The European Families

of the Diptera
Identification, diagnosis, biology

Pjotr Oosterbroek

K N N V Publishing > -

Z ITI
 ocelli
occiput

thorax

mouthparts

closed cell open cell

habitus brachycerous fly head calyptrate fly

F = Frontorbital bristles Iv Iv Oc
ocelli frons IV = Innervertical bristles
compound eye
frontorbital Oc = Ocellar bristles
plate Ov = Outervertical bristles
P Postocellar bristles
parafaci alia
ptilinal suture
facial ridge
fronto-genal suture

head calyptrate fly; eyes in contact head calyptrate fly

eyes separated with each other
(dichoptic) (holoptic)

1st antennal segment (scapus)

2nd antennal segment (pedicel)

3rd antennal segment (Istflagellomere)

6a antenna Culicidae (Nematocera)

style 3rd antennal segment

(1st flagellomere)

flagellomere
2nd (pedicel) (scapus)

antenna Tabanidae antenna calyptrate fly head acalyptrate fly

(lower Brachycera) (higher Brachycera) 1-6 = F-bristles
 Sc-R R-M

ptilinum

arista

hypothetical ground-plan
head Heleomyzidae of the Diptera wing
with extruding ptilinum
postsutural scutum
costal break costal break
presuturai scutum (humeral break) (subcostal break)
acrostichal bristles
dorsocentral bristles
H
1 T ^ ' I/ * / /'

postpronotum
^* 1^ posthumeral bristles
y>presuturai bristle
\ * y~ notopleuron
l

transverse suture lower calypter

intra-alar bristles alula CuA2 I BM-Cu
anal vein
(A1+CuA2)

postalar callus wing Calliphoridae

(Calliphora)

trochanter

thorax Tachinidae
preapical bristle
apical bristles
postsutural
(spurs)
scutum
transverse suture wing base
n_
^ra segments
notopleuron

Diptera leg
postpronotum
subscutellum
haltre
^mediotergite
laterotergite
proepisternum
greater ampulla
proepimeron posterior spiracle
merat bristles
coxa 1 \
katepimeron
anepimeron
meron V
katepi sternum coxa 3
bristle (macrotrichium)
thorax Calliphoridae with socket (alveolus)
CONTENTS

