Entomology 3rd edition - C.Gillott - Chapter 9 doc

9
The Panorpoid Orders
1
. Intr
oduc
t
ion
I
n this and the followin
g
chapter we shall deal with the endopter
yg
ote insects—those tha
t
h
ave a distinct pupal instar in which the insect under
g
oes a drastic metamorphosis fro
m
t
he larval to the adult form. As noted in Chapter 2, Section 3.2, considerable difficult
y
h
as ar
i
sen
i
n
d
ec
idi

ng w
h
et
h
er t
h
een
d
opterygote or
d
ers
h
ave a common or
i
g
i
norare
po
l
yp
h
y
l
et
i
c. T
h
e
fi
ve or

d
ers cons
id
ere
di
nt
hi
sc
h
apter, Mecoptera, D
i
ptera, S
i
p
h
onaptera
,
T
r
i
c
h
optera, an
d
Lep
id
optera, s
h
ow c
l

ear a
ffi
n
i
t
i
es t
h
at ena
bl
et
h
em to
b
e
g
roupe
d
to
g
et
h
e
r
as the
p
anor
p
oid com
p

lex. Within the com
p
lex there are two sister lines of evolution: th
e
Antlio
p
hora (first three orders) and the Am
p
hiesmeno
p
tera (Tricho
p
tera and Le
p
ido
p
tera).
Th
e rema
i
n
i
ng or
d
ers, Mega
l
optera, Rap
hidi
optera, Neuroptera, Co
l

eoptera, Hymenoptera
,
an
d
Streps
i
ptera,
d
ea
l
tw
i
t
hi
nC
h
apter 10, s
h
ow
f
ew a
ffi
n
i
t
i
es w
i
t
h

t
h
e panorpo
id
g
roup
.
2
. Mecoptera
S
ynonyms:
P
anor
p
atae, Panor
p
ina, Panor
p
id
a
C
ommon name:
s
cor
p
ionflies
S
lender medium-sized insects; head usually prolonged ventrally into a broad rostrum with long
filif
orm antennae, we

ll
-
d
eve
l
ope
d
compoun
d
eyes, an
dbi
t
i
ng mout
h
parts; usua
ll
yw
i
t
h
two pa
i
rs
o
f identical membranous win
g
s with primitive venation and carried horizontall
y
at rest; abdomen

w
ith short cerci and, in males, prominent genitalia.
L
arvae usually eruciform with simple eyes, biting mouthparts, and thoracic legs; abdominal legs
present or a
b
sent. Pupae
d
ect
i
cous an
d
exarate
.
This is a small order containin
g
about 500 known species, about 90% of which belon
g
t
o
t
wo families, Panorpidae and Bittacidae. The order is particularly common in the Northern
Hemis
p
here and includes about 7
5
North American and 4 British s
p
ecies. About 30 s
p

ecies
occur
i
n Austra
li
a.
23
9
240
CHAPTER
9
Structur
e
Adult
.
A
ch
aracter
i
st
i
c
f
eature o
f
most Mecoptera
i
st
h
e ventra

l
pro
l
ongat
i
on o
f
t
he
head into a broad rostrum. Incorporated into this structure are the cl
y
peus, labrum, and
m
axillae. Compound e
y
es are well developed, and in most species there are three ocelli.
The antennae are multise
g
mented and filiform. The mouthparts are mandibulate, except in
N
annoc
h
orista,w
h
ere t
h
ey are spec
i
a
li

ze
d
an
d
may
b
e
i
nterprete
d
as
f
ores
h
a
d
ow
i
ng t
h
e
s
uctor
i
a
l
type seen
i
n
l

ower D
i
ptera. T
h
e prot
h
orax
i
s sma
ll
,t
h
e pterot
h
orax we
ll d
eve
l
ope
d.
T
h
e
l
e
g
s are
l
on
g

an
d
t
hi
nan
d
a
d
apte
df
or wa
lki
n
g
.T
h
e
yh
ave a
fi
ve-se
g
mente
d
tarsus. In
Bittacidae the fifth tarsal se
g
ment folds back on the fourth and is used for catchin
g
pre

y.
Two
pairs of full
y
developed, identical, membranous win
g
s are present in most species; the
v
enat
i
on
i
spr
i
m
i
t
i
ve. In Bore
id
ae t
h
ew
i
ngs o
ff
ema
l
es are sma
ll

sc
l
erot
i
ze
d
pa
d
sw
hil
e
t
h
ose o
f
ma
l
es are
h
oo
klik
ean
d
use
d
to grasp t
h
e
f
ema

l
e
d
ur
i
ng mat
i
ng. W
i
ngs are re
d
uce
d
i
n some
f
ema
l
e Panorpo
did
ae an
d
B
i
ttac
id
ae, an
d
a
b

sent
i
n
f
ema
l
e Apteropanorp
id
ae. T
h
e
abdomen of females is 11-se
g
mented and usuall
y
carries 2-se
g
mented cerci (unse
g
mented
i
n Bittacidae and Boreidae). In female Boreidae the 10th ter
g
um is prolon
g
ed and to
g
ethe
r
wi

t
h
t
h
epo
i
nte
d
,sc
l
erot
i
ze
d
cerc
if
orms a
f
unct
i
ona
l
ov
i
pos
i
tor.Inma
l
es segment 9
i

s
bif
urcate an
db
ears a pa
i
ro
fb
u
lb
ous c
l
aspers. Segment 10
i
s
i
nconsp
i
cuous an
db
ears
unse
g
mente
d
cerc
i
.T
h
eae

d
ea
g
us
li
es at t
h
e
b
ase o
f
t
h
ec
l
aspers. In Panorp
id
ae t
h
e term
i
na
l
s
e
g
ments are turned upward and resemble somewhat a scorpion’s stin
g
, hence the common
n

ame for the order
.
T
he foregut has two interesting features. The esophagus contains two dilations that
appear to
f
orm a suc
ki
ng apparatus, an
d
t
h
e crop
i
s prov
id
e
d
w
i
t
hl
ong setae (acant
h
ae
)
t
h
at may act as a
fil

ter. S
i
xMa
l
p
i
g
hi
an tu
b
u
l
es occur. T
h
e nervous system
i
s genera
li
ze
d
,
w
ith three thoracic and between five and ei
g
ht abdominal
g
an
g
lia (males usuall
y

with on
e
m
ore than females). Each testis com
p
rises three or four follicles. The
p
aired vasa defer-
e
ntia open separately into a median seminal vesicle, which also receives paired accessor
y
gl
an
d
s. In
f
ema
l
es eac
h
ovary conta
i
ns 7–19 po
l
ytrop
hi
covar
i
o
l

es (pano
i
st
i
covar
i
o
l
es
in
Nannoc
h
or
i
st
id
ae an
d
Bore
id
ae). T
h
epa
i
re
d
ov
id
ucts un
i

te
b
e
f
ore enter
i
ng a gen
i
ta
l
pouc
h
.
T
h
e
d
ucts
f
rom t
h
e spermat
h
eca an
d
accessor
ygl
an
d
sa

l
so
l
ea
di
nto t
h
e pouc
h.
L
arva an
dP
upa
.
Larvae are typically caterpillarlike, with a distinct head capsule tha
t
b
ears s
i
mp
l
e eyes. Pro
l
egs occur on t
h
e
fi
rst e
i
g

h
ta
bd
om
i
na
l
segments, an
d
t
h
e apex o
f
t
h
e
a
bd
omen
b
ears e
i
t
h
er a suct
i
on
di
sc or a pa
i

ro
fh
oo
k
s. In Bore
id
ae an
d
Panorpo
did
ae
l
arvae
are
g
rublike, lackin
g
prole
g
s and a terminal suction disc. Larvae of Nannochoristidae are
v
er
y
elon
g
ate, lack prole
g
s, but have a pair of apical hooks. Pupae are decticous and exarate
.
Lif

eH
i
story and Hab
i
t
s
Adult scorpionflies are most frequentl
y
encountered in cool, shaded locations, espe-
c
ially among low vegetation, though a few species occur in semidesert habitats. They ca
n
fl
y act
i
ve
l
yw
h
en
di
stur
b
e
d
,t
h
oug
h
t

h
ey norma
ll
y rest on grass, un
d
er
l
eaves, etc. A
d
u
l
t
P
anorp
id
ae
f
ee
d
most
l
yon
d
ea
d
so
f
t-
b
o

di
e
d
art
h
ropo
d
s(
i
nc
l
u
di
ng
i
nsects caug
h
t
i
nsp
i
-
ders’ webs); the
y
also eat nectar, pollen, and fruit
j
uices. Bittacidae, b
y
contrast, are insect
p

redators, catchin
g
their pre
y
either in fli
g
ht or b
y
han
g
in
g
under ve
g
etation till it comes
within range. In members of both of these families much of the food of females is pro
-
vid
e
di
nt
h
e
f
orm o
f
a nupt
i
a
l

g
if
t
b
yama
l
e
d
ur
i
ng courts
hi
p (see C
h
apter 19, Sect
i
on 4.2)
.
T
h
e
f
oo
di
tem may
b
eanart
h
ropo
d

recent
l
yo
b
ta
i
ne
db
yt
h
ema
l
e or a mass o
f
sa
li
va
s
ecreted from the male’s
g
reatl
y
enlar
g
ed salivar
yg
lands. Adult Boreidae feed on mosses,
241
T
HE P

A
N
O
RP
O
ID
ORDER
S
and members of other families ma
y
be herbivorous, sapropha
g
ous, or carrion feeders. To
attract females, male panorpids and bittacids secrete pheromones from
g
lands on the pos-
t
erior abdominal se
g
ments. Visual si
g
nals (win
g
movements and abdominal vibrations)
are a
l
so
i
mportant
i

nc
l
ose-range courts
hi
p
i
nteract
i
ons
i
nt
h
ese an
d
ot
h
er (non-p
h
eromone-
pro
d
uc
i
ng) mecopteran
f
am
ili
es. Eggs o
fbi
ttac

id
s are
d
roppe
d
ran
d
om
l
y; t
h
ose o
f
panorp
ids
an
d
c
h
or
i
st
id
s are
l
a
id i
n
b
atc

h
es
i
nmo
i
st
d
epress
i
ons
i
nt
h
e
g
roun
d
. Bore
id
ae
d
epos
i
te
gg
s
sin
g
l
y

or in small batches in soil ad
j
acent to moss rhizoids. The e
gg
sta
g
ema
y
last from as
little as a week up to several months in species where there is an e
gg
diapause. Larvae ar
e
saprop
h
agous, carn
i
vorous, or moss-
f
ee
d
ers an
d
pass t
h
roug
hf
our
i
nstars

b
e
f
ore enter
i
ng
aqu
i
escent prepupa
l
p
h
ase, usua
ll
y
i
n an eart
h
en ce
ll
.T
h
e
l
engt
h
o
f
t
h

e prepupa
l
p
h
as
e
is varied and ma
y
include a diapause. The pupal stadium usuall
y
lasts 14–
5
0da
y
s. Mos
t
species are univoltine; some are bivoltine; and boreids take 2
y
ears to complete a
g
eneration.
Phylogeny and Classification
Ar
i
c
h
array o
f
mecoptera
lik

e
f
oss
il
s
i
s
k
nown
f
rom t
h
e Lower Perm
i
an, an
d
t
h
emo
d-
ern consensus is that this includes a few
g
enuine Mecoptera. B
y
the Upper Permian, and
extendin
g
throu
g
h the Jurassic, the order was abundant and diverse. Some Upper Permian

scorpionflies from deposits in Australia and Siberia are assi
g
nable to the extant famil
y
Nan
-
noc
h
or
i
st
id
ae, w
hil
ere
p
resentat
i
ves o
f
some ot
h
er mo
d
ern
f
am
ili
es a
pp

ear
i
nt
h
eLowe
r
Jurass
i
c. Recent Mecoptera are arrange
db
yW
ill
mann (1987)
i
ntwosu
b
or
d
ers, conta
i
n
i
ng
n
i
ne
f
am
ili
es w

h
ose poss
ibl
eevo
l
ut
i
onar
y
re
l
at
i
ons
hi
ps are s
h
own
i
nF
ig
ure 9.1
.
F
I
GU
RE 9.1. Proposed phylogeny of the extant Mecoptera. [After R. Willmann, 1987, The phylogenetic syste
m
o
f

t
h
e Mecoptera
,
S
yst. Entomo
l.
12
:519–524. By permission of Blackwell Scientific Publications Ltd.]
242
CHAPTER
9
Suborder Nannomeco
p
tera
Thi
ssu
b
or
d
er conta
i
ns t
h
es
i
ng
l
e, sma
ll

