male nymph; E, adult male ;
eee appar muy F, adult female. All equally magnified. x 20.

suffers an almost com-

plete histolysis, reappearing after a very long period (it is said
it may be as much as seven years) in a considerably altered form.
The post-embryonic development of Aspidiotus nerii has been
studied by Schmidt’ and Witlaczil,” whose accounts agree except
as to some points, such as the number of ecdyses. The young, or
larva, is hatched with fairly well-developed legs, antennae, and
rostrum; there is no external difference between the sexes. The
larva selects some spot on the plant and drives its rostrum therein,
thus becoming fixed; moults occur, and the body excretes waxy
matter from its sides in processes that fell together and form the
shield; the female becomes much larger than the male. The
legs and antennae of both sexes disappear, so that the power of
movement is completely lost. The mouth-parts also atrophy. The
female after this undergoes no further change, except that of
growth in connection with ovarian development. The male,

1 Arch. Naturgesch. li. i. 1885, p. 169. 7 Zeitschr. wiss. Zool. xiii. 1886, p. 156.

596 HEMIPTERA-HOMOPTERA CHAP.

however, continues development ; notwithstanding the impossi-
bility of taking food, owing to the absence of a mouth, it increases
much in size, and the organs of the future perfect Insect
commence to develop from imaginal discs in a manner similar to
that which occurs in the Dipterous genus Corethra ; no mouth-
parts are however developed, these being merely represented by
spots of pigment, or rudimentary additional eyes. The wings
are developed outside the body. Difference of opinion prevails as
to the nature of the instars between the young larva and the
imago. It is clear, however, that Fig. 291, D, corresponds fairly

-Fic. 291.—Development of male of Aspidiotus nerii. A, Newly hatched larva; B,
prae-pupal instar ; C, pupa before ecdysis ; D, pupa shortly before the emergence
of the imago: a, antenna; e, eye; /, wing-rudiment; /, leg; 0, basal part of
mouth-organs. (After Schmidt.) Magnification not definitely stated.

with the pupa of Insects with complete metamorphosis, and the
instars shown in Fig. 291, B, C, may therefore be looked on as
equivalents of the resting-larva stage of ordinary Insects with
complete metamorphosis. Witlaczil considers this development
to be a condition of incomplete, approaching very nearly to
complete, metamorphosis. The condition is perhaps more pre-
cisely estimated if we recollect that winged Insects are divided
into two series, in one of which the wings are developed outside
the body ; in the other, inside the body. The Insects with very
complete metamorphosis all belong to the second of these two
series, while in the male Coccid we have the highest form of
metamorphosis attained by any of the first series. As regards
the development of the female encysted nymph or pupa,
previously alluded to as being found in the “ ground-pearls” of

VII COCCIDAE—-SCALE-INSECTS 597

the genus Margarodes, we can at present offer the reader no
satisfactory account.'

Products of Coccidae——Honey-dew is secreted by Coccidae,
but as a rule not so extensively as by Aphidae and some other
Homoptera ; nevertheless, it is often sufficient tomake the plants
frequented by Coccids very sticky and unclean. Some species
make a really extensive exudation of such matter. Réaumur
records that a Coccid, which is doubtless Lecanium persicae,
excretes a supply of honey-dew that drips to the ground; he says
it tastes sweet and nice. The manna mentioned in the book of
Exodus is pretty certainly the honey-dew secreted by Coccus (now
Gossyparia) mannifera, which lives on Tamarix in many places
in the Mediterranean basin. This substance is still called by the
Arabs “ Man,” and is used as food; in its natural state it is a
substance very like honey; it is doubtless excreted by the
Coccus, and is not produced directly by the Zamariz as some
have supposed. Waxy matters are produced by several Coccidie.
Ceroplastes ceriferus, a Lecaniid, produces white wax in India.
Ceroplastes is a widely distributed genus, and various species of it
have been used for the purpose of producing wax in other parts
of the world. The white wax of China is understood to be pro-
duced by another Lecaniid, Fricerus pela; but little is known as
to this Insect; it is said that the wax is produced by the winged
males. The substance was formerly greatly prized in China, but
is falling into disuse on account of the introduction of Kerosene.
Lac is produced by Carteria lacca, a Lecaniid living in India on
Anona squamosa, as well as on species of Ficus, Rhamnus and
other trees; the lac is the shelly scale produced by the Insect
as a covering; it is composed in larger part of resinous matter,
with which there is mixed a comparatively small quantity of wax
and other substances. The body of this Insect also affords the
red substance called lake. Various species of Kermes formerly
afforded a red dye well known to the Greeks and Romans. These
Insects live on Quercus coceifera in the Mediterranean region. A
medicinal syrup is also obtained from them. Porphyrophora
polonica was used in North and Central Europe for the same pur-
poses as Kermes ; it is a Coccid living on the roots of Polygonum
cocciferum. These European Insects were replaced commercially

1 For summary as to our present knowledge of this curious condition of Insect
life, see Mayet, Ann. Soc. ent. France, 1896, p. 419.

598 HEMIPTERA-HOMOPTERA CHAP.

after the discovery of America by the cochineal Insect, Coccus
cacti, a Mexican Coccid feeding on a Cactus called Nopal
(Opuntia coccinellifera). This Insect was subsequently introduced
to the Eastern hemisphere, and was established with more or
less success in a few spots on the borders of the Mediterranean.
In the Canary Islands it flourished on other species of Cactus,
became acclimatised, and was the object of an extensive commerce.
The colour in the case of all these Coccid dyes was obtained from
the bodies of the Insects, in the tissues of which it is contained.
The dyes have now been largely displaced in commerce by the
derivatives of Aniline. Axin is produced by the Mexican Coccid
Llaveia axinus ; this substance appears to be of a very peculiar
nature ; it is apparently chiefly fatty, and contains a peculiar acid,
axinic acid. Axin is used as an external medicinal application
in various affections; and it is also employed as a varnish; it
dries and hardens on exposure to the air, and is said to be of
considerable value." In our British genus Orthezia the body of
the female is completely covered with a symmetrical snow-white
armour, from which project the pink legs and antennae. This is
one of the forms in which the female preserves the legs to the
end of her life. The objects called ground-pearls, already alluded
to, have long been known in various parts of the world, and in
the island of St. Vincent they are sufficiently large to be collected
and strung for necklaces. These bodies are the encysted pupae
of Coccids of the genus Margarodes; the cyst is said to be of
chitin. Jf vitis commits serious ravages on the vines in Chili
by sucking their roots, and it is probable that all the species are
of subterranean habits; this would partially explain the fact
that very little is known about the history of these pearls,
though naturalists have been acquainted with them for niany
years.

The gall-making Coccids of the group Brachyscelides have
only recently been at all investigated; the galls they give rise
to are sometimes about a foot in length, and there appear to be
numerous species and several genera in Australia; they are
especially abundant on Hucalyptus and Acacias. The females are
highly remarkable from the variable conditions the legs assume,
so that in some cases they may be described as biped Insects, the

1 For additional information as to useful Coccidae, see Blanchard, Bull. Soc. Zool.
France, viii. 1883, p. 217.

VIII ANOPLURA—LICE 599

hind legs remaining, though the others have atrophied.’ Very
little indeed is known as to these Insects. One of the most
peculiar points of their economy appears to be that the galls
giving rise to males are different from those producing females.

Anoplura or Lice.

Small Insects with thin integument; entirely wingless, the
three thoracie segments indistinctly separated; the head
bearing in front a short tube furnished with hooks; from which
tube there can be protruded another very delicate sucking-
tube. Feet terminated by a single long claw. The Anoplura,
Pediculidae, or lice are disgusting Insects about which but little
is known. The most contrary opinions have been expressed as
to their mode of taking their nourishment, which is, without
exception, the blood of Mammals; on the bodies of which they
pass the whole of their life. It is a most
difficult matter to examine their mouth ;
the best information on this point is given
by Schiddte and Graber, but though these
two authorities agree, their results are very
incomplete, and do not warrant us in ex-
pressing a confident opinion as to the nature
of the relationship between Hemiptera and
Anoplura—a question that has been for
long a moot one. The short tube furnished
with hooks in front (Fig. 293, d) is con-
sidered to be the lower lp, and the tube
inside is, it is suggested, a combination of Fic. 292.—Pediculus capi-

5 : tis, 9. Human head.
the homologues of maxillae and mandi- (After Piaget. )
bles; there is also what may be a labrum
(g); and inside the head a framework, at any rate analogous to
if not homologous with, the parts of this kind we have described
as existing in Hemiptera. All the parts, with the exception of
the basal tube or head of the beak, are of the most minute and
delicate nature, so that it is difficult to see their form or com-
prehend their relations. It is evident that they are very different
anatomically from the mouth-parts of Hemiptera; still there is

1 Rubsaamen’s paper on these Insects gives references to most of the previous
literature, Berlin. ent. Zeitschr. xxxix. 1894, p. 199. :

600 ANOPLURA CHAP.

sufficient general resemblance to warrant the belief that the parts
in the two may ultimately be shown to be also morphologically
similar. If Meinert be correct, this
view will, however, not prove to have
any foundation. He considers that
morphologically the mouth of the louse
has no similarity to that of the bug;
the protrusible parts in the former he
considers to be modifications of epi-
pharynx and hypopharynx; and the
rod-like structures to be hypopharyngeal

NPE eee (ee a lamellae; and that they are thus totally
Chitinous envelope into different from the setae of bugs’ He
we the peak can pe wit” considers Lice to be a distinct Order of

drawn ; d, head of the beak,

with crown of spines; g, Insects for which he proposes the name
labrum ; h, delicate tube

protruded (very rarely seen Siphunculata.
in this state) ; m’, unpaired The alimentary canal and nervous

ote system resemble those of Mallophaga
more than they do those of Hemiptera. The oesophagus leads
junto a large stomach bilobed in front; at thé posterior extremity
of this there open the four Malpighian tubes, and behind these
there is a well-marked small intestime. The nervous system
consists of a cephalic ganglion and of three other closely approxi-
mated ganglia, the posterior. one the larger. It remains
doubtful whether or not the first of these three gangha is the
infra-oesophageal one.”

The species of lice, so far as known, are not numerous, some
six genera and about forty species being all that are recorded ;
they occur on various kinds of mammals, including some that live
in water. Seals have.a genus, Echinophthirius, peculiar to them.
Monkeys are specially hable to be affected by lice; the genus
that chiefly occurs on them is Pedicinus, a very distinct one, in
which there are only three instead of five joints to the antennae.
Perhaps the most remarkable louse is Haematomyzus elephaniis,
that of the elephant; it has a long proboscis in front of the head.
As a rule each species of louse is confined to one species of Mam-
malia, or to very closely allied forms. Man is said to be infested

1 Ent. Meddel. iii. 1891, p. 82.
2 Cf. Graber, Zeitschr. wiss. Zool. xxii. 1872, p. 165, and Landois in the same
Journal, xiv. 1864, p. 24.

Vill LICE 601

by three species, Pediculus capitis, P. vestimenti and Phthirius
inguinalis ; Meinert is of opinion that P. capitis and P. vestimente
are only one species, and Schiddte appears also to have thought
this probable. Andrew Murray was of opinion that the heads
of different varieties of men are infested by distinct varieties of
P. capitis. His conclusion was chiefly base\ on examination of
specimens preserved by Charles Darwin ; it requires confirmation.
Very little is known as to the life-history of the louse. Leeuwen-
hoek made himself the corpus vile for an experiment, from which
he concluded that the Pediculus vestimenti is very prolific. That
scientific men did not know whether the louse bites or sucks was
formerly made the ground for a taunt. Schiddte has given an
almost pleasing account of the way in which he settled this,’
showing that the sucking action is beyond all doubt. Accounts
of disease called Phthiriasis, attributed to lice, are to be found in
many old books, but the evidence does not warrant us in believing
anything more than that persons suffering from some disease, and
in a neglected and filthy condition, were horribly infested with
these disgusting Insects.

It is usual to say that Pediculidae are Hemiptera degraded by
a long exclusive persistence in parasitic habits. At present,
however, this must be looked on as a pious opinion, rather than
as an induction from our knowledge of their morphology and
embryology ; for this is at present too imperfect to warrant any
final conclusion.

1 Ann. Nat. History (3), xvii. 1866, p. 213.

NOTE.

Since the remarks on the classification of Hymenoptera were written Mr.
W. H. Ashmead has published several important papers proposing a classi-
fication to a considerable extent new. He adopts 10 superfamilies and 94
families. His views are summarised in P. U. S. Mus. xxiii. No. 1206, 1900,
Pastor Konow has also discussed this subject in Hnt. Nachr. xxiii. 1897, pp.
148-156.

INDEX

Every reference is to the page: words in italics are names of genera or species ; figures
in italics indicate that the reference relates to systematic position ; figures in thick
type refer to an illustration ; f. = and in following page or pages ; n. = note.

Abdomen, of Chrysis, 2; of Coleoptera,
185; of Diptera, 446; of Hemiptera,
538; of Lepidoptera, 318; of Thy-
sanoptera, 528

Abdominal legs, 9

Abeille-perce-bois, 33

Abeille tapissiere, 51

Abispa, 77

Acacia jistulosa, beetles in, 213

Acalyptrate Muscide, 494

Acanthia, 560

Acanthomeridae, 483

Acanthosoma grisewm, 546

Acari, relations to Insects, 220, 223, 238,
530

Acentropus, 425

Acephalous larvae, 449

Achreioptera, 219

Acraeides, 350

Acridium maroccanum, 254

Acrocera globulus, 490

Acroceridae, 489

Acronycta, 418

Actias luna, 374

Actiidae, 510

Acutilingues, 20

Adapted excrement, 284, 284, 380

Adelops, 221

Adensamer, on Ascodipteron, 520

Adephaga, 190, 200 f., 216, 234

Adimeridae, 240

Adimerus setosus, 241

Adlerz, on Formicoxenus, 160 ; on Tomo-
gnathus, 161

Adminicula, 327

Aédes, 455 n.

