LIBRARY

OF Til E

JOHNS HOPKINS UNIVERSITY

COLLECTION
OF TH E

WILLIAM H . WELCH MEDICAL LIBRARY

Anatomical drawings of biting flies

Boris Jobling

Boris Jobling just before his retirement in 1958.

Anatotnical drawings of biting flies

Boris Jobling

Prepared for publication by David J. Lewis

Department of Entomology

British Museum (Natural History)

produced in association with the Wellcome Trust

Contents
Introduction
The artist

7
9

The Jobling collection

10

The species illustrated here

Abbreviations

11

12

The drawings
The sandfly- Phlebotomus papatasi
(Figures 1-113)
The mosquito - Aedes aegypti
(Figures 114-195)
The blackfly - Simulium
(Figures 196---239)

49

69

The deerfly - Chrysops caecutiens

(Figures 240-295)
The stablefly - Stomoxys calcitrans
(Figures 296-356)
References 117

17

81
101

Acknowledgements
Sincere thanks are due to the following. Dr A. J. Duggan facilitated Jobling's later work, conserved
his collection, and made the drawings available for publication. Dr L. A. Mound, Keeper of
Entomology, British Museum (Natural History), provided working facilities for D.J.L., supported
the proposal to publish this work, and provided editorial expertise. Mr Jobling and his son
Mr Edwin Jobling gave much information, and Dr L. G. Goodwin, C.M.G., F.R.S., Honorary
Director of the Wellcome Museum in 1984, provided facilities there. Dr R. P. Lane made many
helpful suggestions. Mr Jobling, while in Egypt, received much help from the late Dr H . Hoogstraal,
American Naval Research Unit No. 3, Mr H. Lomas, Wellcome Foundation Office in Cairo, and Mrs
Lomas.

Introduction
The late Boris Jobling was employed by the Wellcome Organization and made numerous insect and
other drawings of high scientific and artistic quality. Of these, 356 were not published, and are
reproduced here to make them widely available and to draw attention to the whole Jobling
collection of specimens and drawings in the Wellcome Museum of Medical Science, London, which
is now administered by the Wellcome Trust (1985).
Failing eye-sight prevented Jobling from completing a text for the drawings, and in 1981 Dr A. ].
Duggan, then Director of the Museum, consulted me about them . I offered to prepare them for
publication and provide an introductory text. The drawings alone, with occasional reference to
Richards & Davies (1977) or similar text-books on entomology, serve as an anatomical atlas of these
representative species of biting flies. Specialised publications are cited for further information.
The drawings are useful for teaching and illustrate aspects of anatomy, histology, physiology,
taxonomy and medical entomology. Regarding the last, some of the sensilla, for instance, guide a fly
to its host, which may be a reservoir of disease. The structure of the mouth parts determines the
nature and depth of a wound, and thus influences the uptake and subsequent deposition of
pathogenic organisms. The cibarium and pharynx are part of the parasites' route into and out of a
fly. Saliva, secreted by glands, is essential for the feeding process and for transmission of some
parasites, and often causes irritation to man. The crop of sandflies stores sugar which is necessary
for the development of Leishmania. In the alimentary canal some parasites are greatly influenced by
the nature of the epithelium, and by the peritrophic membrane (Gemetchu, 1974; Le Berre, 1967)
which prevents many from developing . In some sandflies an abundant abdominal fat-body is an
indicator of autogeny. The female reproductive organs are used in the basic estimation of physiological age in relation to disease.
One may well ask what is the place of art in a scientific work. Lapage (1961) gives the answer and
points out, inter alia, that some objects depicted already have an element of beauty, that scientific
preparation precedes the artist's work, and that he must exercise his talent without sacrificing
accuracy. Jobling has achieved a vivid realism which calls attention to the often obscure objects
portrayed.

Digitization
I rst became aware of Joblings drawings several years ago during a seminar at Johns Hopkins
University in Baltimore, Maryland. Despite having worked in the medical entomology eld for a
number of years, I had not previously encountered an adequate presentation of the anatomical
features of hematophagous insects. Indeed, it was a common source of frustration in the lab when we
would seek to identify an internal anatomical structure, most often in the context of a marked
transgenic mosquito strain, that we would ultimately be resigned to make inference from the
homologous features of a model insect whose anatomy was thoroughly described and readily
available. Generally, this meant making comparisons with the fruit y Drosophila melanogaster, which
was inadequate not only because it is not hematophagous, but also because its most recent ancestor
shared with mosquitoes is estimated to have lived more than 250 mya.
It was therefore with great excitement that I obtained a copy of this work from The William H.
Welch medical library to use during the preparation of my doctoral dissertation. Joblings
illustrations proved so useful that I sought to acquire my own copy of the book, only to nd out that
it was long out of print. Retaining copies of these beautiful sketches for my own reference was
suciently important that I decided to digitize the entire work myself manually. This electronic
document, representing more than 80 hours of work, is the result of that process.
Each page of the original book, including the front and back covers, was scanned at 300 dpi as an
8-bit grayscale TIFF. The content in each of these 127 images was individually aligned and centered
on an 8.5 x 11 inch page, taking care to retain the scale of the gures on each page, although the
original page size was A4. The text on each page was recognized electronically and made
searachable. The light gray background noise of the scanned images was removed manually during
processing, except in cases where it would aect the texture of the drawing. This resulted in an
interesting eect on the processed drawings that I like to think slightly enhances their
three-dimensional appearance.
It is my hope that converting these drawings to a modern, digital format will aid in their
dissemination among researchers old and new as well as anyone who would not otherwise have had
the opportunity to experience such truly extraordinary works of art.
John R. Clayton
October 2012

Boris Jobling at work in 1955.

The artist
Boris Jobling's grandfather John was born about 1834, served in the Royal Navy, and became a
railway engineer near Newcastle. He was a Chartist, and the unpopularity of the movement led him
to emigrate to Russia. He married in 1865 and his son Edwin John became a mechanical engineer
and married Agripina, a Russian who became British by marriage. Boris was born at Gorbatka,
260 km east of Moscow, . on 26 December 1893. He studied science and art at the Rial secondary
school in the neighbouring town of Murom, and from 1915 to 1919 entomology and protozoology at
the Shaniavsky University, Moscow. He then taught natural history at his old school and became an
assistant at the Freshwater Biological Station on the River Oka. He was imprisoned briefly in the
aftermath of the revolution, and in 1920 he and the late C. A. Hoare (Garnham, 1984) were
repatriated. Boris crossed the Russian frontier into Finland and travelled to the port of Hanko where
he met Hoare, and Ada Braithwaite whom he was to marry in 1925. They sailed in the S.S. Dongola
which was sent by the British Government to fetch refugees, and reached England in March. Boris
lived in the Mitcham Institute for Refugees and later in a rooming house in Charleville Road,
Kensington. He died at Guildford on 26 April 1986 and is survived by his wife and a son and
daughter.
Jobling worked as a private artist for Hoare, and his skill was noticed by the late C. M. Wenyon
who was preparing his great work on protozoology. Jobling was taken on as a temporary artist in
1921 by the Wellcome Bureau of Scientific Research which had been founded in 1913 (Duggan,
1981), and in 1923 was appointed to the staff as entomologist by Sir Henry Wellcome. The Bureau
was a unique etablishment, a scientific institute devoted to fundamental research on problems
connected with tropical medicine and parasitology (Goodwin & Bruce-Chwatt, 1986). Jobling
worked at Wellcome's Entomological Research Station at Wisley, later at Claremont, Esher, and
during the Second World War moved to the Wellcome Research Institution in Euston Road,
London. In the 1950s he visited Egypt to study Phlebotomus papatasi. He retired from the service of
the Wellcome Foundation Limited on 31 December 1958 and became a Fellow of the Institute of
Biology in 1966.
He published 23 papers between 1925 and 1940, 13 between 1947 and 1959, and one in 1976. Most
were on Streblidae (parasites of bats), but other subjects were Phlebotomus (1976), Culicidae (1935a,
1936, 1938b), Culicoides (1928a, 1953), Muscidae (1935b), Glossinidae (1933), Hippoboscidae (1926),
Nycteribiidae (1928b), the tick Ornithodoros (1925) and supports for cover-glasses (1938a). The tick
paper and the first one on Culicoides were described respectively as a 'well illustrated paper' and a
'finely illustrated anatomical study' by Alcock (1925, 1928). Buxton (1955: 51), in his book on the
tsetse flies, used Jobling's 'detailed and beautifully illustrated work' on the structure of the feeding
apparatus.
Jobling contributed illustrations of Protozoa for several publications. Wenyon (1926) referred to
his 'knowledge of biology combined with his artistic skill' and many original drawings. Hoare (1949)
thanked his colleague for coloured plates, and (1972) 'was fortunate in having the assistance of my
friend Mr B. Jobling, F.l. Biol., whose artistic skill and knowledge of the subject ensured faithful
execution of the original illustrations and the reproduction of numerous figures from other sources'.
Jobling painted malaria parasites for James (1929: 273) and James, Nicol & Shute (1938: pl. 11), and
for plate 2 of Shute & Maryon (1960) which was copied for Wilcocks & Manson-Bahr (1972: 959). He
also illustrated Orthoptera for B. P. Uvarov.
Goodwin & Bruce-Chwatt (1986) described Jobling as 'one of the very finest entomological artists
in the classical style'.