,pr
i
m
i
t
i
ve Sout
h
ern Hem
i
sp
h
ere
f
am
ily
NANNOCHORISTIDAE. The ei
g
ht species in this
g
roup differ from other mecopterans in
l
ackin
g
a pump for transferrin
g
sperm from male to female, and in details of their external
g
enitalia, win
g

venation, and mandibular structure. Adults are small and live in the vicinit
y
of
streams or
l
a
k
es. T
h
e
i
r
l
arvae are aquat
i
can
d
carn
i
vorous,
f
ee
di
ng on
l
arva
l
D
i
ptera,

e
spec
i
a
ll
yc
hi
ronom
id
s.
Su
b
or
d
er Pisti
ll
i
f
era
T
he suborder Pistillifera (literally “piston-bearers” in reference to the sperm
pump o
f
ma
l
es)
i
s
di
v

id
e
db
yW
ill
mann (1987)
i
nto two
i
n
f
raor
d
ers, Rapt
i
pe
di
aan
d
O
pisthogonopora. The Raptipedia includes the large (14
5
species) and cosmopolitan fam
-
i
l
y
BITTACIDAE (Fi
g
ure 9.2C) whose members have raptorial tarsi with which the

yg
ras
p
their pre
y
, often han
g
in
g
under ve
g
etation b
y
their forele
g
s. Amon
g
the Opistho
g
ono-
pora, the BOREIDAE (Figure 9.2A) constitute the most distinct family; indeed, Hinto
n
(
19
5
8) placed this holarctic group of about 2
5
species in a separate order, Neomecoptera
.
F

I
GU
RE 9.2. Meco
p
tera. (A)
B
oreus
b
ruma
l
is
(
Bore
id
ae
)
;
(
B
)
Panorpa
h
e
l
en
a
(Panor
pid
ae); an
d

(C
)
B
i
ttacus
p
ilicornis
(
Bittacidae). [A, B, from D. J. Borror, D. M. Delong, and C.A. Triplehorn, 1976
,
An Intro
d
uction to th
e
S
tu
d
y of Insects
,
4t
h
e
d
. By perm
i
ss
i
on o
f
Broo

k
s/Co
l
e, a
di
v
i
s
i
on o
f
T
h
omson Learn
i
ng. C,
f
rom D. W. We
bb
,N.
D
. Penn
y
, and J. C. Marlin, 197
5
. The Mecoptera, or scorpionflies, of Illinois
.
B
u
ll

.I
ll
. Nat. Hist.
S
ur
v.
31
:2
5
1–316.
By permission of the Illinois Natural History Survey.]
243
T
HE P
A
N
O
RP
O
ID
ORDER
S
B
oreidss are small, with hooklike win
g
s in males used to
g
rasp the female durin
g
matin

g,
and scalelike reduced win
g
s in females. The
y
are sometimes called “snow fleas” as the
y
ma
y
be found walkin
g
or
j
umpin
g
on snow patches. Onl
y
two, widel
y
dis
j
unct species,
A
ustromerope pou
l
toni
(
Western Austra
li
a) an

d
M
erope tu
b
e
r
(
eastern Un
i
te
d
States),
are p
l
ace
di
nt
h
e MEROPEIDAE. T
h
ese are coc
k
roac
hlik
e
i
n appearance as t
h
e
i

r
h
ea
d
is lar
g
el
y
hidden beneath the enlar
g
ed pronotum. Males have elon
g
ate
g
enitalia possi
-
b
l
y
used in fi
g
htin
g
for females.
N
otiothauma reed
i
,
found in central Chile, is the onl
y

member of the NOTIOTHAUMIDAE. Similarl
y
,
A
pteropanorpa tasmanica
f
r
o
mT
as
m
a-
n
i
a, w
hi
c
h
resem
bl
es
b
ore
id
s
i
n genera
lf
orm an
dh

a
bi
ts,
i
st
h
eso
l
e representat
i
ve o
f
th
e APTEROPANORPIDAE. CHORISTIDAE, w
i
t
h
on
l
ye
i
g
h
t spec
i
es, are restr
i
cte
d
to

Australia. The two remainin
g
families, PANORPODIDAE and PANORPIDAE (Fi
g
ur
e
9.2B), are the most specialized mecopteran
g
roups. The former includes onl
y
two
g
enera,
Brachypanorp
a
(
four s
p
ecies in the United States) and
P
anorpode
s
(five s
p
ecies in Ja
p
an
an
d
Korea), w

hil
et
h
e
l
atter
i
st
h
e
l
argest mecopteran
f
am
il
y (300 spec
i
es) w
i
t
h
an essent
i
a
ll
y
h
o
l
arct

i
c
di
str
ib
ut
i
on, t
h
oug
hi
t
h
as representat
i
ves
i
nAs
i
a.
Literatur
e
Because o
f
t
h
e centra
l
pos
i

t
i
on t
h
at t
hi
sor
d
er occup
i
es
i
nany
di
scuss
i
on o
f
t
h
eevo
l
ut
i
on
o
f
most en
d
opterygote or

d
ers,
i
nterest
i
nt
h
e Mecoptera
h
as
b
een out o
f
proport
i
on to t
h
e
number of extant species. Information on the biolo
gy
of Mecoptera is provided b
y
Webb
et al.
(1975), Kaltenbach (1978), and B
y
ers and Thornhill (1983). The evolution of th
e
order and its relationship to the other panorpoid groups are discussed by Hennig (1981),
Kr

i
stensen (1981), W
ill
mann (1987), an
d
W
hi
t
i
ng (2002). Keys to
f
am
ili
es o
f
Mecoptera
in
N
ort
h
Amer
i
ca an
d
Austra
li
a are prov
id
e
db

y Arnett (2000) an
d
Byers (1991), respect
i
ve
l
y
.
Th
eBr
i
t
i
s
h
spec
i
es are
id
ent
ifi
a
bl
e
f
rom P
l
ant (1997)
.
A

rnett, R. H. Jr.,
2000,
American Insects: A
h
an
db
oo
k
of t
h
e Insects of America Nort
h
of Mexico
,
2n
d
e
d
., CRC
P
ress
,
Boca Raton
,
FL.
B
yers, G. W., 1991, Mecoptera, in
:
T
he Insects o

f
Australia
,
2nd ed., Vol. 2 (CSIRO, ed.), Melbourne Universit
y
P
ress, Car
l
ton, V
i
ctor
i
a
.
By
ers, G. W., an
d
T
h
orn
hill
, R., 1983, B
i
o
l
o
gy
o
f
t

h
e Mecoptera
,
A
nnu. Re
v
. Entomo
l.
28
:
203–228
.
H
ennig, W., 1981, Insect Ph
y
lo
g
en
y
,
W
iley, New York.
WW
H
inton, H. E., 1958, The phylogeny of the panorpoid orders
,
Annu. Rev. Entomo
l
.
3

:181–20
6
.
Ka
l
ten
b
ac
h
, A., 1978, Mecoptera (Sc
h
na
b
e
lh
a
f
te, Sc
h
na
b
e
lfli
e
g
en),
i
n: Han
db
uc

hd
er Zoo
l
ogie
,V
ol. IV, Insecta
V
V
L
fg
.
28
:1–111, de Gruyter, Berlin.
Kr
i
stensen, N. P., 1981, P
h
y
l
ogeny o
fi
nsect or
d
ers
,
A
nnu. Rev. Entomo
l
. 2
6

:
135–157
.
Pl
ant, C. W., 1997, A
k
e
y
to a
d
u
l
ts o
f
Br
i
t
i
s
hl
acew
i
n
g
san
d
t
h
e
i

ra
lli
es (Neuroptera, Me
g
a
l
optera, Rap
hidi
opter
a
and Mecoptera)
,
Field
S
tud.
9
:179–269
.
W
ebb, D. W., Penny, N. D., and Marlin, J. C., 1975. The Mecoptera, or scorpionflies, of Illinois, Bu
ll
.I
ll
. Nat
.
H
ist.
S
ur
v.

31
(
7
)
:2
5
1–316
.
W
hiting, M. F., 2002, Mecoptera is paraphyletic: Multiple genes and phylogeny of Mecoptera and Siphonaptera
,
Zoo
l
. Scrip.
3
1:
93
–
104
.
W
illmann, R., 1987, The phylogenetic system of the Mecoptera,
WW
Syst. Entomo
l
.
12
:
5
1

9
–
5
24
.
3
. Diptera
S
ynonyms:
n
one
C
ommon names
:
true flies; includes mosquitoes, mid
g
es, black flies, dee
r
flies
,
horse flies
,
house flie
s
244
CHAPTER
9
Generally minute to small soft-bodied insects; head highly mobile with large compound eyes,
a
ntennae o

f
var
i
e
d
s
i
ze an
d
structure, an
d
suctor
i
a
l
mout
h
parts; prot
h
orax an
d
metat
h
orax sma
ll
a
n
df
use
d

w
i
t
hl
ar
g
e mesot
h
orax, w
i
n
g
s present on
ly
on mesot
h
orax,
h
a
l
teres present on metat
h
o-
r
ax; legs with five-segmented tarsi; abdomen with varied number of visible segments, femal
e
g
en
i
ta

li
as
i
mp
l
e
i
n most spec
i
es, ma
l
e
g
en
i
ta
li
a comp
l
ex, cerc
i
present.
Larvae eruc
if
orm an
di
n most spec
i
es apo
d

ous;
h
ea
di
n many spec
i
es re
d
uce
d
an
d
retracte
d
.
P
u
p
ae a
d
ect
i
cous an
d
o
b
tect or exarate, t
h
e
l

atter enc
l
ose
di
na
p
u
p
ar
i
um.
T
he more than 120,000 s
p
ecies of Di
p
tera described to date re
p
resent
p
erha
p
s two-thirds
o
f the world total. The order has a truly worldwide distribution, representatives occurring
ev
e
n in the Antarctic. More than 19,
5
00 species have been described from North America,

a
l
most 8000
f
rom Austra
li
a
,
an
d
c
l
ose to 7000
f
rom Br
i
ta
i
n. Some o
f
t
h
ewor
ld
’s commonest
i
nsects and a lar
g
e number of species of veterinar
y

and medical importance belon
g
to this
o
rder
.
S
tructur
e
T
he
g
reat structural diversit
y
found in the Diptera reflects the variet
y
of niches that th
e
true flies have ex
p
loited.
A
du
lt
.
Adults range in size from about 0.
5
mm to several centimeters and the
y
are

g
enera
lly
so
f
t-
b
o
di
e
d
.T
h
e
h
ea
di
sre
l
at
i
ve
ly l
ar
g
ean
d highly
mo
bil
e. It carr

i
es we
ll-
developed compound e
y
es, which in males are frequentl
y
holoptic. The antennae are o
f
v
aried size and structure and are important taxonomicall
y
. In Muscomorpha-Schizophor
a
t
h
e
r
e
i
sa
∩
-
shaped ptilinal (frontal) suture that runs transversely above the antennae and
e
xten
d
s
d
ownwar

d
on eac
h
s
id
eo
f
t
h
em. T
hi
s suture
i
n
di
cates t
h
e pos
i
t
i
on o
f
t
h
ept
ili
num, a
m
em

b
ranous sac t
h
at
i
s exserte
d
an
ddi
sten
d
e
d
at ec
l
os
i
on
i
nor
d
er to rupture t
h
e pupar
i
u
m
and assist a fl
y
in tunnelin

g
throu
g
h soil, etc. The mouthparts are adapted for suckin
g
an
d
are described in Chapter 3 (see Section 3.2.2 and Fi
g
ures 3.14–3.16). The prothoracic and
m
etathoracic segments are narrow and fused intimately with the very large mesothorax,
whi
c
hb
ears t
h
es
i
ng
l
epa
i
ro
f
mem
b
ranous w
i
ngs. T

h
e
hi
n
d
w
i
ngs are extreme
l
ymo
difi
e
d
,
f
orm
i
ng
h
a
l
teres, sma
ll
,c
l
u
blik
e structures
i
mportant as organs o

fb
a
l
ance (see C
h
apter 14
,
S
ect
i
on 3.3.4). In a
f
ew spec
i
es
h
a
l
teres an
d
w
i
n
g
s are re
d
uce
d
or a
b

sent. T
h
e
l
e
g
sa
l
most
alwa
y
s have five-se
g
mented tarsi, and in some species one or more pairs are modified fo
r
g
raspin
g
pre
y
. Primitivel
y
, there are 11 abdominal se
g
ments, but in most Diptera this number
i
s reduced and rarely more than 4 or 5 are readily visible. Frequently, the more posterior
segments (posta
bd
omen) are te

l
escope
di
nto t
h
e anter
i
or part o
f
t
h
ea
bd
omen (prea
bd
omen)
(
see F
ig
ure 3.29). T
h
e posta
bd
omen t
h
us
f
orme
di
s use

d
as an extens
ibl
eov
i
pos
i
tor. T
h
e
m
ale
g
enitalia are complex and their homolo
g
ies uncertain because of the rotation of the
abdomen and as
y
mmetric
g
rowth of the individual components durin
g
the pupal sta
g
e
.
In most D
i
ptera t
h

ec
ib
ar
i
um
i
s strong
l
y muscu
l
ar an
d
serves as a pump
f
or suc
ki
ng u
p
li
qu
id
s
i
nto t
h
e gut. In t
h
e
bl
oo

d
suc
ki
ng Ta
b
an
id
ae an
d
Cu
li
c
id
ae a
l
arge p
h
aryngea
l
pump
i
sa
l
so present. T
h
ea
li
mentar
y
cana

li
s,
i
n most pr
i
m
i
t
i
ve
f
orms, re
l
at
i
ve
ly
unconvo
l
ute
d
.
In Muscomorpha, however, it is much more coiled because of the increase in len
g
th of the
m
id
g
ut. The esopha
g

us divides posteriorl
y
into the
g
izzard and, usuall
y
, one diverticulum,
the food reservoir (misleadingly called the “crop”). In Culicidae three diverticula are found
.
In Nematocera t
h
em
id
gut
i
sas
h
ort, sac
lik
e structure;
i
n Muscomorp
h
a
i
t
i
s
l
ong an