Aegeria, 387

Aegeriidae, 386

Aegialitidae, 265

Aegocera tripartitu, 411

Aenictus, 159, 179, 180

Aenigmatias blattoides, 495
Aeolothrips fasciuta, 528
Aépophilidae, 559

| Aépus, 206

Aérostatic setae, 408

Aérostats, 449

Aganaidae, 408

Agaristidae, 370, 871, 410

Agdistes, 426

Agdistinae, 426

Agenia carbonaria, A. hyalipennis, 105

Ageronia, 354

Aglycyderes setifer, 298

Aglycyderidae, 297

Agromyzidae, 504

Agrotis, 415; A. spina, 417

Ahuatle, 504

Alaenda, 350

Alaopone, 179, 180

Alar organs—see Wings, Elytra, Tegmina

Aletia xylinae, 416

Aleurodes brassicae, 592; A. immaculata,
591

Aleurodidae, 591

Alucita, 426 ; A. polydactyla, 426

Alucitidae, 371, 426

Alula, 447

Amara, 205

Amber, Insects in, 144, 269, 458

Amblyopone, 180

Amblyoponides, 132, 180

Ambrosia, 295

Ambryllis, 409

Amicta quadrangularis, 394

Ammophila, 111; A. affinis, 111; A.
hirsuta, 111; A. holosericea, 111

Amphicyrtides, 242

Amphidasis betularia, 412, 414

Amphiodont, 193

Amphipneustie, 450

| Amphizou lecontei, 207

604

INDEX

Amphizoidae, 207

Ampulex compressa, 114, 115; A. rufi-
cornis, 115 ; A. sibirica, 114

Ampulicides, 114 f., 169

Amycterides, 291

Anal armature, 328, 416

Anal nervures, 318

Anaphe, 376

Anaspini, 267

Anaspis, 268

Anatomy—see External Structure and
Internal Anatomy

Anchor-process, 459

Ancient, Lepidopteron,
Primitive

Ancylolominae, 425

Andrena, 23, 25, 30, 301, 303 ; hair of,
11; A. labialis, 488 ; A. nigroaenea,
25; A. ovina, 30

Andrenidae, 20

Andrenides, 23

Andrenimorpha, 388

Andrenoides, 20

Androconia, 331 f.

Anergates atratulus, 160 f.

Angelitos, 63

Anisopteryx aescularia, 411

Anisotoma cinnamomea, 222

Anisotomidae, 223

Anisotomides, 222

Anlagen, 143

Anobiides, 247

Anobium, 254; A. paniceum, 247; A.
striatum, 248 ; A. tessellatwm, 248

Anochetus ghiliani, 174

Anomma, 178

Anophthalinus, 205

Anoplura, 599 f.

Anosia erippus, A. menippe, 345; A.
plexippus, 307 ; larva, 324; pupa, 327

Antennae ; of Belostoma, 566 ; of flies, 441;
of Lamellicornia, 191 ; of Lepidoptera,
307 ; of butterflies, 340, 341 ; of Sphin-
gidae, 380

Antherophagus, 235

Anthicidae, 266

Anthidium, 45; A. bellicosum, 47; A.
diadema, 45; A. manicatum, 45; A.
septemdentatum, 47 ; A. strigatuin, 30

Anthocopa, 51

Anthocoridae, 560

Anthomyia angustifrons, A. brassicae, A.
cana, 506

Anthomyiidae, 506

Anthophila, 10 f.

Anthophora, 82, 33; destroyer of, 272,
274; A. personata, 33; A. pilipes, 33 ;
proboscis of, 17

Anthothrips aculeata, 530

Anthracides, 486

Anthrax, 486 f. ; A. fenestralis, 489 ; A.
trifasciata, 44

435 — see also

Anthrenus fasciatus, 241

Anthribidaé, 278, 290

Antisquama, 448

Antitegula, 447

Ant-plant, 138, 139, 168

Ants, 131 f.; and Aphidae, 590; and
caterpillars, 356

Ants’-nest, Insects, 200, 213, 221, 223,
224, 225, 231, 236, 240, 548; larva,
501, 502

Anus, 314, 320

Aorta, 320

Aortal chamber, 320

Apate capucina, 246

Apatela, 418

Apathus = Psithyrus, q.v.

Apatidae, 246

Apatura ; larvae, 354; A. iris, 344

Apaturides, 352

A phaenogaster, 164,165,221 ; A. arenarius,
164 ; A. barbarus, 131, 164 ; A. structor
164, 240

Aphaniptera, 522 f.

Aphanocephalus, 228

Aphelocheirus, 565

Aphidae, 581 f. ; and ants, 181

Aphidiides, 590

Aphis maidi-radicis, 584

Aphomia sociella, 424

Apidae, 10 f., 20, 32

Apioceridae, 492

Apis, 53 ; A. adansonii, 69 ; A. domestica,
68 ; A. dorsata, 69 ; A. fasciata, A.
ligustica, 68; A. mellifica, 65 f.; feet
of queens and workers, 69 ; ligula of,
16 ; worker and hairs, 12

Apoda, 402; A. testudo, ete., 401, 402

Apodidae, 402

Apoica pallida, nest, 83

Aporia crataegi, 322

Appetite, 491

Apterogyna, 96

Apterona, 393, 394; A. crenulella, var.
helix, 395

Apterous, beetles, 187, 263 ; females, 95,
96, 140, 171, 174, 315, 392, 393, 407,
413, 4380, 592—see also Workers ; In-
sects, 95, 96, 505, 474, 495, 496, 518,
531, 581; males, 140, 160, 161, 172,
585, 594

Aquatic ; caterpillar, 377 ; cocoons, 280 ;
larva, 421 f., 425, 504; pupa, 423

Aradidae, 550

Aradus orientalis, 550

Araeocerus, 290

Araschnia levana, A. prorsa, 353

Arbelidae, 369, 396

Arceina, 373

Archiapidae, 21, 22

Archiapides, 27 f.

Arctia caja, 308 ; A. villica, 410

Arctiidae, 370, 404, 408, 410

INDEX

Argiva, 414

Argynnis, larvae, 354; A. paphia, andro-
conia, 332

Argyromoeba sinuata, 76 ; A. trifasciata,
486

Arista, 442

Army-worm, 416

Aromia moschata, 188

Arthropterus, 214

Asclera caerulea, 267

Ascodipteron, 520

Asemorhoptrum lippulum, 160

Asilidae, 491

Asilus, 492 ; A. crabroniformis, 441

Asparagus-beetle, 281

Aspidiotus camelliae, 592; A. nerti, 595,
development of, 596

Aspidiphorus, 246

Aspidomorpha, pupa, 283, 284

Association, of <Amnergates and Tetra-
moriuwm, 160 ; of ants and other Insects
ete., 180 f.; of Formica and Formi-
coxenus, 159 ; of Strongylognathus and
Tetramorium, 162; of Tomognathus and
Leptothorax, 161

Astata boops, 119

A statides, 119

Asteidae, 504

Astomella lindeni, 490

Astynomus, 285

Atemeles, 225 ; and ant, 182

Athericerous, 441

Atheriz, 481; A. ibis, 480

Athous rhombeus, 257

Atractocerus, 254

Atrophy of mouth and stomach, 310

Atta, 137, 164 n., 165, 502

Attacus, 373 ; A. atlas, 373

Attelabides, 291

Attini, 158, 159, 165

Attitude, 381, 384, 385, 388, 412, 413,
425, 429

Atylotus fulvus, 483

Autocrates aenea, 275

Automeris, 373

Axin, 598

Azteca, 158

Azygos oviduct, 321

Badamia exclamationis, 365
Barrett, on increase of melanism, 414
Basket-worms, 393
Bat-parasites, 521, 560
Bataillon, on metamorphosis, 306
Bates, H. W., on classification of butterflies,
344; on homoeochromatism, 351; on
Megacephala, 201
' Batesian mimicry, 337, 339
Bathyscia, 221
Beak, 532
Beauregard, on vesicating Insects, 275
Beaver, Insect on, 219, 221

605

Becher, on mouth of Diptera, 444 n.

Bed-bug, 559; enemy of, 558

Beddard, on animal coloration, 339 n.

Bee (i.e. honey-bee)—see Apis mellifica

Bee-louse or -tick, 520

Bees, 10 f. -

Bees born of carcases, myth, 499

Bees’ nest beetle, 235

Bees, stylopised, 300, 303

Beetles, 184 f.

Bellesme, on buzzing, 19

Belostomidae, 534, 565

Bembecidae, 482

Bembecides, 119 f.

Bembex, 509 ; B. rostrata, 4, 120, 120 f. ;
B. spinolae, 130 n.

Benchucha bug, 559

Berosus, 218

Berytidae, 548

Bibio, 475, 476, 477; B. marci, 477

Bibionidae, 475

Birds and butterflies, 338

Biscuit-weevil, 247

Bitoma crenata, 233

Bittacomorpha, 473

Black-fly, 530

Blanchard, on flies attacking man, 517 n.

Blepharoceridae, 464

Blind beetles, 205, 221, 233

Blissus leucopterus, 548

Blister-beetles, 269

Blochmann, on founding new nests, 145

Blood-sucking, Diptera, 457 ; Mosquitoes,
467

Blood-worms, 468

Blow-tlies, 511

Blue-bottles, 511

Bogus Yucca-moth, 433

Boletophila luminosa, 463

Boll-worm, 416

Bombardier-beetles, 201

Bombus, 53 f.; insect in nest ef, 221 ; fleas
in nests of, 525; parasite of, 94, 497 ;
proboscis of, 13 f.,14; 6. agrorum, 54 ;
B. lapidarius, 54 ; B. muscorum, 57 ; B.
variabilis, 60

Bombyces, 367

Bombycidae, 368, 375, 406

Bombyliidae, 485

Bombylius major, 488

Bombyx mori, 375 ; B. yamamai, 325

Book-worm, 247

Borboridae, 504

Borborus, 505

Borocera madagascariensis, 405

Bostrichidae, 246

Bot-flies, 514

Brachelytra, 224

Brachycera, 441, 454

Brachycerides, 291

Brachyscelides,*592, 598

| Brachytarsus, 290

606

INDEX

Braconidae, 590

Bradypus cuculliger, Tineid on, 430

Brahmaeidae, 368, 374

Brain, 320 ; cephalic and thoracic, 449

Branchiae, 208, 244

Brands, 332

Brassolides, 349

Brathinus, 223

Brauer, on Dipterous larvae, 451 ; on Oes-
tridae, 514

Braula coeca, 520

Braulidae, 520

Breastbone, 459

Breeze-flies, 443, 481

Breitenbach, on proboscis of Lepidoptera,
311 n.

Breithaupt, on proboscis of bee, 15; on
deglutition of bees, 18

Brenthidae, 295

Brenthus anchorago, 297

Brephos notha, 415, 416

Brimstones, 357

Brontes planatus, 234

Brown-tail moths, 407

Bruchidae, 276

Bruchus fabae and B. lentis, 277, B. pisi,
277

Bryophila, 418

Buckell, on development of pattern, 335

Buffalo-gnats, 477

Bugong-moth, 417

Bugonia-myth, 499

Bull-dog ants, 771, 173

Bull’s-horn thorn and ants, 168

Bumble bee—see Bombus

Buprestidae, 261

Buprestis attenuata, supposed larvae of,
262 n.

Burgess, on suction, 311

Burnet-moths, 390

Burrows, of Dasypoda, 27; of Halictus, 24,
25 ; of Odynerus, 74

Bursa copulatrix, 321

Burying-beetles, 221

Buttertlies, 347 f.

Buzzing, 19

Byrrhidae, 242, 255

Byrrhus pilula, 242

Bythoscopidae, 578

Byturus, 241

Cadphises moorei, 391

Calandrides, 289

Calcium oxalate, 406

Calicurgus, 101 ; C. hyalinatus, 102, 106

Caligo eurylochus, 350

Callidea baro, 303 n.

Callidulidae, 370, 400

Calliphora, 448 ; C. erythrocephala, C. vom-
itoria, 511

Cullirhipis dejeani, 256

Callomyia, 496

Callostoma fascipennis, 489

Caloptenus, 270 ; C. italicus, 489, C. spretus,
488, 506, enemies of

Calypter, calypterate, 448

Calyptrate Muscidae, 448, 504

Camberwell Beauty, 352

Camel bot-fly, 515

Camponotides, 144

Camponotus, 145; C. ligniperdus, 138,
145, 147 ; C. pennsylvanicus, 138, 146 ;
C. rubripes, 1381; C. rujipes, 137

Camptosomes, 279, 281

Canephorinue, 394, 395

Cantharidae, 269 f.

Cantharides, 270

Capsidae, 561

Capsus laniarius, 539

Carabidae, 204 f.

Carabides, 206

Caraboidea, 790, 200 f.

Carcinocoris, 554

Carder-bees, 45 f. 45

Cardiocondyla, 161

Cardiophorus, 258

Carlet, on sting, 6; on sound-organs of
Cicada, 574 8

Carnivora, 200

Carotine, 549

Carpenter-bees, 33

Carpenter-worms, 395

Carpets, 411

Carpocapsa juliana, C. pomonella, C. splen-
dana, C. saltitans, 428

Carpophagus, 278

Carteria lacca, 597

Carus, on paedogenesis, 461

Caryoborus, 278

Case, 281, 392, 393, 394, 417, 422, 423,
430, 431

Cassidides, 279, 283

Caste-production, 142

Castnia, 807, 809, 316, 319 ; C. eudesinia,
C. therapon, 372

Castniidae, 369, 371

Castor canadensis, parasite of, 219 .