The Jobling collection

This is in the Wellcome Museum of Medical Science, 183 Euston Road, London NW1 2BP.

General morphology of the head in Diptera

There are 87 drawings, in nine groups, and 55 typed pages of unpublished text on sutures and the
tentorium, and on mouth parts. The drawings include the antennae of Pangonia and Tabanus. The
term sialophore is proposed for the "hypopharynx" of Diptera, which Jobling regards as the
modified dorsal wall of the labium.

Drawings of the morphology of biting flies

These comprise many already published, such as 57 in his 1928 paper on the head of Culicoides, and
the originals of those reproduced here.

Microscope slides
These include thousands of sections used for the drawings.

Protozoa
There are hundreds of drawings, including many for Wenyon's work (1926), the plates of which
were lost in a fire.

The species illustrated here

The drawings were made without a camera lucida and are based on numerous measurements. They
are reproduced here without reduction, as originally intended. In many of them cytological detail is
shown by the difficult use of dots as described by Cannon (1936). No less than 474 parts of the
insects are labelled. For each species the drawings, or groups of them, were numbered, and those in
a group lettered A, B etc. All are now numbered consecutively, but the groups are kept and
indicated by their sets of captions. The scale lines are all in millimetres.
For all species the descriptions of anatomy by Gordon & Lavoipierre (1962) and external anatomy
by Colless & McAlpine (1970), McAlpine (1981) and Matsuda (1965, 1970, 1976) are useful. C. T.
Lewis (1972) and Mciver (1982) discussed sensilla, Downes (1971) and Snodgrass (1944) reviewed
trophic structure, and Detinova (1962) described age-grouping. The species illustrated here represent a wide range of Diptera and of feeding mechanisms. Phlebotomus, Aedes and Simulium are in the
Nematocera, Chrysops is in the Brachycera, and Stomoxys in the Cyclorrhapha. Phlebotomus, Simulium
and Chrysops are pool feeders, Aedes is a capillary feeder, and Stomoxys a rasping pool feeder.
Phlebotomus papatasi (Scopoli) (Psychodidae; Figs 1-113) is an important vector of Oriental sore
(cutaneous leishmaniasis), and an example of the genus which includes vectors of several forms of
Leishmania. Sandfly anatomy was described by Davis (1967; some terminology not widely acepted),
Just (1973), authors mentioned by them, Jobling (1976) and Rudin & Hecker (1982), and external
features by Abonnenc (1972), Forattini (1973), D. J. Lewis (1973, 1975, 1982), Perfil'ev (1968),
Theodor (1958) and others. Magnarelli, Modi & Tesh (1984) studied ovarian changes, and KillickKendrick (1979) discussed leishmania! development in relation to structure of the gut.
Aedes aegypti (L.) (Culicidae; Figs 114-195) is the well known vector of urban yellow fever, and a
major vector of dengue. Christophers (1960) wrote a comprehensive book on the species, Clements
(1963) dealt with features of histology and physiology, and Harbach & Knight (1980) described the
external anatomy of mosquitoes.
Simulium (Wilhelmia) sp. (Simuliidae; Figs 196--239) may comprise more than one species. The
figures, originally labelled S. equinum L., were drawn before it was known that the related S.
lineatum Meigen differs from it in having a sub-ovoid spermatheca (Crosskey, 1981: 147) very like
that shown in Figs 214, 229 and 235. The head of a Wilhelmia was studied by Wenk (1962). One of
these species bites horses in Britain (Edwards, Oldroyd & Smart, 1939), and S. lineatum attacks cattle
in Europe. The simuliids or black-flies include notorious man-biters in many countries, and vectors
of Onchocerca volvulus Leuckart, the cause of river blindness (Crosskey, in press; Laird, 1981) in
Africa and tropical America.
Chrysops caecutiens L. (Tabanidae; Figs 240--295) is a common European man-biting species
(Edwards, Oldroyd & Smart, 1939) and is related to vectors of human loiasis in Africa (Oldroyd,
1957; Service, 1980). The biting process in Chrysops has been described by Gordon & Crewe (1953),
and aspects of anatomy by Lavoipierre (1958).
Stomoxys calcitrans L. (Muscidae; Figs 298-356), the stable fly, is a widespread biting pest of man
and animals (Greenberg, 1971; Service, 1980). Its subfamily has been reviewed by Zumpt (1973), and
its peritrophic membrane studied by Lehane (1976).

11

Abbreviations
Notes
Sial: this is the hypopharynx (of Diptera) of most authors .
Ven: used here for the abdominal part of the mid-gut
9 stn: sternite 9 of male P. papatasi. The coxite arises, not from this, but from the phallobase, according to Just
(1973).
10 stn: sternite 10 of the same. This is sternite 9, according to Just (1973).
Various: in some cases other authors have used different names.
a
abd. gng
abd. nv
abd. spr
ac. gl
adv
aed
aed. p
af
air. s
al. m
amp
an
ant. ar. t
ant-I. t
ant. thor. spr
ant. t. p
ant. w
a. nv
a or
aor. sin
ap
api. eel
api . snl
a. pul. org.
ar
asc
as. nur
a. tr
a. ves
ax . aps
ax. fb
axn

antenna
abdominal ganglia
abdominal nerve
abdominal spiracles
accessory glands
adventitia (pericardium)
aedeagus
aedeagal pouch
antennifer
air sac
alary muscles
ampulla
anus
anterior arm of tentorium
antero-lateral trachea
anterior thoracic spiracle
anterior tentorial pit
anterior wall
antenna! nerve
aorta
aortic sinus
apodeme
apical cell
apical sensilla
antenna! pulsatory organ
arista
ascoids
association neurone
antenna! trachea
antenna! blood vessel
axial apophysis
axial fibre of sense cell
axon

bld . eel
b . mem
br
bs . pl
bs. snl
buc
burs

blood cells or haemocytes

basement membrane
brain
basal plate
basiconic sensilla
bucca
bursa copulatrix

car
cb

cardia
cibarium

cb. lm. tr
cb. pmp
cd
cer
cg. eel
cho
cir. m
cl
cl. ap
de. chn
elf
clge. sut
cl . nv
clsp
cl. sut
clx
cmd. e
em. ovd
cmp. snl
em. sl. d
comp. m
en
cnn. stk
con
cor
cox
cp. cl
cr
ern
crp . al
crp. card
crp . card. nv
cv. con
cv. nv
cv . sc
ex. nv
ex. 1, 2, 3

cibario-labral trachea
cibarial pump
cardo
cerci
corneagen cell
chorion
circular muscles
clypeus
clypeal apodeme
collecting channel
cleft
clypeogenal suture
clypeal nerve
claspette
clypeal suture
calyx
compound eye
common oviduct
campaniform sensilla
common salivary duct
compressor muscle
canal
connecting stalk
connective
corneal lens
coxite
cap cell
crop
cornua
corpus allatum
corpus cardiacum
corpus cardiacum nerve
cervical connective
cervical nerve
cervical sclerite
coxal nerve
coxa of fore, middle and hind legs

d
d. ac. gl
d. cr
dr. ar. t
dr. arh sg
dr. com

duct
duct of accessory gland
duct of crop
dorsal arm of tentorium
dorsal arch of sigma
dorsal commissure