d
c
onvo
l
ute
d
. Genera
ll
yt
h
ere are
f
our Ma
l
p
i
g
hi
an tu
b
u
l
es t
h
at ar
i
se
i
npa
i

rs
f
rom a common
duct on either side of the
g
ut. In the nervous s
y
stem a complete ran
g
e of specialization
245
T
HE P
A
N
O
RP
O
ID
ORDER
S
is seen. In primitive Nematocera three thoracic and seven abdominal
g
an
g
lia occur, bu
t
all intermediate conditions are found between this arran
g
ement and the situation in the

more advanced Muscomorpha where a composite thoracoabdominal
g
an
g
lion exists. I
n
f
ema
l
es t
h
epa
i
re
d
ovar
i
es compr
i
seavar
i
e
d
num
b
er o
f
po
l
ytrop

hi
covar
i
o
l
es. In v
i
v
i
parous
spec
i
es t
h
ere may
b
eon
l
y one or two,
b
ut
i
nt
h
ema
j
or
i
ty o
f

ov
i
parous
fli
es t
h
ere may
b
e
more t
h
an 100. In v
i
v
i
parous
f
orms t
h
e common ov
id
uct
i
s
dil
ate
d
to
f
orm a uterus, an

d
t
he accessor
y
(“milk”)
g
lands produce a nutritive secretion. One to four spermathecae ar
e
alwa
y
s present. In males the testes are
g
enerall
y
small, ovoid, and pi
g
mented. The short
,
pa
i
re
d
vasa
d
e
f
erent
i
a
l

ea
di
nto a muscu
l
ar e
j
acu
l
atory sac. Pa
i
re
d
accessory g
l
an
d
s may
occur.
L
arva an
dP
u
p
a
.
Larvae are usuall
y
elon
g
ate and c

y
lindrical. Bod
y
se
g
mentatio
n
is usually distinct, but in a few groups the true number of segments is masked as a re
-
su
l
to
f
secon
d
ary
di
v
i
s
i
on or
f
us
i
on o
f
t
h
eor

i
g
i
na
l
segments. True t
h
orac
i
c
l
egs are a
l-
w
ays a
b
sent, t
h
oug
h
pro
l
egs may occas
i
ona
ll
y
b
e present on t
h

et
h
orax an
d
/or a
bd
omen
.
I
n primitive Diptera the head capsule is distinct and sclerotized (the eucephalous condi-
t
ion). In most species, however, it is much reduced (hemicephalous) or entirel
y
vesti
g
ia
l
(acephalous) (Fi
g
ure 3.13). The antennae and chewin
g
mouthparts, includin
g
horizontall
y
mov
i
ng man
dibl
es, are we

ll d
eve
l
ope
di
n Nematocera. In ort
h
orr
h
ap
h
ous spec
i
es anten
-
nae an
d
max
ill
ae may
b
ewe
ll d
eve
l
ope
d
,
b
ut t

h
e man
dibl
es are s
i
c
kl
e-s
h
ape
d
an
d
mov
e
i
navert
i
ca
l
p
l
ane. In Muscomorp
h
at
h
e antennae are
i
nt
h

e
f
orm o
f
m
i
nute pap
ill
ae an
d
t
he mouthparts are reduced to a pair of curved hooks (the ori
g
inal mandibles). The internal
structure of larvae
g
enerall
y
resembles that of adults. Dipteran pupae are alwa
y
s adecticous
.
Pu
p
ae o
f
nematocerous an
d
ort
h

orr
h
a
ph
ous s
p
ec
i
es are o
b
tect, w
h
ereas t
h
ose o
f
Musco-
morp
h
a are secon
d
ar
il
y exarate an
d
coarctate,
b
e
i
ng enc

l
ose
di
n a pupar
i
um, t
h
e
h
ar
d
ene
d
cut
i
c
l
eo
f
t
h
et
hi
r
dl
arva
li
nstar.
L
if

eH
i
story and Hab
i
ts
Adult Diptera are active, mostly free-living insects that are found in all major habitats.
Th
ey are pre
d
om
i
nant
l
y
di
urna
l
an
d
usua
ll
y assoc
i
ate
d
w
i
t
hfl
owers or w

i
t
hd
ecay
i
ng organ
ic
matter. W
i
t
h
t
h
e except
i
on o
f
a
f
ew spec
i
es t
h
at
d
o not
f
ee
d
as a

d
u
l
ts,
fli
es
f
ee
d
on
li
qu
id
s
(rarel
y
, also pollen), a habit that ma
y
reflect the use of hone
y
dew from homopterans as
t
heir ancestral ener
gy
source (Downes and Dahlem, 1987). Most Diptera feed on nectar
or the
j
uices from deca
y
in

g
or
g
anic matter, but a few
g
roups are adapted for feedin
g
o
n
th
et
i
ssue
fl
u
id
so
f
ot
h
er an
i
ma
l
s espec
i
a
ll
y art
h

ropo
d
san
d
verte
b
rates. T
hi
s
i
sac
hi
eve
din
some spec
i
es
b
ys
i
mp
l
y cutt
i
ng t
h
es
ki
n or squeez
i

ng prey w
i
t
h
t
h
e
l
a
b
e
ll
aan
d
suc
ki
ng up
th
eexu
d
e
dfl
u
id
.Int
h
ema
j
or
i

t
y
o
fb
o
dy
-
fl
u
id f
ee
d
ers,
h
owever, a
fi
ne pro
b
osc
i
s
i
s use
d
t
o pierce the skin and penetrate directl
y
to the fluid, usuall
y
blood. The habit is usuall

y
confined to females. It is throu
g
h the bloodsuckin
g
habit and the subsequent importanc
e
o
f
t
h
ese
i
nsects as vectors o
fdi
sease-caus
i
ng m
i
croorgan
i
sms t
h
at t
h
eor
d
er
i
s genera

lly
cons
id
ere
d
t
h
e most
i
mportant o
f
t
h
e ent
i
re c
l
ass
f
rom t
h
eme
di
ca
l
an
d
veter
i
nary po

i
nt o
f
v
i
ew . A
l
t
h
oug
h
part
h
enogenes
i
s
i
s
k
nown to occur
i
na
f
ew spec
i
es, most D
i
ptera repro
d
uce

b
isexuall
y
. Copulation is preceded in some species b
y
an elaborate courtship. Usuall
y
females activel
y
search for, and la
y
e
gg
s directl
y
on, the larval food source. Members of
af
ew
f
f
g
roups are ovoviviparous or viviparous. Egg development is normally rapid and
h
atc
hi
ng occurs
i
na
f
ew

d
ays.
Larvae are usua
ll
y
f
oun
di
nmo
i
st
l
ocat
i
ons suc
h
as so
il
,mu
d
,
d
ecay
i
ng organ
i
c matter
,
and plant or animal tissues, thou
g

h a few are trul
y
aquatic. The ma
j
orit
y
are liquid feeders o
r
2
4
6
CHAPTER
9
m
icropha
g
ous. Some aquatic forms trap their food in speciall
y
developed mouth brushes
.
Larvae of man
y
species are of a
g
ricultural or medical importance. Usuall
y
four larval
i
nstars occur, but up to ei
g

ht are found in some species, and in Muscomorpha the fourth i
s
suppresse
d
.Pr
i
or to pupat
i
on,
l
arvae genera
ll
y craw
l
to a
d
r
i
er
l
ocat
i
on. Pupae may
b
ena
k
e
d,
b
u

t
th
ose o
f
many nematocerous an
d
ort
h
orr
h
ap
h
ous spec
i
es are enc
l
ose
di
n a cocoon, an
d
t
h
ose o
f
Muscomorp
h
a are ens
h
eat
h

e
dby
t
h
e pupar
i
um. T
h
ep
h
arate a
d
u
l
tswa
ll
ows a
i
r
to facilitate emer
g
ence from the pupal skin or puparium, the latter splittin
g
len
g
thwise in
o
btect
p
u

p
ae; in coarctate forms, however, the adult
p
rotrudes the
p
tilinum to
p
ush off the
anter
i
or en
d
o
f
t
h
e pupar
i
um
.
P
hylo
g
eny and
C
lass
ifi
cat
i
o

n
F
rom a structural comparison of primitive livin
g
Mecoptera and Diptera and from the
re
l
at
i
ve
l
y scarce
f
oss
il
ev
id
ence,
i
t seems pro
b
a
bl
et
h
at D
i
ptera evo
l
ve

df
rom mecoptera
lik
e
i
nsects
i
nt
h
e Perm
i
an per
i
o
d.
P
ermoti
p
u
l
a
p
atrici
a
,
f
rom t
h
e Upper Perm
i

an o
f
Austra
li
a,
w
as
o
r
igi
na
lly
t
h
ou
gh
tto
b
et
h
eo
ld
est
di
pteran. However,
i
t
i
s now cons
id

ere
d
an ear
ly
M
ecopteran. Similarl
y
, other four-win
g
ed forms (the Protodiptera
P
ermotanytarsus
a
n
d
C
horistotanyderus) from the same strata, which were ori
g
inall
y
considered to be Diptera
(
R
i
e
k
, 1977),
h
ave
b

een reexam
i
ne
d
an
d
, as a resu
l
t, move
df
rom t
h
e
di
pteran stem group
(
W
ill
mann, 1989; Wootton an
d
Ennos, 1989). T
h
us, t
h
e ear
li
est re
li
a
bl

e recor
d
so
ff
os
-
s
il
D
i
ptera come
f
rom t
h
eM
iddl
eTr
i
ass
i
c, t
h
ou
gh
t
h
ese are qu
i
te mea
g

er. B
y
t
h
e
l
at
e
Triassic, the more primitive suborder, Nematocera, had under
g
one a considerable radia-
tion, possibl
y
makin
g
use of the hone
y
dew produced b
y
the alread
y
abundant homopteran
s
(
Downes and Dahlem, 1987). By the Lower Jurassic, well-developed Nematocera (som
e
ass
i
gna
bl

e to recent
f
am
ili
es) an
d
pr
i
m
i
t
i
ve ort
h
orr
h
ap
h
ous
f
orms were present. T
h
e Mus
-
c
omorp
h
a (Cyc
l
orr

h
ap
h
a) evo
l
ve
df
rom ort
h
orr
h
ap
h
an stoc
k
, pro
b
a
bl
y
i
nt
h
e Late Jurass
i
c
.
The
g
reat radiation of the order, and the establishment of man

y
of the structures and habit
s
o
f modern flies, took
p
lace in the Cretaceous
p
eriod. This was, of course, correlated wit
h
the evolution of the flowering plants and mammals. By the Eocene period, the diptera
n
f
auna was s
i
m
il
ar
i
n many respects to t
h
at w
hi
c
h
surv
i
ves to
d
ay. In

d
ee
d
, many Eocen
e
f
oss
il
s are ass
i
gne
d
to mo
d
ern genera. A poss
ibl
ep
h
y
l
ogeny o
f
t
h
eor
d
er
i
ss
h

own
in
F
i
g
ure 9.3.
Classification of the modern Diptera continues to present problems, particularl
y
the
rank assi
g
nable to different
g
roups. The difficult
y
arises in part because the order is ex-
treme
l
yo
ld
;
i
t conta
i
ns many ext
i
nct groups an
d
ot
h

ers t
h
at are
i
n
d
ec
li
ne; an
d
yet t
h
ere ar
e
a
l
so groups t
h
at are st
ill
evo
l
v
i
ng at a rap
id
rate. T
h
us, w
hil

e anc
i
ent
f
am
ili
es
h
ave we
ll
-
e
sta
bli
s
h
e
d diff
erences an
d
are eas
ily
separate
d
, more recent
g
roups, nota
bly
t
h

e
f
am
ili
e
s
and su
p
erfamilies of Muscomor
p
ha, are sometimes little more than convenient divisions
because of the vast number of s
p
ecies that must be considered. The differences between
t
h
ese groups, t
h
ere
f
ore, are re
l
at
i
ve
l
ym
i
nor. T
h

ec
l
ass
ifi
cat
i
on use
dh
ere
i
sa
d
apte
df
ro
m
t
h
ev
i
ews presente
di
nt
h
e Manua
l
o
f
Nearctic Diptera (McA
l

p
i
ne et a
l
., 1981–1989). In
t
hi
s arrangement, t
h
eD
i
ptera are arrange
di
ntwosu
b
or
d
ers, Nematocera an
d
Brac
h
ycera
.
Within the Nematocera, seven infraorders are reco
g
nized. Fi
g
ure 9.3 indicates that Nema
-
tocera are monoph

y
letic. However, it should be noted that some authors believe that this is
a
paraphyletic group, with the Bibionomorpha or Psychodomorpha being the sister group t
o
t
h
e Brac
h
ycera (Yeates an
d
W
i
egmann, 1999). In t
h
eun
d
ou
b
te
dl
y monop
h
y
l
et
i
c Brac
h
yc