Cataclysta lemnata, 423

Caterpillar, 322, 324, 325; of Diptera, 474

Catopomorphus, 221

Cauda, 538, 588

Cave-beetles, 205, 221

Cebrionides, 260

Cecidipta excoecaria, 424

Cecidomyia buxi, 459 ; C. destructor, 460 ;
C. tritici, 460

Cecidomyiidae, 455 n., 458

Cecropia, plant and ants, 158

Cedeocera, 297 n.

Cell, of wing, 317, 318 ; complete and in-
complete, 116 n.

Cells, formation of, by bees, 21, 22, 24,
25, 28, 38, 34, 35, 46, 48, 51, 52, 54, 56,
60; earthen, 72, 106 ; of Coelonites, 89

INDEX

607

Celyphidae, 504

Celyphus, 505

Cemonus unicolor, 128

Cephaloidae, 275

Cephalomyia maculata, 515

Cephaloon, 275

Cephalothorax, 465 ; of Sty/ops-larva, 302

Cephenomyia rufibarbis, 517

Cerambycidae, 278, 285

Cerambycides, 287

Ceramixs, 89; C. lusitanicus, 89

Ceranchia, 374

Cerapachys, 175 n.

Ceratina, 11, 32

Ceratocampidae, 368, 375

Ceratocombidae, 559

Ceratognathini, 194, 295

Ceratonema, 401

Ceratopogon, 469; C. bipunctatus, C. puli-
caris, C. varius, 470

Cerceris, 125); Cl arenaria, 125);
bupresticida, 125; C. labiata, 125; C.
tuberculata, 126

Cercopidae, 577

Ceresa bubalus, C. taurina, 577

Cerocoma schaeffer’, 275

Cerophytides, 260

Ceroplastes ceriferus, 597

Ceroplatus masterst, 463

Cerura vinula, 383

Cervical sclerites, 472

Cetonia, in ants’-nests, 149; C.. floricola,
200

Cetoniides, 195, 199

Chaerocampa, 380; recte Choerocampa

Chaetophorous, 446

Chaetotaxy, 446

Chafers, 194 f.

Chaleosiidae, 391, 420

Chalia hockingi, 394

Chalicodoma, 32, 35: C. muraria, 30, 35
f., 36, 254, 486 ; C. parietina, C. pyre-
naica, 39

Change of habit in larva, 301, 431

Chapman, Dr. T. A., on Chrysis, 3; on
classification of pupae of Lepidoptera,
367 ; on Hepialus, 398; on Metoecus
paradoxus, 268; on pupa of Lepidop-
tera, 327 n.

Charagia, 396

Chartergus chartarius, nest of, 82, 83

Cheilosia chrysocoma, 439

Cheimatobia brumata, 414

Cheliomyrmex, 180

Chelonariides, 242

Chelonia, 410

Chelostoma, 35

Chennium bituberculatum, 224

Chermes, 583, 586, 587; C. abietis, 586,
587, 589

Cheshire, on proboscis of bee, 15

Chigger, 525

Child, on sense-organ, 442

China-marks, 421

Chinch-bug, 548

Chionea araneoides, 474

Chiromyzidae, 479

Chironomidae, 468, 474

Chironomus, 440, 468

Chlamydes, 279

Chlorion, 110

Chloropidae, 504

Chlorops, 504

Choerocampa elpenor, 380

Choerocampini, 381

Cholodkovsky on Chermes, 586

Choragus sheppardi, 290

Chorion, 322

Chrysalis, 326, 344

Chrysauginae, 423

Chrysididae, 1 f.

Chrysiridia madagascariensis, 419

Chrysis bidentata, 3; C. ignita, 33 ¢,
shanghaiensis, 4

_ Chrysochus pretiosus, 279

Chrysocoris grandis, 303 n.

Chrysomelidae, 276, 278 f.

Chrysomelides, 279

Chrysopolomidae, 396

Chrysops, 482

Cicada, 123; C. plebeia, 574; C. septen-
decim, 569

Cicadellinae, 578

Cicadidae, 568 f.

Cicindela hybrida, 202

Cicindelidae, 201 f.

Cicinnus, 878 n.

Cilix glaucata=spinula, 401

Cimex, 560 ; C. lectularius, 559

Cimicidae, 559

Cioidae, 245

Cis melliei, 245

Cistelidae, 264

Cistus salvifolius, beetle-larvae in, 282

Cithaerias, 348

Citheronia, 375

Cixiides, 576

Cixius, 575

Clambidae, 223

Clasper, 314

Classification, of ants, 144 ; of bees, 20 ;
of butterflies, 341 ; of Coleoptera, 189 ;
of Diptera, 454 f.: of Hemiptera, 543 ;
of Hymenoptera Aculeata, 70 ; of Lepi-
doptera, 339 f. ; of moths, 366 f.

Clavicornia, 189, 213, 265

Claviger testaceus, 224

Clavigerides, 224

Clavus, 539 |

Clear-wings, 386

Cleggs, 481

Cleosiris, 400

Cleptes, 2, 4

Cleridae, 253

608

Click-beetles, 256

Clicking butterfly, 354

Clidicus, 223

Clisiocampa neustria, 322

Clothes-moths, 450

Clouded-yellows, 357

Clypeus, 307

Clythra in ants’-nests, 149

Clythrides, 279

Cnemidotus caesus, 209

Cnethocampa processioned, 376

Coarctate larva, 271

Coccidae, 592 f. ; destroyer of, 290

Coccidula, 239

Coccinellidae, 237

Coccus cacti, 598 ; C. mannifera, 597

Cochineal Insect, 598

Cochliopodidae, 402 n.

Cochlophora, 394

Cockchafer, 198

Cockroach, parasite of, 269

Cock-tail, 225

Cocoon, 46, 55, 66, 109, 122; 328, 347,
373, 376, 384, 385, 891, 403, 404, 405,
407, 419, 424, 436, 460, 462, 494;
flax-seed, 460; of ants, 134; aquatic,
280, 377

Cocytia durvillii, 382

Cocytiidae, 382

Codling-moth, 428

Coelioxys, 31

Coelonites, 89 ; C. abbreviatus, cells of, 89

Coenomyia ferrugined, 480

Coenomyiidae, 479

Coenonympha, 348

Colaenis, 351

Coleophora, 431

Coleoptera, 184 f.

Colletes, 22; C. daviesanus, 30

Colobopsis, 138

Colon, 320

Colorado beetle, 278

Colour, corresponding with locale, 201 ;
and surroundings, 337; of larva and
habits, 336 ; of Sphingidae larvae, 381 ;
physiology of larval, 413; of cater-
pillars and sex, 325; development of,
“in Hemiptera, 542 ; of eyes, 440

Coluocera formicaria, 240

Colydiidae, 233, 234

Colydium, 233

Comb, 63, 64, 65, 78, 79

Combs and brushes, 134

Compound pupa, 452

Composmyia, 512

Comstock, on nervures, 317 n.

Conchylidae, 427

Connexivum, 538

Conopidae, 497, 504

Conorhinus sanguisuga, 559

Copiopteryx, 373

Copium clavicorne, 550

Crioceris asparagi, 281 ;

INDEX é

Copius, 547

Copper butterflies, 356

Coprides, 195 f.

Coprini, 195 n.

Copris hispanus, 197

Copulatory pouch, 320, 321

Cordyluridae, 504

Coreidae, 546

Corethra, 467

Corium, 539

Corixa, 567

Corixidae, 567

Corn-leaves, larva on, 281

Coronidia, 419

Corotoca, 227

Corticaria, 240

Corylophidae, 228

Corynetides, 253

Coscinocera hercules, 372

Cossidae, 369, 395.

Cossonides, 294

Cossus, 309; C. ligniperda, 319

Costal nervure, 318

Cotton-stainer bug, 548

Cotton-worm, 416

Courtship, 494 ; of Hepialus, 398 f.

Coxa, 307 :

Crabroy 129)" CG"
stirpicola, 130 n.

Crabronides, 128 f.

Crambidae, 425

Crane-flies, 471

Cratoparis, 290

Cremaster, 327, 328, 344, 426

Cremastochilini, 200

Cremastochilus, 200

Cremastogaster, 213, 165 ; C. tricolor, 165

Crepitation, 213, 214

Criocerides, 279, 280

cephalotes, 129; C.

C. merdigera,
281

Crossocerus, 130; C. wesimaeli, 1389

Cryptocephalides, 279

Cryptocephalus, 282

Cryptocerata, 544; 562 f.

Cryptocerini, 132, 134, 158, 159, 169

Cryptocerus, 138 ; C. atratus, 170

Cryptophagidae, 235, 257

Cryptophagus dentatus, 235

Cryptostomes, 279, 282

Cteniza ariana, destroyer of, 490

Ctenophora, 475

Ctenostylidae, 517

Ctenuchinae, 409

Cuckoo-bees, 22

Cuckoo-spit, 577

Cucujidae, 232, 234

Cucujos, 258

Cuculinae; 20

Culex pipiens, 466

Culicidae, 466 f.

Cultelli, 443

INDEX

609

Cuneus, 539, 540

Cupesidae, 234

Curculionidae, 290

Curtice, on Hypoderma, 516

Curupira, 465

Cut-worms, 415

Cyathoceridae, 243

Cybister laterimarginalis or roeseli, 210 ;
C. tripunctatus, 211

Cybocephalus, 232

Cyclica, 279, 282

Cyclorrhapha, 454; C. Aschiza, 455, 494 f.;
C. Schizophora, 456, 503 f.

Cylidrus, 253

Cylindrotomina, 474

Cymatophoridae, 368, 386

Cymbidae, 410

Cynomyia mortuorum, 510

Cyphagogus segnipes, 296

Cyphanta, 368 vn.

Cyphonia clavata, 576

Cyphonidae, 255

Cyrtidae, 489

Cyrtocorides, 545

Cyrtocoris monstrosus, 546

Dacnides, 237

Dactylopius citri, 595

Daddy-long-legs, 471

Dakruma coccidivora, 424

Dalla Torre, Catalogue of Hymenoptera,
21

Danaides, 344, 247

Danaioid Heliconiidae, 346

Danais archippus, or plexippus, 345

Dances, 351, 464, 493, 554

Darwin, C., on Pelobius, 208

Darwin, F., on proboscis of Lepidoptera,
311 n.

Dascillidae, 243, 255

Dascillus cervinus, 255

Dasychira pudibunda, 408; D.
407

Dasygastres, 20, 35 f.

Dasypoda hirtipes, 27

Dead-leaf butterfly, 353

Death-watches, 248, 254

December-moth, 406

Deer bot-fly, 517

Deer-fly, 518

Delphacides, 576

Deltocephalus inimicus, 578

Deltoidae, 418, 423

Denudatae, 20, 29

Deporaus, 291

Dermatobia noxialis, 517

Dermestidae, 241

Deroca, 400

Derodontidae, 244, 253

Derodontus maculatus, 245

De Saussure, on wasps’ nests, 81

Devil’s coach-horse, 225

ViOls vi

TOSsti,

Dewitz, on development, of sting, 8; of
thoracic appendages, 9

Dexiidae, 510

Diactor bilineatus, 547

Dianeura, 392

Diateina holymenoides, 547

Dichoptic, 440

Dicthadia, 178, 180

Dictyocicada, 543

Dilophus febrilis, 477 ; D. vulgaris, 476

Dimera, 544

Dimorphie, generations, 586 ; males, 161,
Wy

Dimorphism, 139 ; of wings, 549

Dinapate wrightii, 246

Dingar, 70

Dinoponera grandis, 182, 134, 171

Dionychopus niveus, 410

Diopsidae, 503, 504, 505

Diopsis apicalis, 503

Dioptinae, 409

Dioptoma adamsi, 251

Dioscorea batatas, beetle-larvae in,280

Dioxys cincta, 32, 43

Diphyllides, 237

Diplocotes, 248

Diplonychus, 566

Diploplectron, 119

Diploptera, 70, 71 f.

Diplosara lignivora, 429

Diplosis, 459 ; D. resinicolu, 459

Dipsocoris alienus, 559

Diptera, 438 f.

Dipterous parasitic larva, 26

Dirphia tarquinia, 377

Discocellular nervures, 318

Dismorphia, 346, 357

Dissociation of embryo, 70 n.

Diza, 471

Dixidae, 471

Dohrn, Anton, on Hemiptera, 538

Dolichoderides, 157

Dolichopidae, 493

Dolichopus undulatus, 441

Dolichurus haemorrhous, 116

Donacia, 280

Donaciides, 279

Dorsal vessel, 320 ; 529—see also Internal
Anatomy

Dorycera, 504

Doryceridae, 504

Dorylides, 174 f.

Dorylini, 175. 177

Dorilars Ne tee ll tte
178

Doryphora decemlineata, 278

Dragon, 383, 385

Drepanidae, 370, 400

Drepanosiphum platanoides, 585

Drepanulidae, 400

Drilides, 248

Driver ants, 178

179; D. helvolus,

bo
=)

610 INDEX

Drones, 63, 67, 69

Drosophilidae, 504

Drurya, 362

Dryomyzidae, 504

Dryophthorus, 289

Dubois, on luminescence, 259

Dudgeon, on Badamia, 365

Dufour, on host helping parasite, 26

Duke of Burgundy fritillary, 355

Duration, of ant-colonies, 154; of wasp-
colonies, 70 n., 80; of life—see Longevity

Durrant, on moth-cases, 431

Dutch bulbs, larva in, 501

Dyar, classification of larvae of moths,
367

Dycladia, 389

Dynastes, 199

Dynastides, 195, 199

Dysdercus suturellus, 548

Dytiscidae, 210 f.

Dytiscus, 211

Earias, 410

Eau de Javelle, 368 n.

Echinophthirius, 600

Eciton, 159, 175, f; EH. hamatum, 175,
WT

Ecitonini, 774, 175 f.