12

ABBREVIATIONS

dr.
dr.
dr.
dr.
dr.
dr.
dr.
dr.
dr.
dr.
dr.

dorsal cephalic trunk

c. trn
dorsal diaphragm
dphr
dorsal diverticulum
dv
long. tg . m dorsal longitudinal tergal muscle
dorsal mesothoracic nerve
mes. nv
met. nv
dorsal metathoracic nerve
nv
dorsal nerve
dorsal prothoracic nerve
prt. nv
dorsal sinus
sin
dorsal trunk
trn
w . lm
dorsal wall of labrum

ej. d
en. eel
epi
epi. eel
epid
eps
e. rig
e. sut

ejaculatory duct
enveloping cell
epithelium
epithelial cell
epidermis
episternum
epistomal ridge
epistomal suture

f
fa)
fat . b
f-cl
fd. en
fgl. sut
f. gng
f. gng. con
fl
fie
fig
fol
fol. epi
frc
frk
fun

frons
falciform process
fat body
fronto-clypeus
food canal
frontogenal suture
frontal ganglion
frontal ganglion connective
filament
fulcrum
flagellum
follicle
follicular epithelium
furca
fork
funicle

ga
ge. rd
gen. ch
gen . fl
gen. frc
gli
gls
gl. vs . sem
gon
grm
grm. eel
gs cae
gs. nv
gu
gus. snl

galea
genal rod
genital chamber or atrium
genital filament
genital furca
glia
glossa
gland of vesicular seminalis
gonopore
germarium
germ cells
gastric caecum
gastric nerve
gula
gustatory sensilla

h
hin-g

haematin
hind-gut

hist. m
hl. tr
hphy
ht
hy
hyp . gng

histolysed muscles
haltere trachea
hypopharynx
heart
hyoid
hypocerebral ganglion

i. eel
il
in. pre
ins
int
in. tbr
in. to
intrb . chn
in. w
ir. pg. eel

interstitial cells
ileum
inner process
insula
intima
inner tuberosity of mandible
inner tooth
interbifid channel
inner wall
iris pigmented cells

J.

Johnston organ

kl

keel

1. abd. trn
larv. m
lb
lb. g
lbl
lbl. cls
lbl. frc
lbl. gl
lbl. pre
lb. mem
lb. nv
lb. pis . org
lb. tr
lc
le
lg. nv
lg. tr
lig
1. lob

lateral abdominal trunk

larval muscles
labium
labial gutter
Ia bellum
labellar canaliculus
labellar furca
labellar glands
interlabellar process
labial membrane
labial nerve
labial pulsatory organ
labial trachea
lacinia
lever
leg nerve
leg trachea
ligament
lateral lobe of tenth abdominal
segment
lateral longitudinal tergal muscle
labrum
labral nerve
labral trachea
premento-labellar and mandibulogenal links
lobe of eighth sternite
longitudinal dorsal muscle
longitudinal muscle
longitudinal sternal muscle
loral arm
lateral oviduct
lip

1. long. tg. m
lm
lm. nv
lm. tr
Ink
log. 8 stn
long. dr. m
long. m
long. stn. m
lor. ar
1. ovd
lp

13

ABBREVIATIONS

1. pr
1. thor. trn
lv
1. w. lm

lateral process of labial gutter

lateral thoracic trunk
laterovertex
lateral wall of labrum

m:m. tro

muscles
muscle of trochanter
levator of scape, in some abductor of
same
depressor of scape, in some
abductor
retractor of scape; in some may be
abductor
extensor of pedicel
rotator of pedicel
flexor of pedicel
tormo-labral muscle
anterior labral muscle
adductor of mandible
abductor of mandible
tentorial adductor of mandible
tentorio-cardinal muscle or
protractor of maxilla
tentorio-stipital muscle or retractor
of stipes
cranial flexor of lacinia or in some
lacinia retractor of same
geno-stipital muscles or diagonal
adductor of stipes
stipito-palpal muscle or levator of
maxillary palpus; in some may be
abductor of same or retractor of
maxilla
depressor of maxillary palpus, in
some abductor
levator of second palpal segment
depressor of second palpal segment
retractor of prementum, or labium
abductor of first segment of labellum
adductor of first segment of
labellum, in some flexor
abductor of second segment of
labellum, in some extensor
anterior cibarial dilator
posterior cibarial dilator
retractor of cibarial pump
protractor of cibarial pump, in some
depressor
sphincter buccalis
dilator buccalis
precerebral dilator of pharyngeal
pump
postcerebral dorsal dilator of
pharyngeal pump
posterior lateral dilator of
pharyngeal pump

1
2
3
4
5
6
7
8
9
10
11

12
13

14
15
16

17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32

33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50-53

salivary pump muscle

levator of head
compressor muscle of sac of
pompetta
protractor of genital filament
depressor of ninth abdominal
segment
levator of tenth abdominal segment
adductor of style
abductor of style
rotator of style
levator of coxite
abductor of coxite
adductor of coxite
abductor of claspette, in some
depressor
adductor of claspette, in some
levator
depressor of aedeagus, in some
retractor
levator of aedeagus
depressor of claspette
muscles of tenth abdominal segment

Mal. tu
md
md. end
md. max. tr
md. nv
mdv
mem
mes. gng
met. gng
mid . cr. sut
mid-g
mit
msn . inv
mt
mtr. nur
mx
mx. gl
mx. lob
mx. nv
mx. pip

Malpighian tubes
mandible
mandibular condyle
mandibulo-maxillary trachea
mandibular nerve
mediovertex
membrane
mesothoracic ganglion
metathoracic ganglion
midcranial suture
mid-gut
mitosis
mesenteric invagination
mentum
motor neuron
maxilla
maxillary glands
maxillary lobe
maxillary nerve
maxillary palpus

n
n. epi . eel
nephr
nlm
nr. eel
ns. eel
nv
nv. cr
nv. hi
nv. lam

nucleus
nucleus of epithelial cell
nephrocytes of epicardial cells
neurilemma
nurse cells
neurosecretory cells
nerve
nerve to crop
nerve to haltere
neural lamella

14

ABBREVIATIONS

nv. sl. d
nv. wng

nerve to salivary duct

nerve to wing

oc
oc. end
oc. for
ocl. m
oclr. sc
oclr. sut
oclr. tr
oc. sut
oe
oen
olf. p
olf. snl
ooc
oog
oos
op
op. ac. gl
op. sp . d
opt. lob
opt. nv
ost
ost. vl
out. chia
out. pre
out. w
ov
ovl
ovl. ped
ovl. sh
ov. sh

ocelli
occiput
occipital condyle
occipital foramen
occlusor muscle
ocular sclerite
ocular suture
ocular trachea
occipital suture
oesophagus
oenocytes
olfactory pits
olfactory sensilla
oocyte
oogonium
oosome
openings of ducts
opening of accessory gland
opening of spermathecal duct
optic lobe
optic nerve
ostia
ostial valve
outer chiasma
outer process of mandible
outer wall
ovary
ovariole
ovariole pedicel
ovariole sheath
ovarian sheath

ped
peg
pen
pg. eel
pge
pge. brg
pgen. p
pg. gr
pgl
ph
ph. pmp
phr
plf
pls. mem
pls. or
pm
pmt
poe
poe. sut
poe. rig

pedicel
peg
penniculus
pigment cell
postgena
postgenal bridge
postgenital plate
pigment granules
paraglossa
pharynx
pharyngeal pump
phragma
palpifer
pulsatory membrane
pulsatory organ
pompetta
postmentum
pas tocci pi tal
postoccipital suture
postoccipital ridge