-
e
ra t
h
ere are t
h
ree
i
n
f
raor
d
ers; t
h
e
fi
rst two, As
il
omorp
h
aan
d
Ta
b
anomorp
h
a, correspon
d
to
the “Orthorrhapha” while the third, Muscomorpha, is equivalent to the “C

y
clorrhapha” o
f
247
T
HE P
A
N
O
RP
O
ID
ORDER
S
F
I
G
URE 9.3.
A
s
u
gg
este
d
p
hyl
o
g
en
y

o
f
t
h
eD
i
ptera. [A
f
ter J. F. McA
l
p
i
ne an
d
D. M. Woo
d
, coor
di
nators
,
1989
,
M
anual o
f
Nearctic Dipter
a
,V
ol. 3, Agriculture Canada Monograph No. 32. By permission of the Minister
V

V
of Supply and Services Canada.]
earlier s
y
stems. Within the Muscomorpha, there are two easil
y
defined but ver
y
unequall
y
sized sections, the probabl
y
paraph
y
letic Aschiza and the monoph
y
letic Schizophora. Th
e
latter has two subdivisions, Acalyptratae and Calyptratae.
S
uborder Nematocera
Most Nematocera are small, delicate flies, with
6
- to 14-segmented antennae of simple
structure, and 3- to
5
-se
g
mented maxillar
y

palps. Larvae have a well-developed head an
d
chewin
g
mandibles that move in the horizontal plane.
248
CHAPTER
9
FIGURE 9.4
.
T
i
pu
l
omorp
h
a. A crane
fl
y, Ti
p
u
l
a trivittata
(
T
i
pu
lid
ae).
[

From F. R. Cole and E. I. Schlin
g
er, 19
6
9
.
T
h
eF
l
ies of Western Nort
h
A
m
e
r
ica.
By
permission of the Universit
y
of California Press.]
T
he suborder contains the oldest families of Di
p
tera, most of which are now on the
decline. Some, however, like the Culicidae, have undergone relatively recent radiations and
are among t
h
e most success
f

u
l
mo
d
ern groups.
Infraor
d
er Tipu
l
omorp
h
a
M
em
b
ers o
f
t
hi
s group are p
l
ace
di
nt
h
es
i
ng
l
e

f
am
il
y TIPULIDAE (F
i
gure 9.4)
(
crane
fli
es,
d
a
dd
y
l
ong
l
egs), w
hi
c
h
,w
i
t
h
a
b
out 14,000 spec
i
es (

i
nc
l
u
di
n
g
>
1
5
00 in North
A
merica), is the lar
g
est famil
y
of Diptera. Representatives of this worldwide famil
y
are
m
ostl
y
associated with moist, temperate habitats, thou
g
h some occur in open meadows,
rangelands, and deserts. Adults range in size from small (wingspan of 2 mm) to very larg
e
(
w
i

ngspan up to 8 cm). Larvae are
f
oun
di
naw
id
evar
i
ety o
fh
a
bi
tats
f
rom str
i
ct
l
y aquat
i
c
to
d
ry so
il
sw
h
ere t
h
ey typ

i
ca
ll
y
f
ee
d
on p
l
ant mater
i
a
l
or
d
ecay
i
ng organ
i
c matter; occa
-
sionall
y
, the
y
become pests b
y
feedin
g
on seedlin

g
field crops.
In
f
raorder Blephariceromorpha
Almost all members of this small
g
roup are included in the widel
y
distributed famil
y
BLEPHARICERIDAE (net-winged midges) (200 species). Adults are slender flies wit
h
l
ong
l
egs, an
di
n
b
ot
h
sexes t
h
e eyes are
h
o
l
opt
i

c. In some spec
i
es
b
ot
h
sexes
f
ee
d
on nectar
;
i
not
h
ers
f
ema
l
es catc
h
sma
ll
er
fli
es an
df
ee
d
on t

h
e
i
r
h
emo
l
ymp
h
.A
d
u
l
ts are genera
lly
f
oun
d
near
f
ast-
fl
ow
i
n
g
streams; t
h
e aquat
i

c
l
arvae attac
h
t
h
emse
l
vestoroc
k
sw
h
ere t
h
e
y
f
eed on diatoms and al
g
ae
.
In
f
raorder Axymyiomorpha
Containin
g
onl
y
five species in the one famil
y

AXYMYIIDAE, the taxonomic position
o
f this group is controversial, and other authorities have included it in a variety of othe
r
n
ematoceran
f
am
ili
es. Spec
i
es occur
i
n Nort
h
Amer
i
ca, eastern Europe, an
d
S
ib
er
i
a. A
d
u
l
ts
are stout-
b

o
di
e
dfli
es t
h
at resem
bl
eB
ibi
on
id
ae. Larvae
li
ve
i
ncav
i
t
i
es excavate
di
n rott
i
ng
w
o
od, perhaps feedin
g
on fun

g
i or other microor
g
anisms
.
249
T
HE P
A
N
O
RP
O
ID
ORDERS
F
I
G
URE 9.5. Ps
y
c
h
o
d
omorp
h
a. A mot
hfly,
P
s

y
chod
a
sp. (Ps
y
chodidae). [From F. R. Cole
and E. I. Schlinger, 1969,
T
he Flies o
f
Wester
n
Nort
h
America.
By
perm
i
ss
i
on o
f
t
h
eUn
i
vers
i
t
y

of California Press.
]
Infraor
d
er Psyc
h
o
d
omorp
h
a
A
b
out one
h
a
lf
o
f
t
h
e approx
i
mate
l
y 1000 spec
i
es
i
nt

hi
s group are
i
nc
l
u
d
e
di
nt
he
PSYCHODIDAE (moth flies) (Figure 9.
5
), a widely distributed group of small flies recog-
n
i
za
bl
e
by
t
h
e
h
a
i
r
y
w
i

n
g
s
h
e
ld
roo
flik
e over t
h
e
b
o
dy
w
h
en at rest. A
l
t
h
ou
gh
most spec
i
es
d
o not feed as adults, some females feed on blood, includin
g
those of the
g

enus Phlebotomus
(sand flies), s
p
ecies of which are vectors of various virus- and leishmania-induced diseases
.
T
RICHOCERIDAE (w
i
nter crane
fli
es) (110 spec
i
es, wor
ld
w
id
e) are eas
il
y con
f
use
d
w
i
t
h
th
e true crane
fli
es. T

h
ey carry t
h
e pre
f
erence
f
or coo
l
,mo
i
st
h
a
bi
tats to an extreme, an
d
man
y
spec
i
es are common
i
n caves an
d
m
i
nes. A
d
u

l
ts are o
f
ten encountere
di
n
l
ar
g
e swarm
s
d
urin
g
winter. The SCATOPSIDAE form a worldwide famil
y
of about 200 species of mostl
y
v
e
r
y
small flies. Larvae are found in deca
y
in
g
or
g
anic material, both plant and animal. Th
e

ANISOPODIDAE (100 species) is a worldwide group of primitive Diptera whose larvae
are
f
oun
di
n
d
ecay
i
ng an
df
erment
i
ng organ
i
c matter
.
In
f
raor
d
er Ptyc
h
opteromorp
h
a
This ver
y
small infraorder comprises two families, PTYCHOPTERIDAE (phanto
m

crane flies) (60 species), found in all except the Australian and neotropical regions, and
T
ANYDERIDAE (40 spec
i
es), a most
l
y Austra
li
an group o
f
crane
fl
y
lik
e
i
nsects. In
b
ot
h
f
am
ili
es
l
arvae are sem
i
aquat
i
c,

li
v
i
n
gi
nt
h
esu
b
strate at t
h
ee
dg
es o
f
streams
.
In
f
raor
d
er Cu
l
icomorp
ha
I
nc
l
u
d

e
di
nt
hi
s
l
arge group o
f
genera
ll
y sma
ll
an
dd
e
li
cate
fli
es are some extreme
l
y
w
e
ll
-
k
nown D
i
ptera. T
h

e group
i
s
di
v
id
e
di
nto
f
our
l
arge an
d
t
h
ree sma
ll f
am
ili
es
b
y
M
cA
l
p
i
n
e

e
ta
l.
(1981–1989). T
h
e
g
enera
lh
a
bi
ts o
f
t
h
ew
id
esprea
df
am
ili
es CULICIDAE
(mosquitoes) (3000 species) (Fi
g
ure 9.6) and CHAOBORIDAE (phantom mid
g
es) (75
species, often included as a subfamil
y
of Culicidae) (Fi

g
ure 9.7) present an interestin
g
contrast. Larval mosquitoes are filter feeders that strain microorganisms from the water i
n
whi
c
h
t
h
ey
li
ve. A
d
u
l
tma
l
es
d
o not
f
ee
d
,
b
ut
f
ema
l

es are vorac
i
ous
bl
oo
d
suc
k
ers an
d
a
s
suc
h
are respons
ibl
e
f
or t
h
e sprea
d
o
f
some
h
uman an
dli
vestoc
kdi

seases,
f
or examp
l
e
,
malaria,
y
ellow fever, filariasis, West Nile, and equine encephalitis. (Some, incidentall
y
,
also spread the m
y
xomatosis virus of rabbits and are, therefore, of some positive economi
c
value.) Larval Chaoboridae, on the other hand, are predators (particularly of mosquito lar
-
vae!). The adults, however, are nectar feeders. The DIXIDAE form a small (1
5
0s
p
ecies)
b
u
t
w
id
e
l
y

di
str
ib
ute
d
group t
h
at
i
s
f
requent
l
y cons
id
ere
d
asu
bf
am
il
yo
f
t
h
eCu
li
c
id
ae,

ma
i
n
ly
on t
h
e
b
as
i
so
f
t
h
ea
d
u
l
tw
i
n
g
venat
i
on an
d
t
h
es
i

m
il
ar
i
t
yb
etween t
h
e
l
arva
l
an
d
250
CHAPTER
9
F
I
GU
RE 9.6
.
Culicomorpha. Mosquitoes. (A) Anopheles
q
uadrimaculatus;
(
B
)
A
no

p
hele
s
sp. larva; (C
)
A
nop
h
e
l
e
s
s
p. pupa;
(
D
)
A
e
d
es cana
d
iensis;
(
E
)
C
u
l
ex pipiens; an

d
(F
)
C
u
l
e
x
s
p.
l
arva. [A, D, E,
f
rom
S. J. Car
p
enter and W. J. LaCasse, 19
55
, Mosquitoes of Nort
h
America. B
y
perm
i
ss
i
on o
f
t
h

eUn
i
vers
i
t
y
o
f
C
alifornia Press. A, E, drawn by Saburo Shibata. D, drawn by Kei Daishoji. B, C, F, from J. D. Gillett, 1971,
M
osquito
s
,
W
eidenfeld and Nicolson. By permission of the author.]
W
W
251
T
HE P
A
N
O
RP
O
ID
ORDER
S
F

I
GU
RE 9.7.
C
ulicomorpha. The clear lake gnat,
C
haoboru
s
a
stictopus (C
h
ao
b
or
id
ae). [From F. R. Co
l
ean
d
E. I. Sc
hli
nger
,
19
6
9
,
Th
eF
l

ies o
f
Western Nort
h
America. B
y
perm
i
ss
i
on o
f
t
he University of California Press.
]
pupal sta
g
es of the two
g
roups. The CHIRONOMIDAE (TENDIPEDIDAE) (Fi
g
ure 9.8)
constitute a lar
g
e, widel
y
distributed famil
y
of more than 5000 species. Adults are small
,

mosquitolike flies, though they do not feed. They often form massive swarms in the vicin
-
i
ty o
f
water. Larvae are aquat
i
can
d
e
i
t
h
er are
f
ree-
li
v
i
ng or
li
e
b
ur
i
e
di
nt
h
esu

b
strate,
mem
b
ers o
f
many spec
i
es construct
i
ng a spec
i
a
l
tu
b
e. T
h
e CERATOPOGONIDAE (
bi
t
i
n
g
mid
g
es, punkies, no-see-ums) (Fi
g
ure 9.9) form a widespread famil
y

of minute or small
fl
ies, man
y
females of which suck the blood of vertebrates and arthropods, or pre
y
o
n
other insects. Members of most s
p
ecies, however, feed on nectar and/or
p
ollen and render
cons
id
era
bl
e
b
ene
fi
tt
h
roug
h
cross-
f
ert
ili
zat

i
on o
f
t
h
ep
l
ants. Larvae occupy a var
i
ety o
f
mo
i
st
h
a
bi
tats,
i
nc
l
u
di
ng so
il
, moss, un
d
er
b
ar

k
,an
di
n roc
k
poo
l
s; t
h
ey may
b
ea
l
g
i
v
-
orous, saprop
h
a
g
ous, m
y
cop
h
a
g
ous, or pre
d
aceous. T

h
e approx
i
mate
ly
1100 spec
i
es o
f
SIMULIIDAE (black flies, buffalo
g
nats) (Fi
g
ure 9.10) form a widespread famil
y
, females
of which attack birds, mammals, and other insects. Several s
p
ecies of
S
imuliu
m
a
r
eo
f
e
x
-
t

reme
i
m
p
ortance as vectors
f
or t
h
e
fil
ar
i
a
l
nemato
d
e
,
Onc
h
ocerca vo
l
vu
l
us
,
whi
c
h
cause

s
onc
h
ocerc
i
as
i
s(r
i
ver
bli
n
d
ness)
i
n trop
i
ca
l
A
f
r
i
ca, Centra
l
Amer
i
ca, nort
h
ern Sout

h
Amer-
i
ca, an
d
Yemen. Ot
h
er spec
i
es transm
i
t nemato
d
e, protozoan, an
d
v
i
ra
l
pat
h
o
g
ens o
fbi
r
d
s
and mammals, includin
g