Ecpantheria, 409

Ectatomma auratum, 181

Ectrephes kingi, 248

Edible larvae, 287

Egg, 305, 485, 468; as food, 504, 568 ;
of bot-fly, 514, 515 ; of Capsidae, 561 ;
carried, 547, 551, 566; of Mndochus,
558 ; of Reduviidae, 559 ; of flea, 524 ;
laid by pupa, 469 ; of Lepidoptera, 321,
322; of Nepa, 564; numerous, 397,
few, 197 ; standing out string of, 378;
swallowed, 508

Egg-tubes, 321

Eggers, 322, 405

Elaphidion villosum, 286

Elaphomyia, 505

Elateridae, 256

Elaterides, 260

Eleodes, 263

Hlephantomyia, 472

Elmides, 244

Elymnias, 348

Elymniidae, 348

Elytra, 184, 186, 539

Embolium, 539

Embryonic dissociation, 70 n.

Emenadia flabellata, 269

Emery on classification of ants, 144 ; on
polymorphism in ants, 143

Emesiides, 555

Emperor-moth, 374

Empidae, 492, 494

Empodium, 446

Empretia stimulans, 408

Eneyrtus, 34; Enecyrtus fuscicollis, em-
bryology of, 70

Endomychidae, 237, 239

Endotrichiinae, 423

Endromidae, 369, 406

Energopoda, 457, 491

Enhydrus, 216

Entomophila, 20

Enzyme, 259

Epeolus variegatus, 3C

Ephestia kuhniella, 306, 424

Ephydridae, 504

Ephyra pendularia, 412

Epiblemidae, 427

Epicausis smithi, 409

Epichnopteryx, 395

Epicopeiidae, 368, 418

Epicranium, 307 .

Epicypta scatophora, 463

Epidapus scabiet, 462

Epilachnides, 238

Epimeron, 307

Epinotia funebrana, 428 ; E. hypericana,
parasite of, 476

Epipaschiinae, 423

Epipharyngeal sclerites, 14

Epipharynx, 14, 308, 443, 600

Epiplemidae, 368, 420

Epipyrops, 404

Episternum, 307

Epitritus, 170

Epuraea, 232

EKrastria scitula, 417

Erebia, 347 ; E. aethiops, 347

Erebides, 418

Eremochaeta, 457

Eremochaetous, 446

Lremocoris, 548

Ergatandrous, 140 n.

Ergatogynous, 140 n., 142

Ergatoid, 140

Ericerus pela, 597

Eriocephala, 308 ; E. calthella, 434

Eriocephalidae, 433

Eviocera, 472

Eristalis, 499

Ermine-moths, 409

Erotylidae, 235, 236

Erucaeformia, 475

Erycides, 364

Erycinidae, 341, 354, 358

Erycinides, 355

Ethon, 262

Eucephalous larvae, 450

Eucera, 32

Eucharis myrmeciae, 173

Euchloe cardamines, egg, 322; larva, 358,
359 ; pupa, 358; EH. genutia, 358

Euchroma goliath, 261

Eucinetus, 256

Eucleidae, 401

| Buclidia mi, 415

a ey

—

INDEX

O11

Eucnemides, 260

Eudaemonia, 373

Eudamus proteus, 340

Eueides, 351

Eugereonhockingl, 542

Luglossa, 34 ; L. cordata, 35

Eugnoristus monachus, 289

Eulema, 35

Eulen, 414

Eulyes, 558

Eumaeus, 355 i

Eumenes arbustorum, 73; E. coarctata,
73, 74; FE. conica, 74; L. jlavopicta,
72; E. pomiformis, 72; L. ungui-
culata, 73.

Eumenidae, 72 f.

Eumolpides, 279

Eumyiid flies, 456

Euparagia, 89

Kuphoria, 200

KEuplocia, 408

Euploea, 345

Eupoda, 279, 280

Eupsalis minuta, 296

Eupterotidae, 368, 376

Eurygona, larva, 355

Euschemon rafflesiae, 371

Eusemia villicoides, 410

Eusthenes pratti, 533

Euthyrhynchus floridanus, 546

Ezxcoecaria biglandulosa, 424

Excrement as covering, 281, 288, 463—
see also Adapted excrement.

Excremental dwellings, 284, 379

External structure, of Aphidae, 588 ; of
Chrysididae, 2; of Coleoptera, 185 ; of
Diptera, 439 f. ; of fleas, 523 ; of Hemi-
ptera, 534; of Hepialus, 400; of Hymen-
optera Aculeata, 5; of Lepidoptera,
307 ft. ; of Thrips, 527

Exudation of fluid, 238

Eye-collar, 387

Eyes, four in number, 215, 251, 476

Fabre, J. 1., on Ammophila, 111; on An-
thraz, 486; on Bembex, 120 f.; on
Calicurgus, 101; on Chalicodoma, 37 f.;
on Eumenes, 72; on Halictus, 24; on
Miltogramma, 509 ; on Osmia, 48 f. ;
on Scarabaeus, 196 ; on Scolia, 97 f. ;
on. Sitaris, 272; on Sphex, 108; on
Stelis nasuta, 30

False cones of Chermes, 587

Families of moths, key to, 368 f.

Feeding young, 147

Fertonius luteicollis, 130

Fever-fly, 477

Filaria, and Mosquitos, 468

Filicornia, 200

Finding nest, 38 f., 126

Finn, on protected butterflies, 345 n.

Flagella, 384, 442

Flatides, 576

Flax-seed cocoon, 460

Fleas, 522 f,

Fly-disease, 513

Food, abstinence from, 254; of
larvae, 19 ; small in amount, 277

Food-reservoir, 320

Footmen, 409

Forbes, on ant sounds, 155

Forel, Aug., on tropical American Ants,
138 ; on classification of Ants, 144

Forest-fly, 518

Formica exsectoides, 149; F. fusca, 137,
150; 1515 F. rufa, 148, 154; F. san-
guinea, 149 ; F. schaufussi, 152

Formicidae, 131 f.

Formicoxenus nitidulus, 148, 159

Fornaz, 260

Fossil, Ants, 143 ; Beetles, 261 ; Diptera,
458; Hemiptera, 542; Thrips, 531;
Wasps, 88—see also Palaeozoic

Fossores, 7, 10, 90 f., 346 ; classification,
93,

Fossorial solitary wasps—see Fossores

Founding new nests, Ants, 145 f.

Frenuluin, 316, 318, 319, 400

Friederich, on Parnid larvae, 244

Friese, on habits and classification of bees,
21; on hosts and parasites, 30 f.

Fritillaries, 352, 354

Froghoppers, 577

Frog-spit, 577

Frontal ganglion, 320

Fulgora candelaria, 575

Fulgoridae, 543, 574 ; larva living on, 404

Fulgorina, 543

Fumea, 393, 395

Fungus cultivated by ants, 167

Fungus-gnats, 462

Funnel-twister, 292

bee-

Gad-flies, 481

Gahan, J. C., on Praogena, 264

Galea, 309

Galerucides, 276

Galgulidae, 562

Galleria mellonella, 306, 331

Galleriidae, 423

Gall-midge-flies, 461

Galls, 262, 430, 424; of Coccidae, 598 ;
of Aphids and Phylloxera, 587; of
Psyllidae, 580 ; of Thrips, 530

Ganglbauer, on Staphylinidae, 224

Garden-whites, 357

Garman, on mouth-parts of Thysanoptera,
528

Gastropacha quercifolia, 405

Gastrophilus equi, 515

Gavara, 401

Gelechiides, 429

Gena, 185

Generations, 306

612 INDEX

Geometers, 411

Geometridae, 368, 411, 416, 420

Geomyzidae, 504

Georyssidae, 243

Georyssus pyginaeus, 243

Geotrupes, stridulation of, 195

Gerrides, 552

Gerris, 535, 552

thilianella filiventris, 555

Ghost-moths, 396

Giraffomyia, 505

Girdlers, 286

Glands, 321, 323, 331, 363, 399, 538,
553 ; accessory, 320; of Filippi, 324;
mandibular, 216 ; salivary, 326 ; silk-,
325 ; stink-, 257 ; wax-, 589

Glaphyrini, 795 n.

Glaphyroptera picta, 441

Glaucopides, 339

Glossa, 309

Glossina morsitans, 512, 513

Glow-worms ; 248 ; New Zealand, 363

Glyptus, 206

Gnats, 466, 468

Gnophaela, 409

Gnostidae, 223

Goat-moths, 895—see also Cossus

Godart on trumpeter-bee, 58

Godman and Salvin, on spermathecal
bodies, 321

Gold-tail moths, 407

Gonapophyses, 9, 305

Gonin, on development of wing, 328, 329

Goossens, on legs of Lepidoptera larvae,
323 ;

Gossyparia, 597

Graber, ou mouth of louse, 599

Grapholitha sebastianiae, 428

Grapholithidae, 427

Grass-inoths, 425

Grayling, 347

Green, E. E., on classification of Coccidae,
593

Green-bottles, 511

Green-fly, 581 f.

Ground-beetles, 204 f.

Ground-pearls, 592, 598—see also Mar-
garodes

Grypocera, 341

Guest-ant, 159

Gula, 185

Gymnocerata, 544, 544 f.

Gymnodomes, 82

Gyrinidae, 207, 215

Gyrinus, 215

Haase, on mimicry, 339 n.

Hadrus lepidotus, 482

Haematobia serrata, 512

Haematomyzus elephantis, 600

Haematopota, 482; H. pluvialis, 483,
443

Haemoglobin, 468

Haemonia, 280; H. curtisi, 280

Haetera, 348

Hag-moth, 403

Hairs, plumose or feathered, 11, 12; of
Dermestid larvae, 241—see also Setae

Half-loopers, 415

Halias, 410

Halictus, 23; H. lineolatus, 24; H.-

malachurus, 23; H. maculatus, 25; H.
moro, 2h: WHE vguddrieinctus, » 22),
25; H. rubicundus, 26; H. sexcinctus,
24, 269

Haliplidae, 209

Halirytus amphibius, 474

Halobates, 552; H. sobrinus, 551

Halobatodes, 553

Halteres, 438, 448, 593

Halticides, 278, 27

Hamadryas, 347

Hammock-moth, 379

Hampson, on classification of moths, 367
f.; on clicking butterfly, 354 ; on frenu-
lum, 316

Hampsonia pulcherrima, 391

Handlirsch, on Bombus, 58

Harpactorides, 558

Harpalides, 206

Harpalus, 205 ; H. caliginosus, 185

Harpes, 314

Hart, C. A., on larvae of Diptera, 473

Hart, J. H., on the parasol-ant, 142

Harvesting ants, 164

Hatchett Jackson, on colour of larvae,
325

Haustellata, 366

Haustellum, 308

Hawk-moths, 380 f.

Head-vesicle, 442

Hearing, organs of, 191, 313

Heath-buttertly, 347

Hebridae, 551

Hebrus, 551

Hecatesia, 371 s

Heerwurm, 464

Heliconiidae, 346

Heliconiides, 351

Heliconius, 346 ; H. erato, H. melpomene,
Hi. rhea, 351

Heliothis armigera, 416

Helluodes taprobanae, 206

Helochares, 218

Helodes, 255

Helomyzidae, 504

Helopeltis, 561

Helotidae, 235

Hemaris, 383 .

Hemerodromia, 493

Hemidiptera haeckeli, 553

Hemi-elytra, 539

Hemileucidae, 374

Hemiptera, 532 f.

—_—”

INDEX

Henicocephalidae, 504

Hepialidae, 306, 369, 396 f.

Hepialus, 309, 310, 311, 317, 319, 322;
H. humuli and others, 396 f.; HW. lupuli-
us, 397

Heredity, 454

Heriades, 53

Hermatobates haddoni, 553

Hermetia, 479

Hesperiidae, 341, 342, 363

Hessian fly, 452, 460

Hestia idea, 340

Heterocera, 340, 366, f.

Heteroceridae, 279, 243

Heterogenea, 402

Heterogeneidae, 402

Heterogyna, 10, 131 f.

Heterogynidae, 369, 392

Heterogynis, 369 n., 392

Heteromera, 190, 262 f.

Heteroneuridae, 504

Heteronotus trinodosus, 576

Heteroptera, 532, 534, 535, 539, 543

Heterotarsini, 264

Hezatoma pellucens, 441

Heylaerts, on Psychidae, 392

Hibernation, of Vanessa, 352

Hilara, 493

Hilbrides, 405

Hill-ant, see Formica rufa

Hill-grub, 417

Himantopteridae, 392

Himera pennaria, 411

Hippobosca equind, 518

Hippoboscidae, 518

Hippopsini, 288

Hirmoneura obscura, 485

Hispa, 282

Hispides, 279, 282

Histeridae, 230

Histia, 391

Histoblasts, 453

Histolysis, 452, 595

Hockings, on stingless bees, 63

Hoffer, on Bombus, 54

Hollandiidae, 396

Hololepta, 230

Holometopa, 504

Holoptiec, 440

Holoptilides, 557

Holymenia, 547

Homoeochromatism, 337, 351

TIomoeoderus mellyi, 193

Homomorpha, 542

Homophysinae, 421

Homoptera, 532, 534,
parasite of, 303, 497

Honey, 18, 80

Honey-ant, 152

Honey-bee—see Apis mellificu

Honey-dew, 580, 589, 597

Hook-tips, 400

535, 548, 544;

613

Hoplopus, 74

Horn, G. H., on classification of Carabidae,
206 ; of Silphidae, 225

Horn, W., on classification of Cicindelidae,
202 n.

Hornet, 81, 87

Hornet’s-nest beetle, 227

Horus as food, 430

Horse bot-fly, 515

Horse-flies, 481, 518

Hot springs, Insects in, 479

House-fly, 511

Hover-flies, 498 ©

Hubbard, on ambrosia-beetles, 295 ;
Phobetron, 403 ; on Xenos, 303

Huxley, on Aphids, 585, 589 ; onsclerites
of oesophagus, 15

Hybocampa mithausert, 385

Hybotinae, 492

Hydnophytum, 139 ; H. montanum, 138

Hydrocampa nymphaeata, 421

Hydrocampidae, 421

Hydrocampinae, 421

Hydrocores, 562

Hydrocorisae, 562

Hydrocyphon deflexicollis, 255

Hydrometra, 552; H. stagnorum, 551

Hydrometridae, 551

Hydrophilidae, 216 f.