pof. sut
ppd
pre
pre . w. lm
prg
prit. mem
prmt
prn
prnv
prot
prp
prs
pr. sh
prst
prst. sc
prst. te
prt. gng
pst. ar. t
pst-1. trn
pstn
pstr
pstr. rng
pstr. slt
pstr. te
pst. t. p
pst. w
pt
ptl. blad
ptr
pt. sut
pv
pyl
pyl. sph

postfrontal suture
postpedicel
process
preoral wall of labrum
proctiger
peritrophic membrane
prementum
pronotum
perineurium
protuberance
periopticon or lamina ganglionaris
prescutum
peritoneal sheath
prestomium
prestomal sclerite
prestomal teeth
prothoracic ganglion
posterior arm of tentorium
postero-lateral trunk
postnotum or postscutellum
pseudo trachea
pseudotracheal rings
pseudotracheal slit
pseudotracheal teeth
poterior tentorial pit
posterior wall
ptilinum
petiolate blades
peritreme
ptilinal suture
postvertex
pylorus
pyloric sphincter muscle

r
rad. str
rb
rd
rec. nv
rect
rect. p
ref
reg. eel
rem. fol. epi
res
ret. eel
ret. pg eel
rig
rst. mem

ring
radial striation
rabdom
rod
recurrent nerve
rectum
rectal papilla
refractive spheres
regenerative cells
remainder of follicular epithelium
reservoir
retinal or sense cells
retinal pigment cells
ridge
rostrum membrane

sarc
sc
sci
sci. org
sco

sarcosomes
sclerite
scolopophores
scolopophorous organ
scolops

15

ABBREVIATIONS

scp
sctl
sctl. pis . org.
scut
si
sial
sl. en
sl. d
sl. gl
sl. pmp
smt
snl
sns. eel
sns. or
sns. p
so
sam. eel
sp
spat
spd
sp. d
spg
sp. gl
sph. m
spi
spm
s. pm
spr
spr. atr
spr. tr
spt
sr. blad
SIC

sr. rig
st
stl
stom. inv
sub. gng
sus . sc
t
tb
t. brg
teg. nv
ten.
tes
tes. sh
tg-stn . m
th
thor. gng
tn. 9
tn . 10

scraper
scutellum
scutellar pulsatory organ
scutum
siphon
sialophore (hypopharynx)
salivary canal
salivary duct
salivary glands
salivary pump
submentum
sensilla
sense cells
sense organ
sensory pits
sockets of sensilla
somatic cells
spermatheca
spatulate rods
spermatids
spermathecal duct
spermatogonia
spermathecal glands
sphincter muscle
spicules
spermatozoa
sac of pompetta
spiracles
spiracular atrium
spiracular trachea
spermatocytes
serrated blades
sarcostyles or myofibrils
serrated ridges
stipes
style
stomodaeal invagination
suboesophageal ganglion
suspensory sclerite
tentorium
tibia
tentorial bridge
tegumental nerve
taenidium
testis
testicular sheath
tergo-sternal muscle
labial theca
thoracic ganglion
tendon of adductor muscle of
mandible
tendon of abductor muscle of
mandible

tn. 11
tor
tr
trh
tr. 1, 2, 3
tro. 1, 2, 3
tun. pr
tu . sh
v
vas.
vas .
vbr
ven
vii
vl
vnt.
vnt.
vnt.
vnt.
vnt.
vnt.

tendon of tentorial adductor of

mandible
torma
trachea
tracheoles
tracheae of fore, middle and hind
legs
trochanters of fore, middle and hind
legs
tunica propria
tubular sheath

vs. sem
vst

vertex
vas deferens
vas eferens
vibrissae
ventriculus
villarium
valve
ventral arch of sigma
ventral cephalic trunk
ventral commissure
ventral diaphragm
ventral nerve
ventral nerve cord
vesicle
vesicular seminalis
vestige

w. aor
wng. tr

wall of aorta
wing trachea

yolk

2. abd . gng
6. abd. gng
7 & 8. abd.
gng

second abdominal ganglion

sixth abdominal ganglion

de
ef

arh. sg
c. trn
com
dphr
nv
nv. crd

VS

10 abd. s
1. br
2. br
3. br
1. nv
2. nv
1. s. lb. plp
2. s. lb. plp
1. s. lbl
2. s. lbl
1. s. max. plp
1-10 stn
1-10 tg

16

fused seventh and eighth abdominal

ganglion
tenth abdominal segment
protocerebrum
deu tocerebrum
tri tocerebrum
first lateral nerve
second lateral nerve
first segment of labial palpus
second segment of labial palpus
first segment of labellum
second segment of labellum
first segment of maxillary palpus
sternites
tergites

The Sandfly

Phlebotomus papatasi

17

oc
pof.sut
'

oclr.sut
oclr.sc

poc.su
t
I
~_poe

- -oc.cnd

prot

_pst.t.p

'pge

scp--ant.t.p

'cd

'-e.sut
'

asc _
'md
---- - lm
----- --m x'

th

- prmt

I s./b/ /

2s./b l/
2

.3

Figs 1, 2 Phlebotomus papatasi,

S?; anteri or and poster ior views of head.

19

''

'st

'

asc -- _

.05

........ bs.snl

trg.cel
'

II

a.nv
'

'

'\

sn's.cel

pdc, ...

5
Figs 3-7 Phlebotomus papatasi, ~. 3, antenna;
4, terminal segment of flagellum; 5, lateral
section of ascoid; 6, dorsal view of cuticular
process of ascoid; 7, basiconic sensillum.

scp...-

, api .snl
''

_- oc.end

pof.sut
',

~ ~ ~poe.rig

'

_-oclr.se

', ed
I

,:n
pdc.,

't
'

- st

I
I

-- ~ pmt
/Jre.w.lm

tor-

tor . . .

md.end

pre--

md' -

.05

,sus.sc

prmt

mx' '

10

- ant.w

sns.p
''
---1 s.lbl

ern

--2s.lb/
.2
/
I

I
I

ph.pmP
\
\

12

9
0

a
0

'

0
('

(\

,',lb
c

Figs 8-13 Phlebotomus papatasi, 2 except Fig 12. 8, lateral view of head; 9, dorsal view of labrum; 10, ventral
view of distal part of labrum; 11, posterior part of dorsal wall of pharyngeal pump; 12, lateral view of anterior
part of head of cf; 13, spiculate scales from pharyngeal wall.

', lm
.1

"pre.w.lm
sial
/

,dr.w.lm

.1
/

14

sl.pmp
/

I
I

in.tbr

'

'

'

tor

/ ant.w

'

.A/
\

md.cnd,

\
\

~xsus.sc

_tn. 9

15

\cb.pmp

'

cm.sl.'d
cmp.sn/
I

/or.ar

__ @
.1

md.cnd-

\
\
\

I
/

tn./1

16
I

17
/

tn .9'
oe,

tn.l 0

s/.cn,.'"

18

19

.05
Figs 14-19 Phlebotomus papatasi, <(. 14, dorsal view of left mandible; 15, dorsal view of sialophore and salivary
canal and pump; 16, section through mandibular articulation; 17, dorsal view of proximal part of left mandible
and its articulation with gena; 18, distal part of sialophore; 19, lateral view of sialophore.

lm

gus .sn/
,

md \

lb .'g

lb/.prc

23

lb .nv

l.w.lm
lc
I

sl.cn
_
pre .w.lm
04

, 16

24
/

tc,

,dr.w.lm
/.w.lm
I

th

.1

-- st

lb.g

.04

.05

sial

/
lb.tr

22

th/

rtg

25

' 23

lb.nv,
20
mx.gf/
21

26

Figs 20--26 Phlebotomus papatasi, ~. 20, dorsal view of right maxilla; 21, distal part of lacinia; 22, dorsal view of
labium and part of labial gutter; 23--25, transverse sections of second and first segments of labella and middle
part of proboscis; 26, transverse section of proximal part of lacinia.

. . 30
/

e.sut

e.rig \

...