livestock. The larvae are found in swiftl
y
flowin
g
water, attached
F
I
GU
RE 9.8.
C
ulicomorpha.
C
hironomus tentans
(
Chironomi
-
d
ae). (A) Larva; an
d
(B) pupa. [From O. A. Jo
h
annsen, 1937
,
A
quat
i
cD
i
ptera. Part IV. C
hi

ronom
id
ae: Su
bf
am
ily
C
hi
ronom
i
-
n
ae, Mem. Cornell Univ. A
g
ri. Exp. Stn
.
2
1
0
:52 pp. By permission
o
f
Corne
ll
Un
i
vers
i
ty Agr
i

cu
l
ture Exper
i
menta
l
Stat
i
on.]
252
CHAPTER
9
FI
GU
RE 9.9. Cu
li
comor
ph
a. A
p
un
ki
e,
C
u
l
icoi
d
es
d

o
v
e
i
(
Ceratopo
g
o
-
nidae). [From F. R. Cole and E. I. Schlinger, 1969,
T
he Flies o
f
Western
Nort
h
America.
B
y perm
i
ss
i
on o
f
t
h
eUn
i
vers
i

ty o
f
Ca
lif
orn
i
a Press.]
F
IGURE 9.10.
C
u
li
comorp
h
a. A
bl
ac
kfl
y
,
S
imu
l
ium nigricoxum (S
i
mu
liid
ae). (A) Fema
l
e; (B) mature

l
arva;
an
d
(C) pupa. [From A. E. Cameron, 1922, T
h
e morp
h
o
l
o
gy
an
dbi
o
l
o
gy
o
f
a Cana
di
an catt
l
e-
i
n
f
est
i

n
gbl
ac
kfly
,
S
imulium simile
M
all. (Diptera, Simuliidae),
B
ulletin #5—New Series (Technical).
B
y permission of Agriculture
an
d
Agr
i
-Foo
d
Cana
d
a.]
to the substrate by an anal sucker, and are filter feeders or grazers. The THAUMALEIDAE
(
80 spec
i
es) const
i
tute a sma
ll

,pr
i
mar
il
y
h
o
l
arct
i
c
f
am
il
yo
f
m
i
nute m
id
ges w
h
ose a
ffi
n
i
t
i
e
s

are uncerta
i
n. In some
f
eatures
i
ts mem
b
ers resem
bl
et
h
eot
h
er Cu
li
comorp
h
a,
i
not
h
ers t
h
e
B
ibi
onomorp
h
a

.
In
f
raor
d
er Bi
b
ionomorp
ha
T
his lar
g
e and diverse
g
roup includes four ma
j
or families. The BIBIONIDAE (Marc
h
fli
es) (700 spec
i
es) (F
i
gure 9.11A) are ro
b
ust,
h
a
i
ry

fli
es o
f
me
di
um to sma
ll
s
i
ze. Large
swarms, cons
i
st
i
ng
l
arge
l
yo
f
ma
l
es, are seen
i
nt
h
e Nort
h
ern Hem
i

sp
h
ere spr
i
ng (
h
ence, t
h
e
c
ommon name). Larvae o
f
t
hi
s cosmopo
li
tan
g
roup
f
ee
dg
re
g
ar
i
ous
ly
on roots or
d

eca
yi
n
g
ve
g
etation. The MYCETOPHILIDAE (2000 species) and SCIARIDAE (500 species) ar
e
253
T
HE P
A
N
O
RP
O
ID
ORDER
S
F
IGURE 9.11
.
B
ibi
onomorp
h
a. (A)
A
M
arc

hfly
,
B
i
b
io a
lb
ipenni
s
(
B
ibi
on
id
ae
);
and (B) the Hessian fly
,
Ph
y
topha
g
a destruc-
to
r
(Cec
idi
omy
iid
ae). [A,

f
rom D. J. Borror
an
d
D. M. De
l
on
g
, 1971
,
An Intro
d
uction
to the Study o
f
Insects
,
3rd ed. By permis-
si
on o
f
Broo
k
s/Co
l
e, a
di
v
i
s

i
on o
f
T
h
omson
L
earn
i
n
g
.B,
f
rom L. A. Swan an
d
C. S
.
P
app, 1972,
T
he Common Insects o
f
Nort
h
A
merica.
C
opyr
i
g

h
t 1972
b
yL.A.Swa
n
an
d
C. S. Papp. Repr
i
nte
dby
perm
i
ss
i
on o
f
Harper & Row, Publishers, Inc.]
sometimes included in a sin
g
le famil
y
of the former name. The
y
are commonl
y
known a
s
fun
g

us
g
nats from the observation that the larvae feed mainl
y
on fun
g
i and deca
y
in
g
p
l
ant mater
i
a
l
.A
d
u
l
ts are common
l
y encountere
di
n coo
l
,
d
amp s
i

tuat
i
ons. T
he
CECIDOMYIIDAE (ga
ll
m
id
ges)
f
orm a very
l
arge
f
am
il
y (4000 spec
i
es) o
f
m
i
nute
fli
es
,
most o
f
w
hi

c
hf
ee
d
,
i
nt
h
e
l
arva
l
sta
g
e, on p
l
ant t
i
ssues,
f
requent
ly
caus
i
n
g
t
h
e
f

ormat
i
on o
f
g
alls. There are, however, sapropha
g
ous or predaceous species. Within the famil
y
are severa
l
economicall
y
important species, for example, the Hessian fl
y,
Phytophaga
(
=
M
ayetiola
)
destructor (Figure 9.11B), whose larvae feed on wheat shoots. Many species are paedoge-
net
i
c, t
h
e
f
u
ll

-grown
l
arvae
b
ecom
i
ng sexua
ll
y mature an
d
repro
d
uc
i
ng part
h
enogenet
i
ca
ll
y.
As t
h
e young
l
arvae grow, t
h
ey
d
evour t

h
e
i
r parent
f
rom w
i
t
hi
n. Severa
l
generat
i
ons o
f
pae-
d
o
g
enetic larvae ma
y
develop in a season, and the larval population can thus increase
enormousl
y
. Eventuall
y
, the larvae pupate normall
y
and sexual reproduction follows.
S

u
b
or
d
er Brac
hy
cera
Most Brach
y
cera are rather stout flies with antennae havin
g
fewer than seven se
g-
ments an
d
,o
f
ten, an ar
i
sta; max
ill
ary pa
l
ps are unsegmente
d
or two-segmente
d
. Larvae ar
e
h

em
i
cep
h
a
l
ous or acep
h
a
l
ous (maggot
lik
e), w
i
t
h
s
i
c
kl
e-s
h
ape
d
man
dibl
es t
h
at move
i

nt
he
vert
i
ca
l
p
l
ane.
Members of this suborder were traditionall
y
placed in two sub
g
roups, Orthorrhapha
(Brach
y
cera sensu str
i
ct
o
)
and C
y
clorrhapha, which were sometimes each
g
iven subor
-
d
inal rank. It is now clear that the Orthorrhapha is a paraphyletic group, comprising th
e

i
n
f
raor
d
ers Ta
b
anormorp
h
aan
d
As
il
omorp
h
a, an
d
t
h
at t
h
ecyc
l
orr
h
ap
h
ous
f
orms, a

l
t
h
oug
h
monop
h
y
l
et
i
c, mer
i
ton
l
y
i
n
f
raor
di
na
l
status (Muscomorp
h
a)
.
Infraorder Tabanomorpha
As constituted b
y

McAlpine
et al.
(1981–1989), the Tabanomorpha includes seven
families, three of which contain between them about 95% of the species. The lar
g
est famil
y
is the TABANIDAE (Figure 9.12A), with more than 3000 species, which includes those
bl
oo
d
suc
ki
ng
i
nsects common
l
y
k
nown as
h
orse an
dd
eer
fli
es, c
l
egs, Marc
hfli
es (

i
nt
h
e
Sout
h
ern Hem
i
sp
h
ere), an
d
pro
b
a
bl
y many ot
h
er,
l
ess po
li
te names! T
h
e
bl
oo
d
suc
k

ers
b
elon
g
to onl
y
three
g
enera
,
Tabanus, Chr
y
sops,
a
n
d
H
aemato
p
ota,
w
hose evolution ha
s
254
CHAPTER
9
F
IGURE 9.12
.
Tabanomorpha. (A) A horse fly, Tabanus o

p
acus
(
Tabanidae); and (B) a soldier fly,
O
dontom
y
ia
h
oo
d
ian
a
(
Stratiom
y
idae). [A, from J. F. McAlpine, 19
6
1, Variation, distribution and evolution of th
e
Ta
b
anus
(Hybomitra)
f
rontalis
c
om
p
lex of horse flies (Di

p
tera: Tabanidae),
C
an. Entomol. 9
3
:894–924. B
y
permission o
f
the Entomological Society of Canada. B, from F. R. Cole and E. I. Schlinger, 1969, The Flies of Western Nort
h
A
merica
.
B
y
perm
i
ss
i
on o
f
t
h
eUn
i
vers
i
t
y

o
f
Ca
lif
orn
i
a Press.
]
c
losel
y
followed that of the hoofed mammals. Althou
g
h the
y
are known to be capable
o
f transmitting various diseases both human and of livestock, tabanids cause far greate
r
e
conom
i
c
l
osses
b
yt
h
e
i

r
di
stur
b
ance an
di
rr
i
tat
i
on o
fli
vestoc
k
, resu
l
t
i
ng
i
n
l
ower y
i
e
ld
s
of
m
ilk

an
d
meat. On
l
y
f
ema
l
e
h
orse
fli
es suc
kbl
oo
d
,
i
nt
h
ea
b
sence o
f
w
hi
c
h
t
h

ey
f
ee
d,
l
ike males and like members of most tabanid species, on nectar and pollen. Larvae mostl
y
o
ccur in mud, deca
y
in
g
ve
g
etation, and shallow water (movin
g
or still) where the
y
pre
y
o
n other invertebrates. Another lar
g
e and well-distributed famil
y
is the STRATIOMYI-
DAE
(
Figure 9.12B), containing some 1500 species, commonly known as soldier flies. The
w

ea
kl
y
fl
y
i
ng a
d
u
l
ts are encountere
d
among
l
ow-grow
i
ng
h
er
b
age an
d
are most pro
b
a
bl
y
n
ectar
f

ee
d
ers. Mem
b
ers o
f
man
y
spec
i
es are consp
i
cuous
ly
str
i
pe
d
,an
d
some spec
i
es are
w
asp mimics. Larvae, often
g
re
g
arious, are aquatic or terrestrial, occurrin
g

in deca
y
in
g
o
r
g
anic matter, in dun
g
, or under bark. Some are scaven
g
ers, others are predaceous or ph
y-
top
h
agous, t
h
e
l
atter group
i
nc
l
u
di
ng pests o
fl
awns an
d
sugarcane. T

h
e RHAGIONIDA
E
(
sn
i
pe
fli
es) are per
h
aps t
h
e most pr
i
m
i
t
i
ve Brac
h
ycera. T
h
e
f
am
il
y, w
hi
c
h

conta
i
ns mor
e
t
h
an 300 extant spec
i
es, as we
ll
as
f
oss
il
s
f
rom t
h
e Upper Jurass
i
c,
i
sw
id
e
l
y
di
str
ib

ute
d
,
thou
g
h seldom encountered, because of the secretive, solitar
y
habits of adult flies. Mem-
bers of man
y
species are nectar feeders, others are predaceous, and females of some species
suck blood. Larvae occur in damp soil, rotting wood, etc. and are believed to prey on other
i
nsects
.
Infraor
d
er Asi
l
omorp
h
a
Th
e12
f
am
ili
es
i
nt

hi
s very
l
arge taxon
f
a
ll i
nto t
h
ree we
ll
-
d
e
fi
ne
d
super
f
am
ili
es,
A
s
il
o
id
ea, Bom
b
y

li
o
id
ea, an
d
Emp
id
o
id
ea. A
b
out 80% o
f
t
h
eas
il
o
id
s
b
e
l
ong to t
h
e cos-
m
opolitan famil
y
ASILIDAE (robber flies) (Fi

g
ure 9.13A), which with about
5
000 species
i
s amon
g
the lar
g
est of the orthorrhaphous Brach
y
cera. Adults suck the bod
y
fluids of
a
v
ariet
y
of other insects. The
y
are powerful fliers and catch their pre
y
on the win
g
, have well
-
d
eve
l
ope

d
eyes an
d
some
d
egree o
f
stereoscop
i
cv
i
s
i
on, possess strong
l
egs
f
or grasp
i
ng t
h
e
prey an
d
are usua
ll
y
h
a
i

ry, espec
i
a
ll
y aroun
d
t
h
e
f
ace, a
f
eature t
h
at per
h
aps protects t
h
em
durin
g
the stru
gg
le. THEREVIDAE form a widel
y
distributed
g
roup of about
5
00 species

that
g
enerall
y
resemble robber flies, thou
g
h the
y
are not predaceous, feedin
g
instead o
n
255
T
HE P
A
N
O
RP
O
ID
ORDER
S
F
IGURE 9.13
.
As
il
omorp
h

a. (A) A ro
bb
er
fl
y
,
M
a
ll
op
h
orina pu
l
c
h
ra
(
As
ilid
ae); an
d
(B) a
b
ee
fl
y, P
oecila
n
th
r