Hydrophilides, 219

Hydrophilus piceus, 217

Hydroporides, 201

Hydroporus, 211, 212

Hydroscaphidae, 228

Hydrous caraboides, 218

Hygrobia, 208

Hylecoetus dermestoides, 255

Hylemyia strigosa, 506

Hymenitis, 346

Hymenoptera Aculeata, 4 f. ;
feray if:

Hyper-metamorphosis, 270, 488

Hypertely, in Kallima, 354

Hyphydrus, 212

Hypnody, 489

Hypocephalus armatus, 288

Hypochlorite of potash, 368 n,

Hypoderma bovis, H. lineata, 515

Hyponomeuta, parasite of, 70 n.

on

H. Tubuli-

- Hypopharyngeal sclerites, 14, 17

Hypopharynx, 15 n., 324, 443, 524, 600
Hypopygium, 446

Hypsidae, 370, 408

Hypsoides radama, 376

Hystrichopsylla talpae, 523

Tdia fasciata, 513

Idolothrips spectrum, 527
Imaginal dises or folds, 596
Imbauba ant, 158

Individual, 585

Inedible associations, 338, 339

614 INDEX

Infericornia, 548

Infra-oesophageal ganglion, 541

Inquilines, 30, 81

Insects as food, 417, 504, 568

Instars, of Aspidiotus, 596 ; of Coccidae,
595; of Hpicauta, 271

Instinct, 235, 274, 373, 403, 424, 487,
546 ; of Ants, 590; of Bembes, 121;
of Chalicodoma, 37 ; of Dasypoda, 28 ;
of Melipona, 64 ; of Miltogramma and
Bembex, 509; of _Odynerus, 76; of
Osmia, 48 f. ; of Pompilus, 102; of
Rhynchites, 292 ; of Trigona, 64

Internal anatomy, of Diptera, 449; of
Hepialus, 400 ; of Hemiptera, 540 ; of
Lepidoptera, 319 f.; of larva of Lepi-
doptera, 324; of Lice, 600

Internal nervures, 318

Iodine, 213

Ipides, 232

Ischium, 523

Ischnogaster, 82, 88 ; I. mellyi, 87

Isoderminae, 550

Isomera, 190

Issides, 576

Ithomia, 346 ; I. pusio, 346

Ithomiides, 346, 351

Ftuna, 346

Ityraea nigrocincta, 576

Jassidae, 578

Jiga, or ‘‘mimic me,” 92

Johnston’s organ, 442

Jordan, on antennae of butterflies, 841 n. ;
on Thysanoptera, 529 f.

Jugatae, 366

Jugum, 316, 400

Jugurthia, 89

Julodis, 262

Jumping-beans, 428

Kallima, 353

Karbi, 63

Kellogg, on Lepidopterous structure, 307 f.

Kentish glory, 406

Kermes, 597

Knot-horns, 424

Komarovia victoriosa, 99

Koo-chah-bee, 504

Kootchar, 63

Koptorthosoma, 70 n.

Korschelt, on formation of eggs of Wepa,
564

Kiinekel d’Herculais, on Volucella, 501

Kungu cake, 467

Laap Insects, 581

Labella, 443

Labidus, 175, 176, 180

Labium of Lepidoptera, 310

Laboulbéne, on sound-production by
Arctiidae, 410

Lac, 597

Lacinia, 309

Laciniata, 366, 434

Lacon murinus, 257

Lacosoma chiridota, 378

Lady-birds, 237

Lagoa opercularis, 404

Lagoidae, 404

Lagria hirta, 264

Lagriidae, 264

Lake, colour, 597

Lamellicornia, 790 f. ; enemies of, 97

Lamiides, 287, 288

Lamprosomides, 279

Lampyrides, 248

Lampyris noctiluca, 250

Langelandia anophthalma, 233

Languriides, 237

Lantern-flies, 575

Laparosticti, 795 n.

Laphria, 492

Lapidicolous beetles, 205

Lappet-moth, 405

Laricobius, 253

Larra anathema, 117; L. pompiliformis,
117

Larrada, 117 ; L. modesta, 118

Larrides, 116 f.

Larvae, of beetles, 188 f., 188 ; of Chrysis,
3; of Dasypoda, 28 ; of Hymenoptera,
7; of Lepidoptera, 323 ; of Sphex, 109

Lasiocampidae, 369, 375, 405

Lasiorhynchus barbicornis, 297

Lasius alienus, 140; founding nest, 146 ;
L. fuliginosus, 138, 153; L. niger, 158

Latridiidae, 240

Latridius minutus, 240

Latter, O., on Puss-moth, 384

Leaf-cutting ants, 165 f.

Leaf-nests (ants), 155

Lecanium hesperidum, 594; L. oleae,
417 ; L. persicae, 597

Ledra, 545, 578

Legs, abdominal, 9

Leistus spinibarbis, 204

Lema melanopa, 281

Lemoniidae, 354

Léon, on Hemidiptera, 553

Lepidoptera, 304 f.; L. Haustellata, L.
Laciniata, 366

Leptalis, 346, 357

Leptidae, 479

Leptinidae, 220

Leptinillus, 221

Leptinus testaceus, 220

Leptis scolopacea, 441, 481

Leptocircus, 362

Leptoderini, 221

Leptogenys, 171 ; L. falcigera, 171

Leptomastax, 223

Leptothorax acervorum, 161

Lerp Insects, 581

INDEX

Leto, 397 ; L. venus, 396

Leucania unipunctata, 416

Leuckart, on Melophagus, 519

Leucospis, 46; L. gigas, 44

Leuthner, on Odontolabis, 193

Libythea, 542

Libytheides, 355

Lice, 599 f.

Ligula, 16

Limacodes, 489

Limacodidae 370, 401 ; parasite of, 4

Limnas chrysippus, 345

Limnichides, 242

Limnobia intermedia, 472

Limnobiinae, 472

Limochores taumas, 340

Limothrips denticornis, 530

Lindemann, on injuries from Thrips, 530

Lingua, 16

Lingula, 15

Liometopum microcephaluin, 158

Lipara lucens, 128, 451

Liparidae, 406

Liponeura brevirostris, 465

Liponeuridae, 464

Lipoptena cervi, 518

Liptena, 356

Lipteninae, 356

Lita solanella, 430

Lithophilus, 239

Lithosiidae, 408

Lithosiinae, 409

Llaveia axinus, 598

Lobster, 383 ; Lobster-moth, 385

Loepa newara, 374

Lohita grandis, 549

Lomaptera, 200

Lomechusa, 142, 225

Lonchaeidae, 504

Lonchopteridae, 490

Longevity, 33, 135, 286, 306 ; of Cicada,
569 ; of Melolontha, 198 ; of Scarabaeus,
197

Longicorifs, 285

Loopers, 411, 415

Lorum, 14, 14, 16

Lowne, on blow-fly, 449; on
Dytiscus, 211

Lucanidae, 193

Lucanus cervus, 194; antenna of, 191

Luciferase, Luciferine, 259

Lueilia, 511, 512; L. caesar, L. macel-
laria, I. sericata, 512

Luciola, 249 ; L. italica, 249

Ludia delegorguet, 373

Luminescence, 250, 258 f., 463, 469

Lunula, 442

Lutz, on exudation of Coccinellidae, 238

Lycaena baetica, 356

Lycaenidae, 341, 355, 356

Lycides, 248

Lycorea, 346

foot of

615

Lyctides, 246

Lygaeidae, 548

Lymantriidae, 370, 406

Lymexylon navale, 254

Lymexylonidae, 254

Lyonnet, on anatomy of caterpillar, 324 n,

Machaerota guttigera, 578

M‘Cook, on honey-tub ants, 152
Macrocneme, 389

Macroglossini, 380

Macrolepidoptera, 340

Macronychus quadrituberculatus, 244
Macrotoma heros, 287

Maggot, 449

*Malachiidae, 252

Malachius aeneus, 252

Malacodermidae, 248, 252, 266

Malaxation, 110, 126

Malpighian tubes, 320, 334, 429, 449, 460,
466, 473, 529, 540, 588

Mamitllo curtisea, 378

Man—see Manna

Mandibles, of Lepidoptera, 308 ; of pupa,
436, 437

Mandibulata, 434, 536

Manna, 597

Manson, on Mosquitoes, 468

Manticora maxillosa, 203 ; MM. tuberculata,
202

Marane, 377

Marbled-white, 347

Marchal, on embryonic dissociation, 70 n. ;
on Ammophila, 111; on pigments, 254

Margarodes, 595, 597, 598; M. vitis, 598

Marimbouda da casa, 118

Maruina, 471

Masaridae, 88 f.

Masaris, 89 ; M. vespiformis, 88, 89

Mason-bee, 35 f.—see Chalicodoma; of
New Zealand, 107

Mastigus, 223

Matthews, on Hydroscaphidae and Cory-
lophidae, 228 ; on Sphaeriidae, 227

Maxillae, of Lepidoptera, 309

Mayer, Paul, on Hemiptera, 536

Meadow-brown, 347

Meal-worm, 263, 264

Mealy-bugs, 592

Mechanitidae, 346

Medeterus ambiguus, 493

Median nervure, 318

Megacephala, 201

Megachile, 35,51; M. albocincta, 52, 53;
M. anthracina, 62; M. fasciculata, 52 ;
M. lanata, 53; M. proxima, 53

Megalopides, 279, 282

Megalopygidae, 404

Megalybus gracilis, 490

Meganostoma, 543

Megarrhina, 445, 467

Megascelides, 279

,

O16

INDEX

Megasoma, 199

Megathymus, 371

Megistorhynchus longirostris, 485

Meijere, on stigmata, 450

Meinert, on Anoplura, 600; on mouth of
Diptera, 444; on paedogenesis, 461; on
Stylops, 302

Melandryidae, 265

Melanism, 414

Melanitis, 351

Melanophila decostigma, 261

Melecta, 31, 33; M. luctuosa, 31

Meliboeus, larva, 355

Meligethes, 232

Melinaea, 351

Melipona, 53, 62; M. scutellaris, 64

Melitaea, larvae, 354

Mellifera, 10

Mellinus, 123; M. arvensis, 123,124; I.
sabulosus, 124

Meloe, 33, 274

Meloidae, 269

Meloides, 270

Melolontha vulgaris, 194, 198

Melolonthides, 795, 198

Melophagus ovinus, 518, 519

Melophorus inflatus, 153

Melyridae, 252

Membracidae, 576, 578

Membracis foliata, 577 n.

Membrane, of Hemiptera, 539

Menorhynchous, 542

Merodon equestris, 501

Merrifield, temperature experiments, 337

Mesodont, 193

Mesophragma, 312, 445

Mesoscutellum, 312

Mesoseutum, 311

Mesosternum, 307

Mesovelia, 551

Mesozoic beetles, 261

Messor, 164

Metamorpha, 351

Metamorphosis, 529 ; of Alevrodidae, 591; |

of Cicada, 571; of Coccidae, 594 f. ; of
Diptera, 452 ; of Hemiptera, 542
Metapneustic, 450
Metascutellum, 313
Methoca ichneumonides, 96
Metochy, 183
Metoecus paradoxus, 268
Meyrick, classification of
367
Miastor, 461
Mice, insects in nests of, 221
Microdon, 501, 502
Microlepidoptera, 240, 427 ;
Micropezidae, 504
Microphysides, 560
Microptera, 22
Micropterism, 549
Micropterygidae, 369, 435

trophi, 309

Lepidoptera,

|
|
|

Micropteryx, 307 f., 317, 319, 327, 433
435, 436

Midge, 461, 470

Migration, Aphis-, 585

Mik, on Hilara, 493

Milichiidae, 504

Miltogramma, 508, 121

Miimacraea, 356

Mimesa bicolor, 128

Mimesides, 127

Mimicry, 337 f.—see also Resemblance

Mines, Dipteron in, 474

Miscophus, 116

Models, 346

Moesa-blight, 561, 562

Mollusca, eaten by beetle-larvae, 252;
larvae mistaken for, 501—see also Snails

Molossus, parasite of, 560

Monarch-butterfly, 345

Monda rhabdophora, 393

Monema flavescens, 4

Moniez, on fertilisation of Coccidae, 594

Monodontomerus nitidus, 44

Monohammus confusus, 286

Monomera, 544

Monommidae, 265

Monomorium, 560; AL. pharaonis, 163

| Monotomides, 240

Montezuinia dimidiata, nest of, 83
Mordella, 268

Mordellidae, 267

Mordellistena floridensis, 268
Mormolyce, 205

Mormolycides, 206

Morphides, 348

| Morpho, 315, 331, 349 ; M. menelaus, 318 ;

M. achilles, M. epistrophis, larvae of, 349

| Mosquito, 466 f.
| Mosquito-bees, 61

Mosquito-blight, 562

Moth-flies, 470

Moths, 366 f.

Motuea fly, 482, 122

Mouth, absence of, 310, 443, 489, 514,
515, 585, 596

Mouth-parts, of Diptera, 442 f. ; of fleas,
523 ; of Hemiptera, 534, 535; of Hy-
menoptera Aculeata, 13 ; of Lepidoptera,
307 f. ; of Lice, 599, 600

Mud-dauber, 113

Miiggenburg, on Melophagus, 518

Miiller, Fritz, on Imbauba-ant, 158; on
Trigona, 64

Miiller, H., on Dasypoda hirtipes, 27

Miiller, W., on South American larvae,
344 n.