29

26

br

'

31

30

29
~-rx
~-rx

e.rig
\

cb .pmp

25 \

24,
I

'

'

'

lm.nv . . .

'
sub.gng

sl.pmp ,
I

''

cm.sl.d

sJ.gJ

33

27

20

' .nv
lb

prt.gng
.2

j.o
3

21
pdc-"

br

28

4 -- --

lbl

5----2',
27,
'
ant.t.p..._ ' ' '
'

11-- .. .....

md.....
lc

'

9
/
/

.... '

'

-- -- 32

18
.....

''
19/

- ls.mx.plp

'

15

I
I
I

'I

'

' pst.t.p

''

'20

cd

Figs 27-29 Phlebotomus papatasi, 9. 27, longitudinal section of middle part of head and anterior part of thorax;
28, same of right half of head; 29, muscles of posterior part of cibarial pump.
24

 tor',

31

scp,

sl.pmp

26

crp.card .nv- --

'

_ -rec.nv
vnt.c.trn

- - fat.b--

,.

3br"
.1

32

33

rec.nv
lbr

vnt.c.trn

...

aor
31

.05

Figs 30-35 Phlebotomus papatasi, ~; transverse sections. 30, basal parts of labrum, mandibles, sialophore,
laciniae and labium; 31, 32, anterior and middle parts of head; 33, 34, posterior parts of pharyngeal pump and
head; 35, oesophagus, corpus cardiacum, hypocerebral ganglion and aorta.
25

f.gng

cl.nv

24
/

29
28

''

rec.nv

..,..30

ph.pmp

'

, .3br
\

N
0\

lm
',,

\''

'

sial

'

.. ~ v
\

cm.sl.d

'

'

''

\
\

mx.nv

''

'pmt
'
Fig. 36 Phlebotomus papatasi,

2; nervous system of right half of anterior part of head.

'
md.nv

nv.sl.d

pstn
I

hin-g
f

rect

Jtg
/

I
1

\ sl.gl
' ph.pmp thor .gng

cbpmp_

\
\

Ql

OV

6abd .gng

Mal.tu

'sub.gng

cr

I
I

ac.gl
8tg

' tro.2

cm.sl.d

7+8abd .gng

sl.pmp

Fig. 37 Phlebotomus papatasi,

<?; general organisation showing natural position of internal organs.

sp

ph.pmp

~,

30

- oe

I
I

I
I
I
I

mid-g
I

cr

ven

Mal.tu

Fig. 38 Phlebotomus papatasi,

28

~;

alimentary canal.

41

car
'
I

aor
w.aor
'

reg.ce/
I

cir.m
''
b.memven

43
''long.m

'

'oe

cir.m
\

d.cr

42

long.m
I

46

c1r'.m- . . . -

pr.sh

45

prit.mem
44

prit.mem

""

47

.8

Figs 39-47 Phlebotomus papatasi, s>; histology of crop and mid-gut. 39, transverse section of duct of crop; 40,
longitudinal section of empty crop; 41, surface view of stretched wall of same showing thick circular muscles
and thin longitudinal ones; 42, longitudinal section of oesophagus; 43, transverse section of thoracic part of
mid-gut; 44, surface view of wall of same showing disposition of longitudinal and circular muscles; 45,
longitudinal section of abdominal part of mid-gut; 46, 47, the same a few, and about 16 hours, after feeding.

29

-.... msn.inv

vii
'fong.m
j --

I
I

Mal.tu

49
48

pr.sh

51

,pg.gr
/

l~ng.m

50

53

.05

52

b.mem"

Figs 48--53 Phlebotomus papatasi, S?; histology of hind-gut and of Malpighian tubes. 48, posterior part of midgut and anterior part of hind-gut; 49, connection of Malpighian tube with mid-gut and hind-gut and
continuation of villarium into Malpighian tube; 50, 51 , transverse and longitudinal sections of ileum; 52, 53,
longitudinal and transverse sections of middle part of Malpighian tube.
30

Stg
\

ht

6tg

prg

'
7stn

/
/

oen

,
I

6stn

'

'
lob.B.stn

54

.1

7+8abd.gng

long.stn.m

op.sp.d

9stn

55

/ob.B.stn

Figs 54, 55 Phlebotomus papatasi,

S? . 54,

longitudinal section of posterior end of abdomen; 55, transverse

section of ninth abdominal segment.
31

a.nv

md.nv
mx.nv"
.1

rec.nv

57

'
crp.card
.nv

Figs 56, 57 Phlebotomus papatasi, <(. 56, right half of nervous system of head; 57, ventral view of same.

32

.. mtr.nur

mes.gng'

58

e>---------

b.mem
cm.sl.d

.1

60

62

Figs 58-62 Phlebotomus papatasi, 2. 58, nervous system of thorax in lateral view; 59, section of prothoracic
ganglion; 60, salivary glands; 61, 62, sections of right and left glands.

.02

63
nv./am,

'

rec.nv" "

64

ns.cel

mid-g ...

65

bid. sin

.05

crp.card

crp.a/

''

rec.nv

hyp.gng"

phpmp

mid-g

66

d.cr"""'

Figs 63--66 Phlebotomus papatasi, 9. 63, neurosecretory cells; 64, cells of corpus cardiacum; 65, diagram of
dorsal view of stomatogastric nervous system and associated endocrine glands; 66, right half of same showing
histology of glands and their connection with brain and stomatogastric nervous system.
34

2stn

67

long.stn.m
,.
cv.mem

Figs 67-69 Phlebotomus papatasi, <;?. 67, four scolopophorous organs of second abdominal segment, which
control position of abdomen in relation to thorax; 68, nerves and sense organs of basal part of wing; 69,
scolopophorous organ of head.

35

fat.b

1./ong.tg.m

70

.02

75

72

Figs 70-75 Phlebotomus papatasi, S?. 70, transverse section through dorsal half of third abdominal segment; 71,
nephrocytes; 72, part of heart showing its internal structure and attachment to its wall of alary muscles; 73,
blood cells, one of them attached to histolysed larval muscle; 74, 75, nephrocytes of unfed female and after
oviposition.

36

I
I

oc/r.sc

lbr
rec.nv--

76

ph pmp--

pdc

11

78

77

.1

79

cor

irpg.ce/ __

ret. pg.cel-

,cg.ce/

--~----11>.
. . ,.

----~}

.05

,%~r[)J

-ret.cel

b.mem,

out.chia

prp80

Figs 76-80 Phlebotomus papatasi, S? . 76, transverse section of anterior part of head showing antenna! pulsatory
organ which consists of vesicle and two muscular pulsatory membranes; 77, longitudinal section of first
antenna! segment; 78, transverse section of tubular part of pulsatory membrane near vesicle; 79, pulsatory
organ of scutellum; 80, ommatidium of light-adapted compound eye.
37

cb-lm.tr' "

81

Figs 81-84 Phlebotomus papatasi, 2. 81, tracheal system of head in dorsal view; 82, dorsal view of origin of
dorsal and ventral cephalic tracheal trunks from lateral thoracic trunks from near anterior thoracic spiracles;
83, 84, anterior and posterior thoracic spiracles.

/.thor. trn

86
.04

.3

l.abd.trn

85

Figs 85, 86 Phlebotomus papatasi,

S?.

85, tracheal system of right half of thorax and of abdomen; 86, abdominal
spiracle.

39

, lig

.3

ov/...-''

...

'ac.g/
lob.B.stn
op.sp.d

'
op.ac.g 1

Fig. 87 Phlebotomus papatasi, Cj?; reproductive organs, tracheal system not shown.

40

lig

'sp.gl

91

b.mem
~

...

. oog

m
.05

nr.cel

92

ovl

'

\
\

fol ,,

89

90

Figs 88--93 Phlebotomus papatasi, ~; histology of internal reproductive organs. 88, 89, longitudinal sections of
spermatheca and ovary; 90, ovariole with one developing follicle containing seven nurse cells and one oocyte,
and in germarium five dividing oogonia; 91, end of calyx and beginning of lateral oviduct; 92, section of wall of
accessory gland; 93, musculature of ovarian sheath.
41

8tg
,
\Otg
cer

8stn

7stn

"

/ob:8.stn

94
.1

op.sp .d
9tg . .