ax
a
utumna
l
is
(
Bomb
y
liidae). (From F. R. Cole and E. I. Schlin
g
er, 19
6
9, T
h
eF
l
ies of Western Nort
h
America
.
By
p
ermission of the University of California Press.
]
nectar, plant exudates, etc. Therevid larvae burrow in soil, rottin
g
bark, fun
g
i, and rot
-

ti
ng
f
ru
i
t, an
d
are vorac
i
ous pre
d
ators, espec
i
a
ll
yo
fb
eet
l
e
l
arvae an
d
eart
h
worms. O
f
t
he
B

om
b
y
li
o
id
ea, some 4000 spec
i
es
b
e
l
ong to t
h
e BOMBYLIIDAE (
b
ee
fli
es) (F
i
gure 9.13B).
Th
e common name o
f
t
h
ese nectar
f
ee
d

ers
h
as
d
ou
bl
es
ig
n
ifi
cance. F
i
rst, t
h
e
fli
es resem
bl
e
b
umblebees, and second, in man
y
species, female flies deposit e
gg
s at the nest entrance o
f
a solitar
y
bee or wasp so that the larvae ma
y

feed on the pollen, hone
y
, and even
y
oun
g
Hymenoptera. Larvae o
f
ot
h
er spec
i
es searc
hf
or an
df
ee
d
on grass
h
opper eggs, t
h
ere
b
yp
l
ay
-
i
ng an

i
mportant ro
l
e
i
nt
h
e natura
l
regu
l
at
i
on o
f
grass
h
opper popu
l
at
i
ons. In a
f
ew spec
i
e
s
e
gg
s are

l
a
id di
rect
ly
onto
l
arvae o
f
Lep
id
optera or H
y
menoptera. T
h
e NEMESTRINIDAE
(250 species) and ACROCERIDAE (450) species are both widespread and ancient
g
roups,
sometimes considered to be Muscomor
p
ha. In both families adults are found around flowers
an
d
some
f
ee
d
on nectar (ot
h

ers apparent
l
y
d
o not
f
ee
d
as t
h
ey
h
ave vest
i
g
i
a
l
mout
h
parts).
Th
e
l
arvae, w
hi
c
h
un
d

ergo
h
eteromorp
h
os
i
s (see C
h
apter 21, Sect
i
on 3.3.2), are paras
i
to
ids
o
fg
rass
h
oppers an
dl
ocusts (an
d
ma
yb
ean
i
mportant
f
actor
i

n contro
lli
n
g
t
h
e
i
r popu
l
a-
t
ions) and of spiders, respectivel
y
. The Empidoidea contain two lar
g
e families of advance
d
B
rach
y
cera, the EMPIDIDAE (dance flies, balloon flies) and DOLICHOPODIDAE (lon
g-
legged flies). There are about 3000 species of Empididae, which are predaceous in bot
h
a
d
u
l
tan

dj
uven
il
e stages. T
h
e
f
am
il
y
i
s
l
arge
l
y restr
i
cte
d
to t
h
e temperate reg
i
ons o
fb
ot
h
h
em
i

sp
h
eres, an
di
ts mem
b
ers ga
i
nt
h
e
i
r common name
f
rom t
h
e courts
hi
p
b
e
h
av
i
or o
f
most, thou
g
hb
y

no means all, species. In man
y
species males have an elaborate courtshi
p
d
ispla
y
in which a female is offered a
g
ift of food (real or imitation). Presumabl
y
this serve
d
originally as an insurance policy against the male’s life. Dolichopodidae, which constitut
e
af
amily of about 6000 species, appear to be a specialized offshoot of the empid line. They
ff
are genera
ll
y
f
oun
di
n coo
l
,mo
i
st
h

a
bi
tats,
i
nc
l
u
di
ng t
h
e seas
h
ore an
d
sa
l
t mars
h
es. A
d
u
l
ts
an
d
most
l
arvae are pre
d
aceous, espec

i
a
lly
on ot
h
er D
i
ptera;
l
arvae o
f
a
f
ew spec
i
es ar
e
ph
y
topha
g
ous.
Infraorder Muscomorpha
T
h
e Muscomorp
h
a are arrange
di
n two ser

i
es. T
h
e
fi
rst, Asc
hi
za,
i
s per
h
aps a
pol
y
ph
y
letic
g
roup, containin
g
the more primitive members, which lack a ptilinal su
-
t
ure. The second, Schizo
p
hora, includes those flies in which a
p
tilinal suture is
p
resent

.
256
CHAPTER
9
In older classifications a third series, Pupiparia, was often found. McAlpine (in McAlpine
et al.,
1
981–1989), however, has presented stron
g
reasons for considerin
g
this
g
roup as a
m
onoph
y
letic superfamil
y
, Hippoboscoidea, within the cal
y
ptrate Schizophora. The ma
j
or
-
i
ty o
f
Sc
hi

zop
h
ora (
i
.e., t
h
ose t
h
at are w
i
nge
d
as a
d
u
l
ts) can
b
e arrange
di
ntwosu
bdi
v
i
s
i
ons,
t
h
eCa

l
yptratae, w
hi
c
h
conta
i
ns
fli
es t
h
at possess a ca
l
ypter, or
l
o
b
e, at t
h
e
b
ase o
f
t
h
e
f
ore
wi
n

g
t
h
at covers t
h
e
h
a
l
teres, an
d
t
h
e Aca
ly
ptratae, w
h
ose mem
b
ers
h
avenosuc
hl
o
b
e.
Thou
g
h this division is a natural one, that is, it represents a true evolutionar
y

diver
g
ence
,
i
t should be reco
g
nized that (1) some
g
roups have secondaril
yg
ained or lost the cal
y
pter
,
an
d
(2) t
h
ea
d
u
l
ts o
f
some paras
i
t
i
c

f
am
ili
es are w
i
ng
l
ess, t
h
oug
h
t
h
e
i
ra
ffi
n
i
t
i
es are c
l
ear
ly
ei
t
h
er ca
l

yptrate or aca
l
yptrate.
Series Asc
h
i
za
S
uper
f
ami
l
yP
l
atypezoi
d
ea
T
his superfamil
y
has one lar
g
e, cosmopolitan famil
y
, PHORIDAE (2700 species)
,
an
d
severa
l

sma
ll
to very sma
ll
ones, some o
f
w
hi
c
h
are o
f
ten
i
nc
l
u
d
e
d
w
i
t
h
t
h
ep
h
or
id

s
.
Al
t
h
oug
h
many p
h
or
id
s are
f
ree-
li
v
i
ng,
f
u
ll
yw
i
nge
dfli
es
f
oun
d
among

l
ow vegetat
i
on,
o
n or near
d
eca
yi
n
g
or
g
an
i
c matter, on
f
un
gi
,
i
n
bi
r
d
nests, etc., t
h
e
y
seem to pre

f
er t
o
run rather than fl
y
, a feature that foreshadows the brach
y
pterous or apterous condition o
f
the man
y
species that live under
g
round, as inquilines or parasites in ant and termite nests.
Larvae are maggot
lik
e, w
i
t
hdi
verse
h
a
bi
ts. Some are scavengers on
f
ung
i
(an
d

may
b
ecome
pests on mus
h
room
f
arms), carr
i
on, an
dh
uman corpses; ot
h
ers are paras
i
tes o
f
eart
h
worms,
o
ther insects, spiders, and m
y
riapods. PLATYPEZIDAE (2
5
0 species) form a worldwide
g
roup whose members prefer shaded woodland where the
y
feed on nectar. Larvae ar

e
f
un
g
ivorous.
S
uperfamily Syrphoidea
O
ne sma
ll
an
d
one ver
yl
ar
g
e
f
am
ily
ma
k
eupt
h
eS
y
rp
h
o
id

ea. T
h
e PIPUNCULIDAE
(
bi
g
-headed flies) (400 species) are a cosmopolitan
g
roup of small humpbacked flies with
l
ar
g
e heads covered almost entirel
y
b
y
the compound e
y
es. Adults tend to be found hoverin
g
ov
er
flo
w
ers; t
h
e
l
arvae are en
d

o
p
aras
i
tes o
fh
omo
p
terans an
d
,assuc
h
, are
i
m
p
ortant natura
l
c
ontrol agents. The SYRPHIDAE (
5
000 species worldwide) (Figure 9.14) are the well
-
k
nown
h
over
fli
es. T
h

e
yf
orm one o
f
t
h
e
l
ar
g
est an
d
most eas
ily
reco
g
n
i
ze
dg
roups o
f
F
I
GU
RE 9.14
.
S
y
rp

h
o
id
ea. A
h
over
fly
,
E
upeo
d
es vo
l
ucris
(
S
y
r
-
p
hidae). [From a drawing by Charles S. Papp. By permission of the
art
i
st.
]
2
5
7
T
HE P

A
N
O
RP
O
ID
ORDER
S
Diptera. The
y
are
g
enerall
y
bri
g
htl
y
colored, often striped, and man
y
mimic bees or wasps.
I
n some species there are obvious reasons for the preciseness of this mimicr
y
, for the hove
r
fly
la
y
s its e

gg
s in the nests of H
y
menoptera and, because of its similarit
y
, presumabl
y
a
v
o
id
s
d
etect
i
on. For ot
h
er s
p
ec
i
es t
h
e reason
i
s
l
ess o
b
v

i
ous, an
d
no re
l
at
i
ons
hip i
sa
pp
arent
b
etween t
h
em
i
m
i
can
di
ts mo
d
e
l
. In contrast to t
h
e rat
h
er un

if
orm, nectar-
f
ee
di
ng
h
a
bi
ts
o
f
a
d
u
l
t
h
over
fli
es, t
h
ose o
fl
arvae are extreme
ly
var
i
e
d

,p
hy
top
h
a
g
ous, zoop
h
a
g
ous, an
d
sapropha
g
ous species bein
g
known.
S
eries Schizo
p
hora
Subdivision Acal
y
ptrata
e
Super
f
ami
l
y Conopoi

d
ea
T
h
e super
f
am
il
y Conopo
id
ea, t
h
e most pr
i
m
i
t
i
ve o
f
t
h
eSc
hi
zop
h
ora, conta
i
ns t
h

es
i
n-
g
l
e, w
id
esprea
df
am
il
y CONOPIDAE (800 spec
i
es) w
h
ose mem
b
ers typ
i
ca
ll
ym
i
m
i
c wasps
an
db
ees. A
d

u
l
ts are nectar
f
ee
d
ers an
d
are espec
i
a
lly
assoc
i
ate
d
w
i
t
hfl
owers o
f
Compos
i
tae,
L
abiatae, and Umbelliferae. Cono
p
ids are
p

arasites of bees and was
p
s, cockroaches, and
cal
y
ptrate Diptera, the female catchin
g
the host in fli
g
ht and depositin
g
an e
gg
directl
y
o
n
i
ts
b
o
d
y.
Super
f
ami
l
yTep
h
ritoi

d
ea
This
g
roup includes ei
g
ht families in the scheme of McAlpine
et al.
(1981–1989) of
whi
c
h
two are
l
arge an
d
t
h
ree are o
f
me
di
um s
i
ze. T
h
e OTITIDAE (p
i
cture-w
i

nge
dfli
es) are
pre
d
om
i
nant
l
y a nort
h
-temperate group o
f
some 400 spec
i
es. A
d
u
l
ts are common
i
n
d
ense
ve
g
etat
i
on;
l

arvae are t
y
p
i
ca
lly
saprop
h
a
g
ous, t
h
ou
gh
a
f
ew are p
hy
top
h
a
g
ous,
i
nc
l
u
di
n
g

pests of onions and su
g
ar beet. PLATYSTOMATIDAE (1000 species) are worldwide bu
t
most common in Africa, Australia, and Asia. Both adults and larvae resemble members of the
previous family in morphology and habits. The cosmopolitan PYRGOTIDAE (330 species
)
are typ
i
ca
ll
y nocturna
lfli
es t
h
at paras
i
t
i
ze scara
b
ae
id b
eet
l
es. Fema
l
es
l
an

d
on t
h
e
b
eet
l
es
i
n
fli
g
h
tan
d
ov
i
pos
i
tont
h
et
hi
na
bd
om
i
na
l
terg

i
tes
b
eneat
h
t
h
ee
l
ytra. LONCHAEIDA
E
(
5
00 species worldwide)
g
enerall
y
occur in forests; larvae feed on rottin
g
plant material,
rarel
y
flower heads and root crowns. The lar
g
est and best known famil
y
, with some 4000
species, is the TRYPETIDAE (TEPHRITIDAE), the fruit flies, a group that includes some
ma
j

or agr
i
cu
l
tura
l
pests. T
h
e
i
r
l
arvae
f
ee
d
onavar
i
ety o
f
p
l
ant mater
i
a
l
s. T
h
ey may
b

e
l
ea
f
or stem m
i
ners, ga
ll f
ormers,
fl
ower-
i
n
h
a
bi
t
i
ng spec
i
es, or
f
ru
i
tan
d
see
d
eaters. In t
h

e
l
atte
r
cate
g
or
y
are t
h
eMe
di
terranean
f
ru
i
t
fly
,
C
eratitis ca
p
itata
(
Fi
g
ure 9.1
5
), which attacks
citrus and other fruits, an

d
Rhagoletis pomonella, the apple ma
gg
ot fl
y
, whose larvae tunnel
into a
pp
les,
p
ears, etc
.
Super
f
amily Nerioidea
Nerioidea (Micropezoidea) form a small
g
roup of three families, the lar
g
est of which
is the MICROPEZIDAE (500 species), commonl
y
called stilt flies because of their lon
g
l
egs. Mem
b
ers o
f
t

hi
s
b
as
i
ca
ll
y trop
i
ca
lf
am
il
y are
f
oun
di
nwoo
d
e
d
areas; t
h
e
i
r
l
arvae are
pr
i

mar
il
y saprop
h
agous, t
h
oug
h
a
f
ew p
h
ytop
h
agous spec
i
es may
b
ecome pests (e.g., o
f
gi
n
g
er an
dl
e
g
umes)
.
258