Miillerian mimicry, 339

Mundstachel, 527, 528

Murray, A., on Lice, 601

Musca domestica, 511

Muscidae, 511 f., 517; M. Acalyptratae,
503 f.; M. Calyptratae, 448, 504

7
‘

INDEX

Musotimidae, 423

Mustiliidae, 376

Mutilla, 94,95; M. ewropaea, 94

Mutillides, 94 f.

Mycetaea hirta, 239, 240

Mycetaeidae, 239

Mycetobia pallipes, 462, 463

Mycetophagidae, 237

Mycetophilidae, 462

Mydaidae, 491

Mygnimia, 105

Myiasis, 512

Myiatropa florea, 499

Myiodocha tipulina, 557

Myoditini, 267

Myopinae, 497, 498

Myrapetra, 82

Myrmecia, 171, 172, 173

Myrmecocystus hortideorum, 152; M. inel- |
liger, 152; M. mexicanus, 152

Myrmecodia, 139

Myrmecophilous Insects, 181 f.

Myrmedonia, 226

Myrmica laevinodis, 148 ; M. rubra, 133 ;
M. rubra, races ruginodis, scabrinodis,
163

Myrmicides, 158
Myrmicini, 159

Nabides, 556 ao

Nabis lativentris, 556

Nacerdes melanura, 266

Naclia ancilla, 390

Nagana, 513

Nagel, on digestion by injection, 212

Nagoda nigricans, 401

Nanosella fungi, 228

Nassonoff, on Strepsiptera, 301 n., 302

Naucoridae, 565

Necrobia ruficollis, 253

WNecrophorus, 221

Nectar, 18

Neleus interruptus, antenna, 191

Nematobola orthotricha, 431

Nematodes, in Thysanoptera, 53Q

Nematois metallicus, 321

Nemeobiides, 355

WNemeobius lucina, 3385, 355

Vemestrina, 455 n.

Nemestrinidae, 484

Nemocera, 440; N. Anomala N.
456

Nemognatha, 304

Nemosomides, 233

Neocastniidae, 372

Neolepidoptera, 366

Neotropidae, 346

Nepa, 541; N. cinerea, 563

Nepidae, 544, 563

Nepticula, 431

Nervous system, Coleoptera, 191

Nervules, 319, 429

Vera

G17

Nervuration, of Diptera, 447; of Lepi-
doptera, 317 f.
Nervures, 318 , 319; development of, 329
f. ; swollen, 347, 348 ; of Eumenes, 73
Nests, of ants, 186 f.,155; of Pormica fusca
and Solenopsis fugax, 137 ; of Porphyras-
pis, 284 ; of wasps, 79 f. 79, 81, 83, 87

Netrocera, 347 n.

Neuroblasts, 453

Newbigin, Miss, on pigments, 334

Newport, on Meloe, 270

Nicagini, 195

Nicolas, on Halictus, 24

Nilionidae, 265

Nitidula, 232

Nitidulidae, 231, 235

Noctuidae, 311, 370, 410, 411, 414 f.

Node, 131, 134

Nolinae, 409

Nolidae, 410

Nomada, 30; N. lathburiana
sexfasciata, 30

Nonne, 407

Nopal cactus, 598

Nosodendrides, 242

Notocyrtus, 558

Notodontidae, 305, 368, 383

Notodontina, 417

Notonectidae, 567

Number of species: of Apidae, 10; of
Butterflies, 343 ; of Coccidae, 593 ; of
Coleoptera, 184; of Diptera, 438 ;
of Fleas, 525; of Hemiptera, 543 ;
of Lepidoptera, 306; of Lice, 600; of
Thrips, 527

Nun, 407

Nurses, 66, 134

Nyctalemon, 419

Nyctemeridae, 409

Nycteolinae, 410

Nycteribia, 521, 522

Nycteribiidae, 521

Nymph, of Cicada, 569 ; of Thrips, 529

Nympha inclusa, 452

Nymphalidae, 341, 343 f.

Nymphalides, 351

Nymphipara, 518

Nymphula stagnata, N. stratiotata, 423

Nyssonides, 123

30; Wy.

Oak-Pruner, 286

Obtusilingues, 20

Oceanic bugs, 552

Ocelli, 325 ; O. compositi, 325
Ochthiphilidae, 504

Ocneria, 407 n. ; O. dispar, 408
Ocnerodes, 489
Ovcnogyna, 409
Ocypus olens, 225
Odontolabis sinensis, 193
Odontomachi, 177, 173
Odour, of bugs, 541

‘

618

INDEX

Odynerus, 74 f., 269; O. antilope, 75 ;
O. callosus, 76; O. parietum, 33; O.
punctum, 773; O. reniformis, 73, 75 ;
O. spinipes, 3, 76

Oecocecis guyonella, 430

Oecodoma, 137, 164,165 ; Oe. cephalotes, 133

Oecophylla smaragdina, 147

Oecophoridae, 429

Oedeagus, 314 [recte aedeagus]

Oedematopoda princeps, 387

Oedemeridae, 266

Oestridae, 514

Oestrus ovis, 517

Ogcodes, 489, 490

Oil-beetles, 269

Oinophila v-flavum, 430

Olethreutidae, 427

Olibrus bicolor, 231

Oligarces, 461

Oligonephrous, 542

Oligoneura, 461

Ommatophorinae, 414

Oncideres, 286

Opetia, 496

Ophideres fullonica, 311

Opomyzidae, 504

Opostega, 429

Orange-tip, 357—see also Huchloe

Orectochilus, 216

Oreta hyalodisca, 400

Origin of parasitic bees, 32

Orneodidae, 340, 371, 426

Ornithoptera, 360 ; O. brookiana, 362 ; O.

. paradisea, 360, 361, 362
Orphnephilidae, 470

Ortalidae, 504, 506

Ortalis, 447

Orthogenya, 494

Orthezia, 541, 598

Orthogonius, 206

Orthoperus atomarius, O. brunnipes, 228

Orthoptera, 533 ; parasite of, 497

Orthorrhapha, 454 ; O. Brachycera, 478 f.;
O. Nemocera, 455, 458 f.

Oryctes nasicornis, 199

Oscinidae, 504

Osmeterium, 363

Osmia, 47, 48 f.; enemy of, 100; hair of,
11; O. cyanoxantha, 32; O. leucomelana,
29; O. tricornis, 48; O. tridentata, 48 f.

Osten Sacken on Bugonia, 499; on
Chaetotaxy, 446; on classification of
Diptera, 456; summary
schinsky, 512

Othniidae, 265

Otiorhynchides, 437

Ourocnemis, 355, 356

Ovaries, 541, 602; in larva, 325

Oviduct, 320, 321

Oviposition, of Cicada, 571; of Notonecta,
567 ; of Tachina, 507

Ovipositor, 436, 506, 531, 539

|
|

of Port-

Owlet-moths, 414
Ox-warbles, 517
Oxybelus, 129
Oxychirotinae, 425
Ozaenides, 214

.
Pachypus, 187
Packard, on Hymenopterous metamor-
phosis, 7

Pad, of Lepidoptera, 314

Paedogenesis, 3038, 461

Pagiopoda, 544

Palaeolepidoptera, 366

Palaeomicra, 435

Palaeotropinae, 347 ©

Palaeozoic Insects, 311, 542

Palmer-worm, 323

Palpicornia, 279

Paltostoma, 465

Palustra, 377

Pangonia longirostris, 482

Panomoea, 237

Panurgides, 20

Papilio, 359 ; P. ajax, forms of, 335; P.
antimachus, 362; P. zalmoxis 3, .

Papilionidae, 342, 357, 359

Paracelyphus, 505

Paragia, 89

Parallel series in Aphidae, 585

Parandra, 288

Parapompilus gravesii, 105

Paraponera clavata, 172

Paraponyx, 423

Paraptera, 312

Parasites, among ants, 183; of mason-
bee, 43; of larva of Andrena, 26 ; of
Odynerus, 76

Parasitic, bees, 23, 29 f.; Prosopis, 32 ;
Diptera, 507

Parasyscia, 175 n.

Parmula, 501

Parnassius, 342, 362

Parnidae, 219, 243, 255

Parnopes carnea, 4

Parnus, 244

Parthenogenesis, 24, 85, 86, 395, 430, 469,
530, 588, 594

Parthenogenetic young, 139 n.

Passalidae, 192; larva, 192, 262

Passaloecus, 128

Patagia, 311, 312

Pattern, formation of, 335

Patula, 414

Paurometabola, 542

Paussidae, 207, 213

Paussus cephalotes, etc., 214

Pavonia, 350

Pea-weevil, 277

Peacock butterfly, 352

Peal, on sound-producing ant, 156

Peckham, on Fossores, 130 n. :
stinct, 70, 99

on in-

INDEX

Pectinicornia, 194

Pedicinus, 600

Pediculidae, 599

Pediculus capitis, 599, 601; P. melittae,
274; P. vestimenti, 601

Pedilidae, 266

Peduncle, 133, 134

Pegomyia inanis, 79

Pelobiidae, 207

Pelobius tardus, 208

Pelogonidae, 562

Pelopaeus, 110, 112 n.; P. bilineatus,
114; P. laetus, 113, 117; P. madra-
spatanus, 113; P. spirifex, 112

Peltasticta, 245

Peltides, 233

Pemphigus, 589

Pemphredon lugubris, 128

Pemphredonides, 127

Penis, 314

Pentamera, 190

Pentanephria, 466

Pentatoma, 541; P. rufipes, 535

Pentatomidae, 533, 545, 546

Pepsis, 104, 389; P. formosus, 105

Pérez, on bee-parasitism, 32; on Halictus, 24

Pericoma canescens, 470

Pericopinae, 409

Peridrepana, 401

Perilitus, 282

Peripneustic, 450

Peritracheal spaces, 332, 333

Perkins, R.C.L., on bee and acarid sym-
biosis, 70 ; on Odynerus, 76 ; on Prosopis,
2h

Perophora batesi, P. melsheimeri,
sanguinolenta, 379

Perophoridae, 377

Perothopides, 260

Peytoureau, on morphology of abdomen,
313, 314

Phaeism, 837.

Phalacridae, 231

Phalacrocera replicata, 474

Phaloniadae, 427

Phanaeus splendidulus, antenna of, 191

Pharynx, 320

Phaudinae, 392

Pheidole, 165

Pheidologeton laboriosus, 159 ; P. diversus,
167

Phenax, 575

Phengodes hieronymi, 249

Phengodini, 251

Philaenus spumarius, 577

Philanthides, 124 f.

Philanthus apivorus, 127 ; P. triangulum,
125

Phileurus didymus, antenna, 191

Philonthus nitidus, 225

Phlebotomus, 470

Phloea corticata, 545

SHH Clive La

619

Phloeides, 545

Phloeothrips frumentarius, 530

Phobetron pithecium, 403

Phonapate, 246

Phoridae, 494

Phosphaenus hemipterus, 249

Phosphorescence, 250—see also Lumin-
escence

Phragma, 307, 313

Phragmocyttares, 81

Phthanocoris, 543

Phthiriasis. 601

Phthirius inguinalis, 601

Phycitidae, 424

Phycodromidae, 504

Phyllocnistis, 431

Phyllomorpha laciniata, 547, 548

Phyllorhina, parasite of, 520

Phylloxera, 587

Phylogeny of butterflies, 343 n.

Phymatidae, 554

Phymatopus, 399

Physapoda or Physopoda, 531

Phytomyzidae, 504

Phytophaga, 190, 237, 276 f.

Phytophthires, 544

Phytoscopic effects, 336

Pierella, 348

Pieridae, 341, 357

Pieris brassicae, 340 ; development of wing,
328, 329, 333

Piesma, 550

Pigments, 330, 334, 357

Pilifer, 308

Pill-beetles, 242

Pinaridae, 405

Piophilidae, 504

Pipunculidae, 496

Pipunculus, 496

Pison, 118

Pit-falls, 481

Plagiolepis trimeni, 153

Plagithmysus, 287

Plant-lice, 581 f.

Plants and ants, 183

Plataspides, 545

Platycnema, 496

Platypezidae, 496

Platyphora lubbocki, 495

Platypides, 289, 295

Platypsyllidae, 219

Platypsyllus castoris, 219

Platysoma depressum, 230

Platystomidae, 504

Plea minutissima, 567

Pliny, on Mason-bee, 44

Ploiaria pallida, 556

Plume-moths, 426

Plumules, 3351

Plusia, 415

Pochazia, 576

Podilegidae, 32

620

INDEX

Poecilocampa populi, 406

Poecilocapsus lineatus, 542

Poecilocyttares, 81

Pogonomyrmex barbatus, 154

Pogonostoma, 203

Poison, of Hymenoptera, 7 f.

Poison-glands, 2

Poisonous caterpillars, 376, 403, 404, 405

Polistes, 86; P. and Stylops, 301, 303

Pollen-gathering and -carrying, 11, 12

Polybia, nests of, 81, 82, 83

Polyctenes fumarius, 560

Polyctenidae, 560

Polyergus lucidus, 151 ; P. rufescens, 150 f.

Polymorpha (Coleoptera), 789, 190, 213 f.

Polymorphism, 139 f., 143

Polyphylla fullo, antenna, 191

Polyploca, 386

Polyplocidae, 386

Polyplocotes, 248

Polyrhachis, 155; P. pandurus, 156 ; P.
spinigera, 138

Pompilidae, 93, 101 f.

Pompilus, 103; P. polistoides, 104 ;
sericeus, 106

Ponera contracta, P.ergatandria, P.puncta-
tissima, 172

Ponerides, 132, 170 f.