95

Figs 94--96 Phlebotomus papatasi, <j?; genitalia. 94, lateral view of posterior end of abdomen; 95, dorsal view of
part of eighth segment; 96, ventral view of ninth and tenth segments.

42

fl

vas.def

99

rd

,35
/

97

.1

ej.d

35' ....

~~

.....

s.pm,

tu.sh-

98

gen.{/ "

.05
.03

100

101

gon

,. ,.

,.

102

Figs 97-102 Phlebotomus papatasi, cf' . 97, reproductive organs; 98, section of pompetta showing inner sac and
opening of ejaculatory duct; 99, horizontal section of same showing compressor muscles of sac and protractors
of pompetta and genital filaments; 100-102, genital filaments.

spm

api.cel
''

- pr.sh

mi

b.mem

spd

ej.d,

''

'

b.mem

m,

pr.sh, ,
epi"

104
.05

105

103

Figs 103-105 Phlebotomus papatasi, d; histology of testis and vesicula seminalis. 103, longitudinal section of
testis; 104, longitudinal section of vesicula seminalis and its glands, each extending from vas deferens, and
latter with narrow canal full of spermatozoa; 105, transverse section of ejaculatory duct.
44

7tg
/

.1

c/sp
-1 Ostn

9tg~

108
.05

109

107

Figs 106-110 Phlebotomus papatasi, cf; genitalia. 106, end of abdomen after torsion; 107, ventral view of ninth
abdominal segment; 108, tenth abdominal segment; 109, longitudinal section of tactile sensillum of style; 110,
penniculus.

45

39
'

41

stl

.2
I

8tg

aed

' 46

aed-113

50

.05

Figs 111-113 Phlebotomus papatasi, d'; musculature of genitalia . 111, inner view of right half of ninth and tenth
abdominal segments and of right coxite; 112, transverse section of tenth abdominal segment; 113, same
through basal parts of coxites.

46

The Mosquito

Aedes aegypti

47

_ mid-cr.sut

clge.sut
'

115

.3

mx.plp
c/ge .sut
,

lc

--- th

116

pge -_./bl
114

.3

oe

cv.sc
117

Figs 114-117 Aedes aegypti, S?. 114, anterior view of head and proboscis; 115, lateral view of anterior part
head showing connection of anterior arm of tentorium with ciypeus, position of torma, and connect!on
cibarial pump by ciypeal apodeme with clypeus; 116, lateral view of head, left compound eye removed
show position of pharyngeal pump; 117, middle part of ventral wall of head showing internal position
stipites, occipital foramen and fused postgenae of which anterior parts form postgenal bridge.

of
of
to
of

--I

-~?J-- op
{

'

.05

- lm

sial--l.w.lm-' dr.w./m
.5
.1

lm
.1

cl

cl.ap- __ _
---- sl.pmp

33
118

''cm.sl.d

ph.pmp---

119

122

121

buc

Figs 118-122 Aedes aegypti, ~. 118, posterior and anterior parts of sialophore; 119, lateral view of clypeus,
cibarial pump, pharyngeal pump, sialophore and labrum; 120, ventral view of anterior part of labrum; 121,
dorsal view of cibarial pump and posterior part of labrum, showing attachment of clypeal apodemes to lateral
margins of cibarial pump, and sense organs of anterior wall of latter; 122, scale-like processes of posterior
parts of walls of pharyngeal pump .

lbl.prc

125

dr.w.lm

21 ,,,

123
.05
.1

126
cl
.05

.1

7
I s.mx.plp

fb.nv
.nv

124
128

---lb.tr

127

Figs 123-128 Aedes aegypti, 2. 123, anterior part of prementum and two-segmented Iabella, left Ia bellum
showing ventral wall; 124, internal structure of same, showing three pairs of muscles and their attachment to
segments of Iabella, labial nerves and tracheae; 125, transverse section of first segment of Iabella; 126, S3me of
anterior part of proboscis; 127, same of middle part of proboscis showing position of syn trophium (combination of mouth parts) in labial gutter; 128, same of anterior part of clypeus and base of mouth parts.
'

- ... ge
----16
130

sns. org
/

\
~

sub.gng ....
ph pmp,.,

.1

aor

132

Figs 129-133 Aedes aegypti, 9. 129, transverse section of anterior part of head; 130, same of posteriL1r end of
cibarial pump and of anterior part of bucca, showing origin of protractor muscles of cibarial pump on
tentorium and their insertion together with retractor muscles of the same on cornua; also showing origin of
salivary pump muscles on loral arms; 131, sense organ (?) of posterior part of antenna! pulsatory organ; 132,
section of posterior part of pharyngeal pump and its circular sphincter muscle; 133, section of posterior part of
head showing position of pharyngeal pump and insertion on its plates of postcerebral dorsal dilators and
posterior lateral dilators.

31

cl .nv

f.gng

'\, 29 ... _ '

Im.nv'

fd.cn
\

cv.con

' sl.d

em~

/b.nv sl.pmp

/b.g

th

Fig. 134 Aedes aegypti,

S?;

.3

right half of head showing sucking apparatus comprising food canal of labrum,
cibarial and pharyngeal pumps, and their muscles .

52

bs.snl

.05

137

'
.05

st-

27

,,

12 ..

136
135

Figs 135-137 Aedes aegypti, S?. 135, distal end of mandible in dorso-lateral view; 136, musculature of mandible
and maxilla; small drawing shows articulation of mandible by mandibulo-genallink; 137, distal end of lacinia.
53

scut
'

long .dr .m

scut/

fat .b
ht
mid-g
I

7tg

Mal.tu
.1

hin~g

vnt.dphr

'

7f.Sabd.gng

Fig. 138 Aedes aegypti, <j?; general organisation .

2/

' sf.pmp

ten

141
139

sl.d ,...

b.mem

.05
Figs 139-143 Aedes aegypti, 2. 139, salivary gland immediately after emergence of mosquito from pupa; 140,
salivary glands of two-day-old mosquito, and longitudinal and transverse sections of common salivary canal;
141, section of middle gland showing intracellular duct and its supporting disc; 142, 143, longitudinal and
transverse sections of lateral gland.

...........

146

'

.05

.2

n.epi .eel

c:r

~-

\
\

mid-g

I'

.1

''

.05
I

int

'

hin.g ~ _

----... -- ... ___

144

...._-

------an

Figs 144-147 Aedes aegypti, S!. 144, alimentary canal; 145, diagram of dorsal diverticulum showing thick
circular and thin longitudinal muscles and very thin longitudinal folds in wall; 146, wall of ventral diverticulum or crop; 147, section of walls of oesophagus and crop.
56

oe

148

,' '

cir m

pr.sh

d.cr'
.1

''

...
,

pr.sh

c1r.m

~il

' epi

'

long.m'

'
reg.cel
150

149

151

____ .vii

et>i

152

cr

153
Figs 148-153 Aedes aegypti, ~; histology of mid-gut. 148, median longitudinal section of anterior part; 149
longitudinal section of abdominal part, or ventriculus, of unfed mosquito; 150, section of narrow thoracic part
fixed during feeding; 151, same of posterior part of ventriculus; 152, middle part of ventriculus of fully fed
female, showing strongly flattened epithelium and villarium; 153, same 24 hours after feeding, showing
peritrophic membrane.
1

ven

-- -~pyl.sph
b.mem
j

Mal.tu ---

.05

epi '

___ /ong.m

-----. hin.g------- ---

.05

155

154

.05

157

rad.str --

epi .eel
long.m

int----

b.mem _.,

pr.sh
,,

158

;'

long.'m

160
, epi

159

long.m

cir.m

Figs 154-161 Aedes aegypti, <j>; histology of hind-gut and Malpighian tubes . 154, posterior part of mid-gut and
anterior part of ileum; 155, same from a different angle, showing origin of Malpighian tube from mid-gut and
the same structure of their epithelial cells; 156, 157, longitudinal and transverse sections of middle part of
Malpighian tube of unfed female; 158, longitudinal section of colon and its short narrow posterior part, in both
of which epithelial cells lack spicules; 159, longitudinal section of thick part of colon (see Fig. 171); 150, 161,
transverse and longitudinal sections of posterior part of rectum.