CHAPTER
9
FI
GU
RE 9.15. Tephritoidea. The Mediterranean fruit fl
y,
C
eratitis
c
a
p
itat
a
(Tephritidae). [From F. R. Cole and E. I. Schlinger, 1969
,
Th
e
F
l
ies of Western Nort
h
America
.
By
perm
i
ss
i
on o
f

t
h
eUn
i
vers
i
t
y
o
f
C
alifornia Press.]
S
uper
f
amily Diopsoidea
D
iopsoidea (Tan
y
pezoidea) make up another small superfamil
y
whose members mostl
y
f
all into the PSILIDAE (200 mostl
y
holarctic species) and DIOPSIDAE (150 mostl
y
tropical
s

p
ecies). Psilid larvae feed on roots and stems and a few are
p
ests, for exam
p
le,
P
sila rosa
e
(
carrot rust
fl
y), on carrots, ce
l
ery, an
d
ot
h
er root crops. A
d
u
l
t
di
ops
id
s are ca
ll
e
d

sta
lk
-eye
d
fli
es
b
ecause o
f
t
h
e
l
atera
l
extens
i
ons o
f
t
h
e
h
ea
d
w
hi
c
hb
ear t

h
e compoun
d
eyes; t
h
e
i
r
l
arvae are sapropha
g
ous or ph
y
topha
g
ous (especiall
y
on Graminae), the latter occasionall
y
becomin
g
minor pests
.
S
uper
f
ami
l
y Sciomyzoi
d

e
a
M
ost members of this small superfamily belong to the cosmopolitan families
S
CIOMYZIDAE (
55
0 species) or SEPSIDAE (240 species). Sciomyzid larvae feed o
n
terrestr
i
a
l
or aquat
i
cMo
ll
usca, a
li
ve or
d
ea
d
,oront
h
e
i
r eggs an
d
em

b
ryos. Larvae o
f
Sep
-
sidae scaven
g
eindun
g
, includin
g
sewa
g
e slud
g
e, or in deca
y
in
g
plant or animal material
.
S
uper
f
amily Lauxanioide
a
L
auxanioids were formerl
y
included in the previous superfamil

y
. The
g
reat ma
j
orit
y
o
f species belong to the LAUXANIIDAE (1200 mainly tropical species), adults of whic
h
are se
d
entary an
d
co
ll
ect on
l
ow-grow
i
ng vegetat
i
on, espec
i
a
ll
ya
dj
acent to water. Larvae
are saprop

h
agous, occurr
i
ng
i
n
d
ecay
i
ng vegetat
i
on,
l
ea
fli
tter, an
dbi
r
d
nests
.
S
uper
f
ami
l
y Opomyzoi
d
e
a

T
his is possibl
y
a pol
y
ph
y
letic
g
roup that, as constituted b
y
McAlpin
e
et al.
(1981–
1
989),
i
nc
l
u
d
es a
b
out a
d
ozen
f
am
ili

es. Ot
h
er aut
h
or
i
t
i
es se
p
arate t
h
e
f
am
ili
es
i
nto t
h
ree
super
f
am
ili
es, Opomyzo
id
ea
s
en

s
u
s
tricto, Agromyzo
id
ea, an
d
Aste
i
o
id
ea. Most o
f
t
h
e
f
am
ili
es are very sma
ll
an
dh
ave a restr
i
cte
ddi
str
ib
ut

i
on. A
d
u
l
t CLUSIIDAE (220 spec
i
es
w
orldwide) occur around rottin
g
lo
g
s and feed on nectar, exudates of rottin
g
material, etc.
Their larvae are found in rottin
g
wood, and in the tunnels of termites and bark beetles. Som
e
1
800 species of AGROMYZIDAE are known, including pests of shade trees, vegetables
,
an
dfl
owers. Larvae o
f
t
hi
s cosmopo

li
tan
f
am
il
y are most
l
y
l
ea
f
or stem m
i
ners; some
f
ee
din
see
d
s,
b
ore
i
nwoo
d
, or are ga
ll f
ormers. ASTEIIDAE (100 spec
i
es) are w

id
e
l
y
di
str
ib
ute
d.
Larvae of these tin
y
flies appear to be scaven
g
ers in rottin
g
plants or fun
g
i
.
2
5
9
T
HE P
A
N
O
RP
O
ID

ORDERS
Superfamily Carnoidea
T
h
ere
l
at
i
ons
hi
ps o
f
t
h
en
i
ne
f
am
ili
es
i
nc
l
u
d
e
di
nt
h

e Carno
id
ea (C
hl
oropo
id
ea) rema
i
n
d
ebatable, and frequentl
y
some of the families are placed in the Opom
y
zoidea or Eph
y-
d
roidea. The
g
reat ma
j
orit
y
of species belon
g
to the cosmopolitan families MILICHIIDAE
(300 s
p
ecies) and CHLOROPIDAE (2000 s
p

ecies). Adult milichiids are found at flowers;
some spec
i
es r
id
eonsp
id
ers an
d
pre
d
atory
i
nsects suc
h
as as
ilid fli
es an
d
re
d
uv
iid b
ugs,
h
e
l
p
i
ng t

h
emse
l
vestot
h
e
j
u
i
ces t
h
at exu
d
e
f
rom t
h
e
i
r
h
ost’s prey. Larvae are saprop
h
agou
s
or
d
un
gf
ee

d
ers,
i
nc
l
u
di
n
g
some t
h
at
li
ve
i
nt
h
e
f
un
g
us
g
ar
d
ens o
fl
ea
f
-cutt

i
n
g
ants. A
d
u
lt
chloropids t
y
picall
y
occur in vast numbers on folia
g
e, and mostl
y
feed on sap exudates,
h
one
y
dew, etc. However, the so-called e
y
e
g
nats (species of
H
ippelete
s
a
n
d

Siphunculina
)
are attracte
d
to woun
d
san
d
secret
i
ons o
f
t
h
e eyes, nose,
li
ps, an
d
s
ki
nan
d
are vectors o
f
con
j
unct
i
v
i

t
i
san
d
s
ki
n
di
seases. Larvae may
b
e saprop
h
agous or p
h
ytop
h
agous an
d
some o
f
th
e
l
atter are
i
mportant pests,
f
or examp
l
e

,
Oscine
ll
a fri
t
(
European frit fl
y
) (Fi
g
ure 9.1
6
B)
on cereals
.
Superfamily Sphaeroceroide
a
I
nc
l
u
d
e
di
nt
h
eSp
h
aerocero
id

ea (He
l
eom
y
zo
id
ea) are t
h
e cosmopo
li
tan
f
am
ili
es
SPHAEROCERIDAE (700 species, includin
g
about 120 that are brach
y
pterous or apterous
)
and HELEOMYZIDAE (400 species), to
g
ether with several small
g
roups. Sphaerocerids
are associated with dung, decaying plant material, fungi (occasionally becoming pests on
mus
h
room

f
arms), seawee
d
,an
d
carr
i
on. Larva
lh
e
l
eomyz
id
s, too, are
f
oun
di
n
d
ecay
i
n
g
organ
i
c matter an
df
ung
i
,aswe

ll
as
i
nt
h
e nests o
fbi
r
d
san
d
mamma
l
s, an
db
at caves
.
Super
f
ami
l
yEp
h
y
d
roi
d
ea
Also called Drosophiloidea, this group includes about 2600 species, almost all of which
fall into two large families. The EPHYDRIDAE (shore flies, brine flies) (Figure 9.16A),

wi
t
h
more t
h
an 1000
d
escr
ib
e
d
spec
i
es,
i
swe
ll k
nown
b
ecause o
f
t
h
e remar
k
a
bl
evar
i
ety

of habitats that its members occup
y
. The
y
are t
y
picall
y
found near water, both fresh an
d
F
I
GU
RE 9.16.
E
phydroidea and Carnoidea. (A) An ephydrid with raptorial forelegs
,
O
chthera manti
s
(Ephy-
d
r
id
ae); an
d
(B) t
h
e European
f

r
i
t
fl
y
,
O
scine
ll
a frit (Chloropidae). [A, from F. R. Cole and E. I. Schlinger, 19
6
9,
T
h
eF
l
ies of Western Nort
h
America.
By
perm
i
ss
i
on o
f
t
h
eUn
i

vers
i
t
y
o
f
Ca
lif
orn
i
a Press. B,
f
rom L. A. Swa
n
a
nd C. S. Papp, 1972
,
T
he Common Insects o
f
North America. Copyright 1972 by L. A. Swan and C. S. Papp.
Repr
i
nte
db
y perm
i
ss
i
on o

f
Harper & Row Pu
bli
s
h
ers, Inc.]
260
CHAPTER
9
salt, and in man
y
species the larvae are trul
y
aquatic. Larvae of some species feed on al
g
ae
,
but
g
enerall
y
the
y
and the adults are carnivorous or carrion feeders, sometimes almos
t
to the point of bein
g
parasitic. Two examples, illustratin
g
the extreme habitats in which

Ep
h
y
d
r
id
ae are
f
oun
d
,ar
e
E
p
hyd
ra ripari
a
,
w
hi
c
hi
s
f
oun
di
nt
h
e Great Sa
l

tLa
k
eo
f
Uta
h
(
see a
l
so C
h
apter 18, Sect
i
on 4.3), an
d
Psi
l
opa (He
l
aeom
y
ia) petro
l
ei
,
w
h
ose
l
arvae

li
ve
i
n
poo
l
so
f
cru
d
e petro
l
eum
i
nCa
lif
orn
i
a. T
h
ec
l
ose
ly
re
l
ate
d
DROSOPHILIDAE (pomace o
r

f
ruit flies) are small flies
g
enerall
y
seen in the vicinit
y
of deca
y
in
g
ve
g
etation or fruit, or
n
ear breweries and vine
g
ar factories. The larvae are mostl
y
fun
g
ivorous, thou
g
hafewar
e
l
ea
f
m
i

ners or prey on ot
h
er
i
nsects. Var
i
ous spec
i
es o
f
D
rosop
h
i
la
h
ave, o
f
course,
b
een
e
xtens
i
ve
l
y use
df
oraw
id

e range o
fbi
o
l
og
i
ca
l
researc
h.
S
u
bd
ivision Ca
ly
ptrata
e
How the Cal
y
ptratae should be subdivided remains debatable; some authors lump al
l
f
am
ili
es
i
nt
h
e Musco
id

ea w
hil
eot
h
ers,
i
nc
l
u
di
ng McA
l
p
i
ne et a
l.
(1981–1989),
b
e
li
eve
t
h
at t
h
ere are t
h
ree monop
h
y

l
et
i
csu
b
groups
i
nt
h
esu
bdi
v
i
s
i
on, Musco
id
e
a
s
en
s
u
s
tricto
,
O
estro
id
ea, an

d
H
i
ppo
b
osco
id
ea (
=
P
up
i
par
i
ao
f
ear
li
er aut
h
ors).
S
uper
f
ami
l
y Muscoi
d
e
a

In the cosmopolitan family MUSCIDAE (3000 species) are many common pests,
f
or examp
l
e, t
h
e Austra
li
an
b
us
hfl
y(
M
usca
v
etust
i
ss
i
ma
)
,
h
ouse
fli
es
[
M
u

s
ca
d
ome
s-
tica (F
i
gure 9.17A) an
d
Fannia canicu
l
ari
s
], an
dbl
oo
d
suc
ki
ng spec
i
es suc
h
as t
h
e sta
bl
e
fl
y(

Stomox
y
s calcitran
s
)
and face fl
y
(
M
u
s
ca autumnali
s
)
. However, the
g
reat ma
j
or
-
i
t
y
of species are non-pestiferous. Adults are predaceous, sapropha
g
ous, pollenopha
g
ous,
hematopha
g

ous, or feed on exudates of mammals. Females are t
y
picall
y
oviparous, thou
gh
af
ew
f
f
a
re ovov
i
v
i
parous or
l
arv
i
parous. Larvae are most
l
y saprop
h
agous or
d
ung
f
ee
d
ers,

t
h
oug
h
some are pre
d
aceous an
d
a
f
ew are ectoparas
i
tes on
bi
r
d
s. Most o
f
t
h
e 1000 spec
i
es
of
t
h
epr
i
mar
ily h

o
l
arct
i
c
g
roup ANTHOMYIIDAE (root ma
gg
ot
fli
es) are p
hy
top
h
a
g
ous
in
the larval sta
g
e and, as a result, man
y
are economicall
y
important, for example, the cabba
g
e
F
I
GU