Porphyraspis tristis, 284

Porphyrophora polonica, 597

Porrorhynchus, 216

Portschinsky, on habits and development of
Muscidae, 507, 512

Post-metamorphie growth, 141

Post-scutellum, 307, 312

Potamogeton pectinatus, beetle on, 280

Potamopihilus acuminatus, 244

Potassium hydroxide, 328, 384

Potato-beetle, 278

Pouch, abdominal, 350, 362

Poulton, on colours, 339, 336

Praecostal nervures, 319

Praescutum, 312

Pratt, on imaginal dises, 453 n.; on Jfelo-
phagus, 519

Prepona, larva, 354

Prey, of Fossores, table, 92

Pria dulcamarae, 232

Primitive, beetles, 251, ; Diptera, 475

Priocnemis affinis, 5 ; P. bicolor, 107

Priodont, 193

Prionides, 287

Proboscis, 13, 14, 16, 17, 304, 307, 309,
511, 448, 482, 485, 532

Processional maggots, 464

Processionary caterpillars, 376, 408

Prodoxidae, 432 ; mouth, 309

Prodozus, 433

Pro-legs, 323

Prolific, Aphids,
Hepialus, 397 ;
Stylops, 301

P:

252

ae

589); Coccids, 5
Lice, 601 ; AMeloe, 2

Prominents, 383

Pronuba, 321; P. yuecasella, 432 ; P. syn-
thetica, 432

Pronymph, 453

Propodeum, 131, 133

Propolis, 63

Propygidium, 187

Prosopis, 21, 22; hair of, 11; proboscis
of, 17; P. signata, 21

Protection, 43, 413; of trees by ants,
158 ; of plant by ants, 168

Proterhinidae, 298

Proterhinus lecontei, 298

Protolepidoptera, 356

Protoparce carolina, 309

Protopaussus, 214

Psammorycter vermileo, 481

Pselaphidae, 223 ; and ants, 182

Psen atratus, P. concolor, 127

Psephenus, 244

Pseudholoptic, 440 n.

Pseudocorylophidae, 228

Pseudodicthadia, 180 ; P. incerta, 177

Pseudodoxia limulus, 431

Pseudomeria graeca, 99

Pseudomorphides, 206, 205

Pseudomyrma bicolor, 168

Pseudomyrmini, 168

Pseudoneuroptera, 527

Pseudopaedogenesis, 303

Pseudopod, 188, 264, 267, 290, 449, 492

Pseudopontia paradoxa, 357

Pseudo-pupa, 271, 273

Pseudotetramera, 790

Pseudotrimera, 239

Pseudovespa, 88

Pseudovarium and Pseudovum, 584

Pseudovitellus, 588

Psilidae, 504

Psiliglossa, larva, 8

Psilocephala, 484

Psilura monacha, 407 q

Psithyrus, 53, 57, 59 f.; P. campestris,
60; P. vestalis, 60

Psocidae, 248

Psyche helix, 394

Psychidae, 369, 392

Psychina, 394, 393, 404

Psychoda, 466

Psychodidae, 470

Psylla pyricola, 579, 580; P. succincta,
579; P. buxi, 580

Psyllidae, 578

Pterocheilus, 76

Pterodecta, 400

Pterophoridae, 371, 425, 340, 426

Pterophorinae, 426

Pterostichus, 205

Pterothysanidae, 369, 406

Pterygodes, 312 :

Pterygogenea, 542

Ptilinum, 442, 503, 520

INDEX

621

Ptilinus pectinicornis, 253

Ptilocnemus sidnicus, 557

‘Ptilomacra, 395

Ptilomera laticaudata, 553

Ptinidae, 246

Piomaphila lacrymosa, 222

Ptychoptera, 466

Ptychopterinae, 472

Ptyelus goudoti, 577

Pugs, 411

Pulex avium, P. irritans, P. serraticeps,
525; P. obtusiceps, 525

Pulicidae, 522 f.

Puliciphora lucifera, 495

Pulvillus, 446

Pupa, of beetles, 188 f. ; coarctata, 452 ;
obtecta, 227, 327, 451; depositing
eggs, 469; hairy, 407, 426; of Lepi-
doptera, 326 f.; with mandibles, 436

Puparium, 452

Pupation, of Badamia, 365 ; of Parnassius
and Thais, 363

Pupipara, 456, 513, 517

Purple emperor, 344

Pusci, 504

Puss-moth, 328, 383, 406

Pygidium, 187

Pyralidae, 340, 370, 420

Pyralidina, 420, 426

Pyrameis atalanta, 353; P. cardui, 353 ;
P. tammeamea, 363

Pyraustidae, 421

Pyrochroa rubens, 266

Pyrochroidae, 266

Pyrophorus, 258 ; P. noctilucus, 259

Pyrrhocoridae, 549

Pyrrhocoris apterus, 535, 549

Pyrrhopyge, 364

Pythidae, 265

Pytho depressus, 266

Quartenia, 89
Queen, 66, 67, 69, 140

Races, of Apis, 68

Radial nervures, 319

Raffray, on Pogonostoma, 204 ; on celassi-
fication of Paussidae, 214 ; on classifica-
tion of Pselaphidae, 224

Railway-beetle, 251

Ranatra linearis, 563

Raptorial legs, 493, 554, 556

Réaumur, on Nylocopa, 33

Receptaculum seminis, 140

Rectal cauda, 538

Red admiral, 352

Red ant—see Formica rufa

Reduviidae, 555 f. 537

Reduvius personatus, 558

Régimbart, on Gyrinidae, 216

Resemblance,- between ant, wasp, and
spider, 169; between Anthophora and

Bombus, 33; between Arctia villica
and Husemia villicoides, 410; between
beetle-larva and Termite, 206 ; between
Bombus and Eulema, 35; between Bom-
byliidae and Hymenoptera, 489; between
bug and ant, 556, 557 ; bug and Tipulid,
556, 558 ; bug and fly, 547 ; in butter-
flies, 348; between Callidulidae and
Lycaenidae, 400 ; between Ce/yphus and
Hemiptera, 505 ; between Cleridae and
insects they destroy, 254; between
Dioptinae and Ithomiides, 409 ; between
Diptera and Hymenoptera, 499, 500 ;
between Lpicopeia and Papilio, 418;
between Flatides and Lepidoptera, 576 ;
of flies and bees, 502; to galls, 403 ;
between host and parasite-bees, 30;
between Insects of different Orders, 339 ;
of Ithomiides to other butterflies, 346 ;
between lady-bird and Endomychid, 237 ;
of larva to a colony of larvae, 418;
of larva of Odynerus and of Chrysis, 4 ;
between larvae, 162; in Limacodidae,
401; of Lobster caterpillar, 385 ; between
Longicorns and Hymenoptera, 287 ;
of moth to bird-excrement, 401; of
parasite and prey, 95 ; of Pericopinae and
Heliconiides, 409; between protected
butterflies, 345 ; between Psithyrus and
Bombus, 59; of Reduviid and Pepsis,
558; between Rhyphus- and Mycetobia-
larvae, 463; of Syntomids to other
Insects, 388 f. ; of Tipulids and Hymen-
optera, 475; between two kinds of
ants, 162

Resting-larva, 306

Retinaculum, 316, 319, 420

Retort-shaped bodies, in mouth of Hemi-
ptera, 535

Reuter, E., on classification of butterflies,
345 n.

Rhachicerus, 480

Lthagovelia plumbea, 552

Rhaphiorhynchus, 483

Rhegmatophila alpina, 305, 386

Rheumatobates bergrothi, 553

Rhinomacerides, 291

Rhinopsis, 115; R. ruficornis, 169

Rhinosimus, 266

Rhipicera mystacina, 256
Rhipiceridae, 256

Rhipidiini, 267

Rhipidioptera, 543
Rhipiphoridae, 267
Rhipiphorides, 268

Rhipiptera, 298 [in error for Rhiphiptera]
Rhizophagus, 232

Rhiztrogus, 191

Rhodoneura, 405

Rhogmus, 179, 180
Rhopalocera, 340, 341 f.
Rhopalomelus angusticollis, 206

622

INDEX

Rhopalomeridae, 504

Rhopalosoma poeyi, 100

Rhopalosomides, 70)

Rhygchium brunneum, R. carnaticum, I.
nitidulum, R. oculatum, 77

Rhynchites betulae, 291, 292

Rhynchitides, 291

Rhynchophora, 790, 277, 288 f.

Rhynchophorous series, 240

Rhynchopsylla pulex, 526

Rhynchota, 532

Rhyphidae, 478

Rhyphus, 463

Rhysodidae, 201, 234

Riley, on Hpicauta, 270, 271: on sperma-
thecal bodies, 321; on Yucca-moth,
432

Ripidius pectinicornis, 269

Rippe, of Schiffer, 334

Robber-flies, 491

Rose-chafers, 200

Rosema, 401

Rostrum, 291, 472

Rothney, on Ampulex, 115 ;
169 ; on Sphex, 110

Rothschild, N. C., on morphology of fleas,
523 n.

Royal jelly, 66

Rozites gongylophora, 167

Ruby-wasps, 1 f.

Rurales, 342

Rutelides, 295, 198

on Sima,

Saccoderes, 558; S. tuberculatus, 537

daccus, 314

Sagra splendida, 279

Sagrides, 279

Salda, 560

Saldidae, 544, 562

Salivary, duct, 320; gland, 320

Sand-flea, 525 ; -tlies, 477

Saperda populnea, 285, 286

Sapromyzidae, 504

Sapyga quinquepunctata, 100

Sapygides, 99

Sarcophaga, 510; S. carnaria, 510

Sarcophagidae, 510

Sarcophila magnifica, S. wohlfahrti, 510

Sarcopsylla gallinacea, S. penetrans, 525

Sarginae, 479

Saropoda, 32

Sarrothripus, 410

Sasaki, on parasite of silkworm, 508

Saturnia, 307, 310; S. pavonia, 313, 374

Saturniidae, 368, 372

Satyrides, 347

Sauba, or Sauva, ant, 137

Saunders, E., on feathered hairs, 11; on
proboscis of bees, 16

Saunders, Sir 8., on Hymenopterous larvae,
8

Scale, 131, 133, 315, 467 ; development

iy BPN) Gee
331

Scale-Insects, 592 f. ; enemy of, 356, 357,
417, 425, 430

Scalpella, 443

Scape, 441

Scaphidiidae, 229

Scaphisoma agaricinum, 229

Scaphium, 314

Scapulae, 312

Scarabaeidae, 194 f.

Scarabaeini, 196

Scarabaeus sacer, 196, 187

Scatomyzidae, 504

Scatophagidae, 504

Scatopse, 477

Sceliphron, 112; S. nigripes, 91

Scenopinidae, 484

Schaffer, on structure of wings and uer-
vures, 330

Schiddte, on Anoplura, 599 f.; on Hemi-
ptera, 543

Schistocerca peregrina, enemy of, 506, 514

Schizocarpus mingaudi, 220

-formation, 333; -holder,

Schizometopa, 504 .

Schizoneura, 589

Schizophora, 455

Schmidt-Schwedt, on Donacia, 280

Schoenbergia, 360, 361

Schoenobiinae, 425

Sciapteron, 387

Sciara militaris, 464

Sciomyzidae, 504

Sciophila unimaculata, 462

Scolia bifasciata, S. haemorrhoidalis, 97

Scoliidae, 93, 94 f.

Scoliides, 97 f.

Scolytidae, 294

Scopariidae, 421

Scopariinae, 421

Scopelodes, 401

Scopulipedes, 20, 32 f.

Screw-worm, 512

Scudder, on butterflies, 543

Scutata or Scutati, 546

Scutellerides, 545

Scutelligera, 501

Scutellum, 307, 312, 537

Scutum, 307; S. proboscidis, 443

Scydmaenidae, 223

Seymnites, 238

Scymnus minimus, 238

| Seasonal dimorphism or variation, 335

| Seitz, on Syntomid resemblances, 388, 389
Semi-loopers, 415
Seminal duct, 321

| Semi-pupa, 271

| Semper, on development of wing, 333

| Sense-organ, 442, 448; thoracico-abdo-

| minal, 414 ; in Uraniidae, 419

| Sepsidae, 504

| Sericaria mori, 375

INDEX

623

Sericteria, 325

Seroot-fly, 482

Serricornia, 789, 213, 255

Sesia, 387 ; S. scoliaeformis, 321

Sesiidae, 386, 388, 370 (for Syntomidae),
389

Setae, 534 f. ; aérostatic, 408

Setina, 410

Seventeen-year Cicada, 569

Sex, differences, 92, 95; production of,
32, 67 ; in larva, 325

Sexuparous, 586

Sharp, on classification of Dytiscidae, 213 n.

Sheep, bot-fly, 517 ; -tick, 518

Shield, 592

Shoulder, -lappets, 312; -tufts, 312; of
wing, 319, 316

Siagona, 206

Siculodidae, 423

Siebold, von, on Strepsiptera, 301

Sigara minutissima, 568

Sight, of Pompilus, 104

Silk-glands, 325

Silkworm, 375; affected by parasitic fly,
507, 508 ; Madagascar-, 405

Silpha, 221 ; Silpha atrata, S. laevigata,
S. lapponica, 222; S. obscura, 222; 8.
opaca, S. thoracica, 222

Silphidae, 221 f., 252, 256

Sima, 168 ; 8S. leviceps, stridulating organ,
169; S. rufonigra, 169.

Simuliidae, 477

Simulium columbaczense, 477

Sinodendron cylindricum, 194

Siphon, 563, 581, 589

Siphonaptera, 522 f.

Siphonophora, 239

Siphunculata, 600

Sitaris, 33; S. humeralis, 272

Sitodrepa, 247

Skippers, 363

Slave-making ant, 149, 150, 163

Sloth, 430

Slug-worms, 402

Smallest Insect, 228

Smerinthini, 380

Smerinthus populi, 309, 381

Smith, F., on Mellinus arvensis, 123

Snails, enemies of, 205, 222, 510 ; parasite
of, 495

Social, bees, 35 ; wasps, 78, 84

Sociales, 20, 53 f.