.05

aus
'tg -stn .m

~I.

thor .trn

163
cir.m
'

b.mem'
.5

164
.5
- -tro.l

162
Figs 162-164 Aedes aegypti, 2. 162, slightly oblique transverse section of anterior part of thorax, showing
position of salivary glands, oesophagus and corpora alata; 163, corpora alata on sides of aorta; 164, transverse
section of oesophagus .

59

. crp.card
crp.al

aor

.r

0'0

oe

suH.gng

.2

13/
Fig. 165 Aedes aegypti,

9; stomatogastric nervou s system and endocrine organs.

.3

'
I

6stn

', vntdphr

vnt.nv.ct d

166

.05

167

\
\

al.m

.05
.03

168

dr.dphr

169
Figs 166-170 Aedes aegypti, S? . 166, transverse section of sixth abdominal segment showing position of heart
and attachment of its alary muscles to intersegmental membrane, etc.; 167, part of heart with its cavity
exposed to show ostia and ostial valve, dorsal diaphragm and nephrocyte among muscles and elastic tissue;
168, section of ventral diaphragm; 169, four haemocytes from surface of mid-gut and three from ovaries; 170,
oenocytes in abdominal fat body.

.2

.8tg

p'gen .p

J?;f
oen
\
I

7+8 abd .gng

8stn

' fat.b

'
7stn
Fig . 171 Aedes aegyp ti,

2; long itudi nal section of posterior end of abdo men .

173

tr

'.l.ovd

8 abd.gng
.3

cm.ovd--

.05

ac.gl -pgen. P
pr.sh"

., .,

174

'rect

175

.3

IOstn. __

gen.ch/
177

Figs 172-177 Aedes aegypti. 2. 172, reproductive organs of eight-hour-old female; 173, transverse section of
accessory or collaterial gland; 174, longitudinal section of duct of same; 175, longitudinal section of distal end
of bursa copulatrix; 176, posterior end of abdomen; 177, ventral view of same.

.05

'' ::-- -fl

'\

.1

cho
ns.cef. __

~-ov.sh

ooc --

179

fol.epi ----

y ..
180

Figs 178-180 Aedes aegypti, 2. 178, surface view of ovarian sheath, showing its branching and anastomozing
muscles; 179, part of ovary of just-emerged female; 180, ovariole with germarium and three follicles .

64

181

182

__.sp.gl

.03

d.ac.gl

I
I

.05

long.m
I

cm.ovd

gon

184

.05

Figs 181-185 Aedes aegypti, S?. 181, section of spermatheca and its duct; 182, spermathecal gland; 183,
transverse section of spermathecal ducts, surrounded by strong longitudinal muscles; 184, longitudinal
median section of posterior end of abdomen, showing migration of spermatozoa from bursa copulatrix
through spermathecal ducts into spermathecae after copulation; 185, transverse section of common oviduct,
showing sphincter muscles and two dilator muscles arising on eighth sternite.

ac.gl

187

vs.s

fat.b
\

186

.1

aed .. --int

188

pr.sh

',

''

spm --

-b'.mem
',m

.05

189
190

Figs 186--190 Aedes aegypti, d. 186, reproductive organs showing two testes, one with fat body removed to
show division into apartments; 187, transverse section of accessory glands and vesicula seminalis, its lumen
filled with spermatozoa; 188, longitudinal section of wall of vesicula seminalis; 189, same of accessory glands;
190: same of vesicula seminalis and accessory gland, showing opening of latter.

\.

fl ----- -,

,, grm .eel
spg

'

.1

spt

m t----

spd

tes .sh

fat .b-

spm

Fig. 191 Aedes aegypti, d; longitudinal section of testis showing its division into apartments or cysts.

192

~I sp

stl '

.3

194

, ej.d

.05

Figs 192-195 Aedes aegypti, cf. 192, lateral view of genitalia of Aedes sp., left coxite removed; 193, ventral view
of same of A. aegypti; 194, parts of tenth abdominal segment; 195, right half of aedeagus showing its
connection with membrane and ejaculatory duct.

68

The Blackfly
Simulium

69

.1

197

Figs 196, 197 Simulium (Wi/he/mia) sp .,

2.

196, anterior view of head; 197, tentorium of same.

70

198

.2

.05

.15

202

Figs 198-204 Simulium (Wilhelmia) sp ., S?; mouth parts. 198, labrum; 199, left mandible; 200, right maxilla; 201,
lateral view of anterior end of labrum showing position of scraper; 202, right half of labium showing labial
muscles; 203, ventral view of labium; 204, dorsal view of sialophore, pharynx and oesophageal pump.

.1

205

206

.1

209
st. pmp

.05
.05

.1

Figs 20~211 Simulium (Wilhelmia) sp., 2. 205, transverse section through first segment of labella; 206, same
through middle part of proboscis; 207, same through oesophageal pump; 208, same through anterior part of
salivary pump; 209, same through anterior part of pharynx and posterior part of salivary pump; 210, section of
salivary glands; 211, same of reservoir of .salivary glands.
72

.2

Figs 212-213 Simulium (Wilhelmia) sp ., 2; sections of head showing internal structure and musculature. 212,
longitudinal, through middle part; 213, oblique transverse.

73

1.0

Fig. 214 Simulium (Wilhelmia) sp .,

2; right half of body showing natural position of internal organs .

Fig. 215 Simulium (Wi/helmia) sp.,

75

2; alimentary system.

card
. gng

216

217

crp. a/

218

Figs 216--219 Simulium (Wilhelmia) sp., <j?; hypocerebral ganglion, corpus alia tum and oenocytes. 216, transverse section through anterior part of hypocerebral ganglion; 217, through middle part of same; 218, section
through corpus allatum; 219, group of oenocytes from posterior part of abdomen.

.1

220

222
.05

.1

221
.05

224

223

Figs 220-224 Simulium (Wilhelmia) sp., S?; histology of alimentary system. 220, longitudinal section of anterior
part of mid-gut; 221 , 222, transverse and longitudinal sections of wall of mid-gut; 223, 224, longitudinal and
transverse sections of hind-gut (ileum).

77

.1

.1

226

225

.1

9tg

Figs 225--228 Simulium (Wilhelmia) sp., <j?; histology of pylorus, Malpighian tubes and rectum. 225, longitudinal section of pylorus and mesentarium; 226, 227, transverse section of Malpighian tubes; 228, median
longitudinal section of posterior part of abdomen showing genital opening of common oviduct.
78

230

.03
.05

232

.05

Figs 229-234 Simulium (Wilhelmia) sp., S?; reproductive organs. 229, general structure; 230, transverse section
of duct of accessory gland; 231, section of spermathecal duct; 232, section of accessory gland; 233, longitudinal
section of spermatheca; 234, ovariole showing its attachment to muscular sheath, and different stages of
development of follicles.

.05

: .:..

~stl

236

235

238
b.mem
m

!.~--cir.

Figs 235-239 Simulium (Wilhelmia) sp., d; reproductive organs. 235, general structure; 236, longitudinal
section of testis; 237, transverse section of vasa differentia and vesicula seminalis; 238, section of wall of
vesicula seminalis; 239, median longitudinal section of posterior end of abdomen, showing genital opening,
structure of ductus ejaculatorius, etc.

The Deerfly

Chrysops caecutiens

81

pv
1.0
I

oc

00
N

lm

' 240

241

fbi

242

Figs 240-242 Chrysops caecutiens, <(; anterior, lateral and posterior views of head , in Fig. 241 left eye removed
to show oesophageal pump and tentorium .

--{\

248

Figs 243--248 Chrysops caecutiens, 5?; mouth parts . 243, labrum and its distal end; 244, left mandible; 245, right
maxilla in ventral view; 246, sialophore and cibarial and pharyngeal pumps; 247, labium in dorsal view; 248,
collecting channel and pseudotrachaea of left labellum.