RE 9.17.
M
uscoidea. (A) The house fly, Mu
s
ca
d
ome
s
tic
a
(Muscidae); (B) the wheat bulb fly
,
H
y
lem
y
a
c
oarctata (Ant
h
omy
iid
ae); an
d
(C) a
d
ung
fl
y
,

C
or
d
i
l
ura cri
ddl
ei (Scat
h
op
h
ag
id
ae). (A,
f
rom V. B. W
i
gg
l
eswort
h,
19
5
9, Metamorphosis, pol
y
morphism, differentiation
,
S
cienti
fi

c America
n
, Februar
y
19
5
9. B
y
permission of
Mr. Eric Mose, Jr. B, C, from F. R. Cole and E. I. Schlinger, 1969
,
T
he Flies o
f
Western North Americ
a
.B
y
perm
i
ss
i
on o
f
t
h
eUn
i
vers
i

ty o
f
Ca
lif
orn
i
a Press.]
2
6
1
T
HE P
A
N
O
RP
O
ID
ORDER
S
root fl
y,
Hylemya brassicae, and the wheat bulb fl
y,
H. coa
r
ctata
(
Fi
g

ure 9.17B). Others
are sapropha
g
ous, dun
g
feeders, or are inquilines in the burrows of solitar
y
H
y
menoptera
or rodents. The small, primaril
y
holarctic famil
y
SCATHOPHAGIDAE (dun
g
flies) (50
0
spec
i
es)
i
s cons
id
ere
d
t
h
e most pr
i

m
i
t
i
ve o
f
t
h
eCa
l
yptratae. A
d
u
l
ts (F
i
gure 9.17C) ar
e
most
l
y pre
d
aceous on ot
h
er
i
nsects, t
h
oug
h

some
f
ee
d
on t
h
e
j
u
i
ce o
fd
ung. Larvae
h
ave var
i
e
dh
a
bi
ts; man
y
are
l
ea
f
an
d
stem m
i

ners, ot
h
ers pre
d
aceous, an
d
some
f
ee
d
on
d
un
g
.
Superfamily Oestroide
a
F
i
ve
f
am
ili
es are
i
nc
l
u
d
e

di
nt
hi
sver
yl
ar
g
e
g
roup o
f
D
i
ptera. CALLIPHORIDAE
(1000 species, 80% of which are restricted to the Old World) is a cosmopolitan
g
roup
t
hat includes blow flies,
g
reen- and bluebottles, and screwworm flies. Amon
g
members
o
f
t
h
e
f
am

il
y, w
hi
c
h
may
b
e parap
h
y
l
et
i
c (Rognes, 1997), a comp
l
ete spectrum o
fl
arva
l
f
ee
di
ng
h
a
bi
ts can
b
e seen, rang
i

ng
f
rom true carr
i
on
f
ee
d
ers, t
h
roug
h
spec
i
es t
h
at
f
ee
d
o
n
exu
d
ates or open woun
d
so
fli
v
i

n
g
an
i
ma
l
s, to tru
ly
paras
i
t
i
c
f
orms. Ca
lli
p
h
or
id
ae o
f
me
d
-
ical or veterinar
y
importance include the sheep blow flies
(
Lucilia

s
pp
.), the screwworm
s
[
C
ochliomyi
a
(
Callitroga) spp.] (Fi
g
ure 9.18A), and bluebottles (Calliphor
a
s
pp
.), which ar
e
vectors o
fh
uman
di
seases. C
l
ose
l
yre
l
ate
d
to t

h
eca
lli
p
h
or
id
s are t
h
e SARCOPHAGIDAE
(
fl
es
hfli
es) (F
i
gure 9.18B), w
h
ose
l
arvae
f
ee
d
on
d
ecay
i
ng an
i

ma
l
t
i
ssue or are true par-
a
s
i
tes o
f
art
h
ropo
d
s, mo
ll
us
k
s, or anne
lid
s. Most o
f
t
h
e 2000 spec
i
es
i
nt
hi

s cosmopo
li
-
t
an
g
roup are viviparous, depositin
g
first-instar larvae directl
y
into the food source. Th
e
TA
CHINIDAE (Fi
g
ure 9.18C,D), with some 8000 species worldwide, form the second
l
argest
f
am
il
yo
f
D
i
ptera. W
i
t
h
out except

i
on, t
h
e
l
arvae are paras
i
t
i
conot
h
er art
h
ropo
d
s
,
ma
i
n
l
y
i
nsects. An egg, or
i
nt
h
e many v
i
v

i
parous spec
i
es, a
l
arva,
i
s
f
requent
l
y
d
epos
i
te
d
di
rect
l
yont
h
e
b
o
d
yo
f
t
h

e
h
ost. A
l
ternat
i
ve
l
y, t
h
eegg
i
s
l
a
id
on t
h
e
h
ost’s
f
oo
d
p
l
ant. T
he
h
ost usuall

y
dies as a result of the parasitism, and there is little doubt that tachinids pla
y
a
role equal to that of man
y
parasitic H
y
menoptera in controllin
g
the population level of
certain species. Not surprisingly, some have been employed as biological control agent
s
a
gainst pests (for examples, see Table 24.6). Included in the OESTRIDAE (bot flies an
d
warble flies) (Figure 9.18E–G), a group of about 1
5
0 species, are four well-defined sub-
families that are often
g
iven famil
y
rank. OESTRINAE are holarctic and African flies that
larviposit in the nasal and phar
y
n
g
eal cavities of lar
g

e herbivores; HYPODERMATINA
E
h
ave a similar distribution, their hosts include rodents as well as herbivores, female
s
ov
i
pos
i
tont
h
e
h
ost’s s
ki
n, an
d
t
h
e
l
arvae
d
eve
l
op su
b
cutaneous
l
y; CUTEREBRINAE

a
re restr
i
cte
d
to t
h
eNewWor
ld
w
h
ere t
h
ey ov
i
pos
i
t
i
np
l
aces
f
requente
db
yt
h
e
h
ost, t

h
e
l
arvae
h
atc
hi
n
gi
n response to ra
di
ant
h
eat
f
rom an a
dj
acent
h
ost an
dd
eve
l
op
i
n
g
su
b
cuta-

neousl
y
on primates, rodents, and la
g
omorphs; and the cosmopolitan GASTEROPHILINAE
mostl
y
oviposit on the le
g
s or near the mouth of horses, zebras, and elephants, the lar-
v
ae e
v
entua
ll
yma
ki
ng t
h
e
i
r way to t
h
e
h
ost’s stomac
h
or
i
ntest

i
ne w
h
ere t
h
ey comp
l
et
e
d
eve
l
opment.
Super
f
ami
l
y Hippo
b
oscoi
d
ea
Four families are included in this group, GLOSSINIDAE, HIPPOBOSCIDAE,
STREBLIDAE, an
d
NYCTERIBIIDAE, t
h
e
l
ast t

h
ree
f
ormer
l
y
b
e
i
ng cons
id
ere
d
to con-
st
i
tute t
h
e Pup
i
par
i
a. In a
ll
o
f
t
h
e
f

am
ili
es t
h
ea
d
u
l
ts are
bl
oo
d
suc
ki
ng paras
i
tes o
fbi
r
d
so
r
mammals, and the larvae mature, one at a time, entirel
y
within the
g
enital tract of the female
,
262
CHAPTER

9
F
I
G
URE 9.18. Oestro
id
ea. (A) T
h
e screwworm
fly,
C
oc
hl
iomyia
h
ominivorax
(
Ca
lliph
or
id
ae); (B
)
S
arcop
h
aga
kell
y
i

(
Sarcopha
g
idae), a parasite of
g
rasshoppers; (C) the bean beetle tachinid,
A
plom
y
iopsis epilachna
e
(Tachinidae); (D
)
A
.e
p
ilachnae
l
arva inside bean beetle grub; (E) the sheep bot fly
,
O
estrus ovis (Oestridae)
;
(F) t
h
e catt
l
ewar
bl
e

fly,
Hypo
d
erma
b
ovi
s
(Oestr
id
ae); an
d
(G) t
h
e
h
orse
b
ot
fly
, Gasterop
h
i
l
us intestina
l
is
(Oestridae). [A, G, from M. T. James, 1948, The flies that cause m
y
iasis in Man, U.S. Dep. A
g

ric., Misc. Publ
.
#631
.
B
y permission of the U.S. Department of Agriculture. B, from L. A. Swan and C. S. Papp, 1972
,
T
he Com
-
mon Insects of Nort
h
America. Cop
y
r
igh
t 1972
by
L. A. Swan an
d
C. S. Papp. Repr
i
nte
dby
perm
i
ss
i
on o
f

Harper
&
R
ow
Publishers, Inc. C, D, b
y
permission of the U.S. Department of A
g
riculture. E, F, from A. Castellani and
A. J. Chambers, 1910
,
M
anual o
f
Tropical Medicine
.
By permission of Bailliere and Tindall.
]
bein
g
nourished b
y
secretions of the accessor
y
(“milk”)
g
lands. Pupation immediatel
y
fol-
l

ows birth and occurs off the host. Glossinidae (tsetse flies) is a very small but well-known
f
am
il
yo
f
a
b
out 20 spec
i
es o
f
Gl
ossin
a
f
rom trop
i
ca
l
A
f
r
i
ca (exc
l
u
di
ng Ma
d

agascar). T
h
e
y
are vectors o
f
var
i
ous spec
i
es o
f
T
r
y
panosom
a
th
at cause s
l
eep
i
ng s
i
c
k
ness an
d
nagana
in

h
umans an
d
catt
l
e, respect
i
ve
ly
.T
h
eH
i
ppo
b
osc
id
ae (
l
ouse
fli
es an
dk
e
d
s) (F
ig
ure 9.19A
)
resemble tsetse flies in several wa

y
s and the two probabl
y
are sister
g
roups. However, in
2
63
T
HE P
A
N
O
RP
O
ID
ORDER
S
F
IGURE 9.19. H
i
ppo
b
osco
id
ea. (A)
L
ync
h
ia american

a
(
H
i
ppo
b
osc
id
ae), a paras
i
te o
f
ow
l
san
dh
aw
k
s; (B
)
a
b
at
fly,
Stre
bl
a vesperti
l
ioni
s

(
Stre
blid
ae
)
;an
d(
C
)
Cyc
l
opo
d
ia gree
fi
(N
y
cter
ibiid
ae). [A,
f
rom F. R. Co
l
ean
d
E
. I. Schlinger, 1969
,
T
he Flies o

f
Western North America.
B
y permission of the University of California Press. B,
from Q. C. Kessel, 1925, A synopsis of the Streblidae of the world, J.
N
. Y. Entomo
l
. Soc.
33
:11–33. By perm
i
ss
i
on
of the New York Entomolo
g
ical Societ
y
. C, from H. Oldro
y
d, 19
6
4,
Th
e Natura
l
History o
f
F

l
ies
,W
eidenfeld and
W
W
Nicolson. By permission of Mrs. J. M. Oldroyd.
]
contrast to tsetse
fli
es, w
hi
c
h
are strong
fli
ers,
hi
ppo
b
osc
id
s rare
l
y
fl
y(
i
n
d

ee
d
, some s
h
e
d
th
e
i
rw
i
ngs a
f
ter sett
li
ng on a
h
ost). T
hi
s group o
f
a
b
out 330 spec
i
es
h
as a cosmopo
li
tan

di
str
ib
ut
i
on. T
h
e
y
ma
i
n
ly
paras
i
t
i
ze
bi
r
d
s
b
ut
i
nc
l
u
d
e some un

g
u
l
ates
(
Me
l
op
h
a
g
us ovinus
,
t
he sheep ked is a ma
j
or pest) and other mammals amon
g
their hosts. Most of the 1
6
0 specie
s
of Streblidae (bat flies) (Fi
g
ure 9.19B) have win
g
s, thou
g
h these are pleated to facilitate
movement through the host’s fur. Females of Ascodipteron species shed their wings an

d
l
egs an
db
urrow
i
nto t
h
e
h
ost’s s
ki
n. Most stre
blid
s are assoc
i
ate
d
w
i
t
h
co
l
on
i
a
l
spec
i

es o
f
b
ats t
h
at roost
i
n caves or
f
orests an
d
are
f
oun
di
n trop
i
ca
l
an
d
su
b
trop
i
ca
l
reg
i
ons. T

h
e
Ny
cteribiidae (2
5
0 species) (Fi
g
ure 9.19C) are win
g
less parasites of bats found mostl
y
in
w
armer re
g
ions of the world.
L
i
teratur
e
There exists a massive volume of literature on Diptera, includin
g
man
y
books, too
numerous to mention specificall
y
, on particular
g
roups of flies or aspects of their biolo

gy
.
Oldroyd (1964) and Volumes 1 and 2 of the series edited by McAlpine
et al.
(1981
–
1989)
ar
e
exce
ll
ent sources o
fi
n
f
ormat
i
on on t
h
e
bi
o
l
ogy o
f
t
h
e group. Ro
hd
en

d
or
f
(1974), aut
h
ors
i
n
V
o
l
ume3o
f
McA
l
p
i
n
e
e
ta
l
.
(
1981–1989
)
, Michelsen
(
199
6)

, Friedrich and Tautz
(
1997
)
,
and Yeates and Wie
g
mann (1999) discuss the ph
y
lo
g
en
y
and classification of the order
.
G
riffiths (1994) [Brach
y
cera], Oosterbroek and Courtne
y
(1995) [Nematocera], Na
g
atom
i