Solanuin dulcamara, beetle on, 232

Soldiers, 132 :

Soldier-ant, 150

Solenobia, 395, 430

Solenopsis fugax, 187

Solitary wasps—see Fossores and Eume-
nidae

Song, of Cicada, 572

Soronia, 23:

|

|
Sound-organs, 448 ; of Ageronia, 354; of |

Hecatesia, 371; of Cicada, 573, 574—
see also stridulating orgaus

Sound-production, 155, 156 ; by Aegocera,
411; by Arctiidae, 410 ; by Sphingida:,
382—see also Stridulation

Spalacopsini, 288

Spatula, 459

Spencer, Herbert, on Weismann, 143

Spercheus emarginatus, 218, 219

Spermatheca, 320, 321

Spermophila, 506

Sphaeridiides, 2279

Sphaeriidae, 227

| Sphaerites, 223

Sphaeritides, 229

Sphaerius acaroides, 227

Sphaerocarides, 279

Sphecia, 387

Sphecius speciosus, 123

Sphecodes, 21, 22; S. gibbus, 23, 23; S.
rubicundus, 22; S. subquadratus, 23

Sphegidae, 93, 107 f.

Sphegides, 107 f.

Sphex coeruleus, 110; S. flavipennis, 108 ;
S. lobatus, 110; S. maxillosus, 108

Sphindidae, 245

Sphingidae, 309, 315, 316, 368, 380 f.

Sphing ligustri, 380

Spider parasite, 490

Spilosoma, 408

Spinneret, 324, 325, 403, 417

Spirachtha, 227

Spiracles, 188, 191; of Diptera, 449 f. ;
of Hippoboscidae, 519 ; of Lepidoptera,
313, 314; of Lipara, 451; of Nepa,
564 ; of Thrips, 528

Spondyliaspis, 581 *

Spondylidae, 288

Springing plant-lice, 579

Spuler, on nervures, 317 n.

Squama, 448

Squeakers, 209

Staetherinia, 401

Stag-beetles, 193

Stalk, 317, 319

Staphylinidae, 223, 224 f.

Staudinger, Schatz and Rober, on butter-
flies, 343 n.

Stauronotus maroccanus, 489

Stauropus fagi, 385

Stelis minuta, 29; S. nasuta, 30, 43; 8.
signata, 30

Stelocyttares, 81

Stenamma westwoodi, 159

Stenopteryx hirundinis, 519

| Stephostethus, 240

Sternorhyncha, 544
Sterrhopteryz, 394
Stethopathidae, 496
Stigmata—see Spiracles
Stigmatomina, 180
Stigmus pendulus, 128

624

INDEX

Sting, 4, 5, 6, 58, 144; development of, |
8,9

Stinging, 98; by Calicurgus, 102; by
Pompilidae, 104; by Sphex, 109

Stingless bees, 61

Stink-gland, 257, 533 ; -vessel, 225

Stipes, 309

Stizinae, 123

Stomach, 320

Stomoxys calcitrans, 512

Stratiomyidae, 478

Stratiomys, 452 ;

Straus-Durckheim, on Melolontha, 198 |

Strawberries, eaten by beetles, 205

Streblidae, 521

Strepsiptera, 789, 298 f. |

Streptoperas, 401

Stridulating organ, of Myrmica, 133 ; of
Heterocerus, 243; of Passalus-larva,
192 ; of Sima leviceps, 169

Stridulation, of ants, 134 ; of Corixa, 568 ;
of Criocerides, 281 ; of Dynastides, 199;
of Geotrupes, 195 ; of Ipides, 232; of
Lomaptera, 200; of Longicorns, 287 ; of
larva of Lucanus cervus, 194 ; of Mega-
lopides, 282 ; of Melolontha-larva, 198 ;
of Mutilla, 94; of Pelobius, 208; of
Phonapate, 246 ; of Phyllomorpha, 548 ;
of Praogena, 264 ; of Siagona, 206 ; of |
Sino, W8)s)

Strigil, 568

Striphnopterygidae, 376

Strongylognathus huberi, S. testaceus, 162

Strumigenys, 170

Style, 442

Stylopidae, 298

Stylopised bees, 26

Stylops dalii, 299

Styx infernalis, 358, 340

Suana, 405

Subcostal nervure, 318

Submedian nervure, 318

Suboesophageal ganglion, 320

Sucking-stomach, 311, 449

Suction by Lepidoptera, 311

Suctoria, 526

Supericornia, 546

Swallow-flies, 519

Swarming of wasps, 70 n.

Swarms, 62, 65 67, 80, 135, 467, 505,
584

Swift-moths, 396

Symbiosis, of ants and plants, 139; of |
bee and Acarid, 70. See also Ants’-
nest Insects, and Association

Symbius blattarum, 269

Symphily, 183

Synecthry, 183

Synemon, 371

Synoeca cyanea, nest, 82

Syntelia westwoodi, 229

Synteliidae, 229

Syntomidae, 339 n., 369, 388
Syntomis phegea, 390
Syringe, 535, 536

Syrphidae, 439, 498 f.
Systoechus oreas, 489
Systropus crudelis, 489

Tabanidae, 481, 492 P

Tabanus, 482

Tachinidae, 507, 514

Tachysphex panzeri, 117

Tuchytes, 116; destroyer of, 275: 7.
australis, 1138, 117 ; T. pectinipes, 117

Tachytides, 116

Taenia, fleas as hosts of, 526

Tajuria diaeus, pupa, 357

Taleporia, 395

Taleporiidae, 430

Tanypezidae, 504

Taphroderides, 296

Tapinoma erraticum, 157

Tarantula-killer 105

Tarphius, 233

Tarsolepis, 383

Taschenberg, on anatomy of flea, 523 n.

Tascina, 372

Tea-plant bug, 562

Teara melanosticta, 408

Tegula, 71, 187, 307, 311, 312, 447

Tegmina, 539

Teleodont, 193

Telephorides, 248

Telmatophilus, 235

Temnochila coerulea, 232

Temnochilides, 233

Tenebrio molitor, 263

Tenebrionidae, 263

Tenebroides mauritanica, 232

Tentacle, maxillary, 309, 432

Tenthredinidae, parasite of, 4

Terebrantia, 531

Termites, 203, 206, 227, 231

Terrifying attitude, 384

Tesseratomides, 546

Testes, 321, 324, 400, 429 ; in larva, 325

Tetanocera ferrugined, 504

Tetanoceridae, 504

Tetragona, 53, 61

Tetramera, 190

Tetramorium caespitum, 160, 163

Tettigometrides, 567

Teucrium, bug and galls on, 550

Thais, pupation of, 363

Thanaos, 342 ; T. tages, androconia, 332

Therevidae, 484

Thiridopteryx, 420

Thomas, on androconia, 331

Thorictidae, 236

Thorictus, 236

Thrips, 526 f.; Thrips
secalina, 530

(ify SB 2 ie

/ Throscides, 260

INDEX

Thyatira batis, T. derasa, 386

Thymaridae, 392

Thynnides, 96

Thyreophoridae, 504

Thyrididae, 570, 404

Thyridopteryx ephemeraeformis, 394

Thysanoptera, 526 f.

Tiger-beetles, 201 f.

Tiger-moths, 409

Tillus elongatus, 253, 254

Tinaegeriidae, 370, 387

Tinea, 305; T. pellionella, 429, 430; 7.
vastella, 430 ; T. vivipara, 430

Tineidae, 340, 370, 394, 427, 428

Tineodidae, 423

Tineola biselliella, 430

Tingidae, 549

Tipula brobdignagia, 475

Tipulidae, 471 f.; T. Brevipalpi, 472, 473 ;
T. Longipalpi, 472, 475

Tipulinae, 475

Tiresias serra, 241

Titanus giganteus, 287

Tithorea, 346

Tomicides, 295

Tomognathus sublaevis, 161

Tongue, 309

Tortoise-shell butterflies, 352

Tortricidae, 340, 395, 427, 432

Tortricina, 395

Toxorrhina, 472

Toxotrypana, 506

Trechus, 205

Trichiini, 200

Trichocera, 473

Trichodes alvearius, T. ammios, T. apiarius,
254

Trichophaga tapetzella, 430

Trichoptera, 306, 425

Trichopterygidae, 227

Trichopteryx fascicularis, 227

Trichroism, 351

Trichterwickler, 294

Trichura, 389, 390

Trictenotomidae, 275

T-iecphora, 543

T igona, 53, 61; T. carbonaria, 63; T.
crassipes, 65; T. mosquito, 62

Trimera, 238, 544

Trimeria, 89

Trineura aterrima, 494

Triodites mus, 489

Trioza rhamni, 580

Triphaena, 415

Triphleps, 580

Tritoma bipustulata, 236

Triungulin, 262, 268, 270, 271, 272, 299,
300

Trochalopoda, 543, 544

Trochanter, 307 ; divided, 123

Trochilium, 387

Trogini, 795

VOL. VI

Trogositidae, 232, 235
Trogosita mauritanica, 232
Tromoptera, 457
Trophi—see Mouth-parts
Troz, stridulation, 195
Truftle-beetle, 222
Trumpeter bumble-bee, 58
Trypanaeus, 230
Trypanidae, 395
Trypetidae, 504, 506
Trypoxylonides, 118
Trypozylon, 118; TZ. albitarse, 118; T.
figulus, 119
Tse-tse fly, 512, 513
Tubulifera, 7 f.
Tubulifera (Thrips), 531
Tulip-tree, tubes on, 578
Turkey-gnats, 477
Turnip-flea, 278
Tusser, or Tussore, silk, 374
Tympanoterpes gigas, 572
Typhlatta, 179, 180
Typhlopone, 178, 179, 180

Ugimyia sericariae, 507
Ulidiidae, 504

Ulopa, 578

Uncus, 314

Urania rhipheus, 419
Uraniidae, 368, 419
Uric acid pigments, 357
Urodon, 278

Uzel, on Thysanoptera, 527

Vanessa, 352; larva, 854 —see also
Pyrameis and Araschnia

Vanessula, 256

Vapourer-moths, 407

Variation, of Anomma burmeisteri, 179 ;
of Bombus, 58; of larvae, 336; of
Sphecodes, 23; of male and worker
ants, 160; of workers and females,
162; due to parasites, 26; of larva
and imago, 408 ; generic, 401 ; local,
398 ; in nervuration, 414 ; and dimor-
phism in Geometrid-larvae, 412 ; of
mandibles of Lucanidae, 193 ; in colour
of Psyllidae, 579 ; trichroism of hind
wings, 351 ; in size of Brenthidae, 297 ;
of time and form in Cicada, 570; in
wings, 540; as to winged or wingless,
531; change in, 414; seasonal, 335

Vasa deferentia, 321

Veils, 493

Veins—see nervures

Velia currens, 552

Velleius dilatatus, 227

Verhoeff, on Agenia, 106; on Halictus,
25; on Siphonophora, 239; on Stedis
minuta, 29; on terminal segments of
beetles, 186

Vermileo degeeri, 481

626

Vermipsylla alakurt, 523, 526°

Verson, on rudiments of wings, 328

Vertebrates, larvae of Diptera, attacking,
HOC DLO Hane St Silasw olin mo20:
tick-fleas on, 526

Vespa, nests of, 79, 83; V. austriaca, 81,
88 ; V. crabro, 81; V. germanica, 79

Vespidae, 78

Viviparous, Aphids, 583 ; fly, 506, 511,
513, 518 f.; moths, 430 ; Staphylinidae,
227

Voice—see Song, Sound-organs, Stridula-
tion

Volucella, 500; V. bombylans, 441

Wagner, on morphology of fleas, 523 n. ;
on paedogenesis, 460

Walker, J. J., on Halobates, 552

Wallace, on flight of Hesperiidae, 364

Walsingham, Lord, on Tortricidae, 427

Walter, on mouth of Lepidoptera, 308,
310

Wandering ants, 175 f.

Wanzenspritze, 536

Wasmann, on Ants’-nest Insects, 181 n.,
183; on Lomechusa, 142, 226; on
Weismann, 143

Wasps, 71 f.

Wasps’-nest, beetle, 235 ; Insect, 268

Water-scorpion, 563

Wax, 65, 570, 5/6, 597

Wax-glands, 589

Wax-hairs, 580

Wedde, on mouth of Hemiptera, 535

Weeping-trees, 577

Weevil, biscuit-, 247; pea-, 277

Weinland, on halteres, 448

Wet- and dry-season forms, 336

Whirligig-beetle, 215

White wax, 576, 597

Whittell, on Pelopaews and Larrada, 117

Wielowiejski, on luminous organs, 250

Wing-cases, of beetles, 186, 270

END OF VOL.

INDEX

Wing, of bugs, 539; of Diptera, 447 ;
of Lepidoptera, 315 f. ; development of,
328 ; structure of, 329

Wingless—see Apterous

Wingless and winged Aphids, 584

Wing-nervures—see Nervures

Wing-rib, 330, 353

Wing-veins—see Nervures

Winter-gnats, 473

Winter-moth, 414

Winter-mother, 586

Wire-worm, 258

Wood-ant—see Formica rufa

Wood-leopard moth, 309, 395

Woodpecker, Diptera in, 506

Workers, 54, 66, 67, 79, 85, 132, 140

Worm-eaten furniture, 248

NXantharpyia straminea, parasite of, 521,
522

Xenos, 303 ; X. rossti,

NXestobiuin, 248

Xylocopa, 32, 34, 70 ; submentum of, 14 ;
X. chloroptera, 34; X. violacea, 33

Xylodiplosis, 458, 459

Xylophagidae, 479

Xylophaginae, 480

Xylophilidae, 266

NXylotrupes gideon, 199

299, 301

Yellow-fever-fly, 464
Yolinus, 558

Young carried, 556
Yucca-moth, 432

Zabrus, 205

Zaitha anura, 566

Zelotypia staceyi, 396

Zemioses celtis, 296

Zeuzera aesculi, 309, 395
Zeuzeridae, 395

Zygaenidae, 369, 388, 390, 392, 394
Zygia, 253

VI

Printed by R. & R. Crark, Limitep, Edinburgh.

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