249

250

I
.05

251

Figs 249-253 Chrysops caecutiens , S?; internal structure of labium. 249, labial glands; 250, transverse sections of
middle part of Iabella and pseudotracheae; 251, transverse section of middle part of labium showing position
of other mouth parts; 252, sensilla coeloconica of inner wall of Iabella; 253, sensilla trichodea of outer wall of
Iabella .

.1

.05

.1

Figs 254--259 Chrysops caecutiens, S?; transverse sections. 254, most anterior part of head; 255, posterior part of
pharyngeal pump; 256, posterior part of head showing oesophageal sphincter (circular muscles) etc.; 257,
oesophagus; 258, duct of crop; 259, crop.

cb.

lm

sial
ph. pmp

mid-g

an
Fig. 260 Chrysops caecutiens, Cj2; alimentary canal and salivary glands.

86

262
.05

'

'

263

264

Figs 261-264 Chrysops caecutiens, <j?; salivary pump and salivary glands. 261, transverse section of anterior part
of common salivary duct; 262, same of pharyngeal and salivary pumps; 263, longitudinal and transverse
sections of posterior part of common salivary duct; 264, transverse section of salivary gland.

.01

265
Fig. 265 Chrysops caecutiens,

2; longitudinal sections of anterior part of mid-gut and stomodaeal invagination.

'\.0

thor . g
n9

_
vdnpt.
hr

.5

.05

Figs 267-270 Chrysops caecutiens, ~; histology of narrow thoracic, and abdominal, parts of mid-gut. 267,
transverse section of middle of abdomen showing Malpighian tubes, heart, ventral diaphragm, ovary and
parietal and visceral fat-body; 268, same of narrow thoracic part of mid-gut; 269, 270, longitudinal sections of
wall of ventriculus showing discharge of secretory globules by epithelial cells.
90

.05

272
273

ven

Figs 271-274 Chrysops caecutiens, <jl; histology of Malpighian tubes and anterior part of hind gut. 271 ,
longitudinal section of Malpighian tube showing its connection with ventriculus and anterior part of ileum;
272, transverse section of distal part of Malpighian tube; 273, same of ileum; 274, longitudinal section of
pylorus, posterior part of ventriculus and ileum.
91

Fig. 275 Chrysops caecutiens,

2; heart and respiratory system.

92

nephr

Figs 276, 277 Chrysops caecutiens, c;?; histology of heart and ventral diaphragm . 276, transverse section of heart;
277, abdominal part of ventral diaphragm.

cbl.
pmp

cl

1.0

th

lm

279
278

Figs 278, 279 Chrysops caecutiens.

~;

278, internal structure of head; 279, nervous system.

94

280

Figs 280--282 Chrysops caecutiens, 2; histology of anterior part of ventriculus and corpus allatum, transverse
sections. 280, anterior part of thorax; 281, caecum of anterior part of ventriculus; 282, corpus allatum, corpus
cardiacum and aorta.

95

284

.1

Figs 283-286 Chrysops caecutiens,

9. 283, histolysis of larval muscles by haemocytes; 284, oenocytes; 285, 286,

parietal and visceral fat-body.

\0

'-1

Fig. 287 Chrysops caecutiens,

S?; longitudinal section of posterior part of abdomen.

fig

1.0

Fig. 288 Chrysops caecutiens,

2; reproductive organs.

.05

289

290

292
.1

cir. m

293
Figs 289-295 Chrysops caecutiens, 2; histology of reproductive organs. 289, transverse section of accessory
glands; 290, same of posterior part of abdomen; 291, ovariole; 292, transverse section of wall of spermatheca
showing unicellular glands and their intracellular ducts; 293, 294, same of glandular and posterior parts of
spermathecal duct; 295, same of accessory gland duct.

The Stablefly

Stomoxys calcitrans

101

pt. sutf-

Fig. 296 Sto111oxys calcitrans,

102

2; a nterior view of head .

Fig . 297 Stomoxys calcitrans,

2; internal stru cture of hea d and proboscis.

ern

.1
pre.w .lm

298

304
Figs 298-304 Stomoxys calcitrans, S?; mouth parts. 298, distal end of sialophore; 299, distal end of labrum; 300,
proboscis; 301, 302, transverse sections through anterior and middle parts of proboscis; 303, ventral view of
Iabella, left half showing internal structure; 304, transverse section through most posterior part of same.

mal. t

Fig. 305 Stomoxys ca/citrans,

2; dorsal view of dissected fly showing position of internal organs.

105

Fig. 306 Stomoxys calcitrans,

106

S?; alimentary canal.

OL,
int

307

309

308
.1

311
.1

312

Figs 307-313 Stomoxys calcitrans, 9; structure of salivary glands, oesophagus and crop. 307, transverse section
through duct of salivary glands; 308, 309, transverse sections through anterior and posterior parts of salivary
gland; 310, 311, transverse sections through most anterior and cerebral parts of oesophagus; 312, histological
structure of wall of crop; 313, transverse section through crop.
107

.02

].02
314

16

315
.1
.05

321
.03

.1

317

.03

vii

pr. sh

cir. m

320

b.mem
reg. eel

319

322

Figs 314-322 Stomoxys calcitrans, <;?;structure of mid-gut. 314, 315, transverse sections of cardia and its dorsal
wall; 316, cells secreting substance of peritrophic membrane; 317, transverse section of mid-gut immediately
behind cardia; 318, muscles of mid-gut; 319, transverse section of broadest part of mid-gut; 320, part of same;
321, transverse section of narrow posterior part of mid-gut; 322, part of same.
108

.2

323

.1

325

.05

int

long. m

326
327

Figs 323-327 Stomoxys calcitrans, ~; structure of anterior and posterior intestine. 323, rectal valve; 324, rectal
papilla; 325, 326, transverse sections of wall of ileum and posterior part of rectum; 327, longitudinal section of
posterior part of abdomen.

109

prit. m

if

328
.1

.03

vii

331

''

'\

'
.02
330

332

Figs 328-332 Stomoxys calcitrans, <;?; structure of Malpighian tubes. 328, longitudinal section of proctodaea!
(pyloric) valve showing attachment of Malpighian tubes; 329, longitudinal section of middle part of Malpighian tube; 330, transverse section of Malpighian tube; 331, transverse section of lime sac of Malpighian tube
showing calcospherites; 332, calcospherite.
110

mal. tu
.1

333

.03

336
long. m

334
.03

.02

337
Figs 333-337 Stomoxys calcitrans, S?; structure of heart. 333, transverse section through posterior part of heart;
334, part of heart showing ostial valves and alary muscles with nephrocytes or pericardia! cells; 335,
pericardia! cells; 336, haemocytes; 337, fat-body with two oenocytes.

111

Fig. 338 Sto111oxys cnlcitrnns,

<?;

ventra l view of h ea rt.

339

.03

340
Figs 339, 340 Stomoxys calcitrans, <;?; corpus alia tum and hypocerebral ganglion, transverse and longitudinal
sections.

113

1.0

epi

sph. m

342
m

346

344
343
Figs 341-346 Stomoxys calcitrans, C(; reproductive organs . 341, general structure; 342, transverse section of
accessory glands; 343, 344, same of anterior and posterior parts of its duct; 345, 346, transverse sections of
spermathecae and their ducts.
114

348

.1

.5

349
rem. fol. epi

.03

350

Figs 347-350 Stomoxys calcitrans, 2; histology of ovary. 347, two ovarioles, one containing remainder of
disintegrated epithelium of first follicle after oocyte has left follicular tube; 348, transverse section of lateral
oviduct; 349, surface view of ovarian sheath showing anastomosing muscles; 350, posterior end of egg
showing oosome.

.02

tes. sh

355

Figs 351-356 Stomoxys calcitrans, d'; reproductive organs. 351, general structure; 352, 353, transverse sections
of vesicula seminalis and vas deferens; 354, same of posterior part of ejaculatory duct; 355, longitudinal
section of testis; 356, wall of same.
116

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ANATOMICAL DRAWINGS OF
BITING FLIES

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