Introduction 6

Classification 8

Terminology 12

Identification key 25

Family descriptions 110

Checklists of European countries 173

References 174

Figure details 193

Figure credits 201

Acknowledgements 202

Register (including synonymy) 203

Publishing data 205

Abstract of key inside backcover

INTRODUCTION
S o m e 25 y e a r s a g o , a k e y t o t h e E u r o p e a n
families o f the Diptera w a s published, in
D u t c h ( O o s t e r b r o e k 1981). T h e latest
checklist of the Diptera of The
N e t h e r l a n d s ( B e u k 2002) w a s r e a s o n t o
c o m p l e t e l y r e v i s e a n d u p d a t e this 1981
k e y , e x t e n d e d for e a c h f a m i l y w i t h a s h o r t
chapter o n characters, b i o l o g y and identi
fication r e f e r e n c e s ( O o s t e r b r o e k e t al.
2 0 0 5 ) . O f this b o o k , also p u b l i s h e d i n
D u t c h , an E n g l i s h v e r s i o n w a s p r e p a r e d .
Subsequently, this w a s substantially
i m p r o v e d b y the valuable corrections,
c o m m e n t s and a d d i t i o n s r e c e i v e d f r o m
o v e r 30 m a i n l y E u r o p e a n d i p t e r i s t s , l e a d
i n g t o t h e p r e s e n t k e y for and o v e r v i e w o f
the families o f the Diptera occurring in
E u r o p e , f r o m Iceland a n d F e n n o s c a n d i a i n
the n o r t h , the Ural in the east, and
Macaronesia and the Caucasus in the
s o u t h . O t h e r r e c e n t l y p u b l i s h e d k e y s are
M a t i l e 1 9 9 3 - 1 9 9 5 (in F r e n c h ) , P a p p &
Schumann 2000 (with a limited number
o f i n c l u d e d figures) and N a r t s h u k 2003
(in R u s s i a n ) .
Identification o f D i p t e r a is o f t e n c o n
s i d e r e d t o b e difficult. E x p e r i e n c e s h o w s
that in particular the smaller species can
present the student w i t h various prob
l e m s . T h i s is p a r t i a l l y d u e t o t h e q u e s t i o n
w h e t h e r t h e v e i n w h i c h f o r m s t h e front
m a r g i n o f t h e w i n g (the costa) s h o w s o n e
or t w o b r e a k s , or n o n e at all. T h i s charac
ter is u s e d i n m a n y i d e n t i f i c a t i o n k e y s
b e c a u s e o f its r e l i a b i l i t y t o d i s t i n g u i s h
b e t w e e n ( g r o u p s of) f a m i l i e s . It is n e v e r
t h e l e s s a l s o s o m e t i m e s a rather d i f f i c u l t
character. Therefore, the k e y i n c l u d e d
h e r e h a s b e e n d e s i g n e d t o e m p l o y this
character as little as p o s s i b l e . T h i s a l s o
a p p l i e s t o s o m e o t h e r characters t h a t c a n
b e difficult to a s c e r t a i n , s u c h as t h e b r i s
tles n e x t t o t h e p o s t e r i o r spiracle i n t h e
S e p s i d a e (fig. 1 9 6 , 1 9 7 ) . A n o t h e r a s p e c t o f
t h e D i p t e r a is t h e o v e r w h e l m i n g d i v e r s i t y
o f c h a r a c t e r s . In v i e w o f t h e a m b i t i o n t o
c o v e r all E u r o p e a n g e n e r a ( s o m e 2.320)
a n d s p e c i e s ( a b o u t 1 9 . 2 0 0 ) , a fair n u m b e r
of families k e y out more than once.
6
A n o b v i o u s difference w i t h r e s p e c t t o
the key published 25 y e a r s ago
( O o s t e r b r o e k 1981), is t h e n u m b e r o f f a m
ilies, w h i c h r o s e f r o m 107 t o 132. T h i s
increase is d u e to t h e fact t h a t several f a m
ilies h a v e b e e n split u p , w i t h these ' n e w '
families being internationally accepted.
S o m e examples: the former Tipulidae
have b e e n split up into 4 families;
B i b i o n i d a e i n t o 3; M y c e t o p h i l i d a e i n t o 5,
E m p i d i d a e i n t o 4 o r 5. M o r e o v e r , s u b s t a n
tial p r o g r e s s h a s b e e n m a d e i n the s t u d y o f
t h e D i p t e r a . S i n c e 1981, t h e M a n u a l o f the
N e a r c t i c D i p t e r a (J.F. M c A l p i n e et al. 1981-
1989), t h e C a t a l o g u e o f Palaearctic D i p t e r a
( S o o s & P a p p (eds) 1 9 8 4 - 1 9 9 3 ) as w e l l as
the Contributions to a Manual of
Palaearctic D i p t e r a (Papp & D a r v a s 1 9 9 7 -
2000) have been published. O u r under
s t a n d i n g o f t h e s y s t e m a t i c s also h a s b e e n
m u c h i m p r o v e d ( r e v i e w s i n Y e a t e s ck.
W i e g m a n n 1 9 9 9 , 2 0 0 5 ) , as h a s o u r
k n o w l e d g e o f d i s t r i b u t i o n a n d faunistics
(see t h e list o f c h e c k l i s t s o f t h e separate
countries).
The Nematocera (mosquitoes, midges,
g n a t s , etc.) a n d B r a c h y c e r a (true flies)
together constitute the insect order
Diptera, a w o r d that derives from the
Greek di = t w o and pteron = w i n g . A c o m
m o n character o f all D i p t e r a is t h e r e d u c
t i o n o f t h e s e c o n d set o f w i n g s (the p o s t e
rior pair) t o a pair o f h a l t e r e s or w i n g
knobs. This reduction has not impaired
t h e f l y i n g c a p a c i t y o f t h e s e i n s e c t s . In fact,
m a n y D i p t e r a are o n a par w i t h o t h e r g o o d
fliers l i k e d r a g o n f l i e s , s o m e b e e s and
w a s p s w i t h r e s p e c t to their m a n o e u v r a
b i l i t y and s p e e d . T h e transfer o f t h e f l y i n g
a c t i o n t o j u s t t h e anterior pair o f w i n g s
h a s led t o a s t r o n g d e v e l o p m e n t o f the
middle part of the thorax, n a m e d the
m e s o t h o r a x . T h i s character affords i d e n t i
fication o f t h e s e v e r a l D i p t e r o u s s p e c i e s
t h a t h a v e l o s t their w i n g s i n t h e c o u r s e o f
e v o l u t i o n , in o n e sex or b o t h . T h e s e
flightless s p e c i e s i n c l u d e parasites t h a t
h a v e their b o d y a d o p t e d in s u c h a w a y
t h a t t h e y w o u l d n o t o t h e r w i s e b e easily
r e c o g n i s e d as b e l o n g i n g to t h e D i p t e r a .
T h e D i p t e r a d i s p l a y a h u g e array o f
s h a p e s and s h o w a great v a r i e t y i n their
biology. T h e n u m b e r of species and that o f
i n d i v i d u a l s is o v e r w h e l m i n g . A t n e a r l y all
l o c a t i o n s , t h e D i p t e r a are t h e m o s t
n u m e r o u s i n s e c t s . For e x a m p l e , t h e n u m
ber o f D i p t e r a s p e c i e s k n o w n f r o m t h e
N e t h e r l a n d s (4,967 after B e u k 2002) is
more than t w i c e that o f the Lepidoptera
(2,336, K u c h l e i n & D e V o s 1 9 9 9 ) . It is also
larger t h a n t h a t o f t h e C o l e p t e r a ( 4 , 0 4 1 ,
V o r s t & H u i j b r e g t s 2002). T h e s e figures,
however, do n o t m e a n that in general the
D i p t e r o u s fauna is w e l l k n o w n . S e v e r a l
families h a v e b e e n relatively w e l l s t u d i e d
b u t there is a c o n s i d e r a b l e terra i n c o g n i t a
c o m p r i s i n g , for e x a m p l e , f a m i l i e s r i c h i n
species like the Limoniidae, C e r a t o p o -
gonidae, Mycetophilidae, Phoridae,
Lauxaniidae, A g r o m y z i d a e , Ephydridae,
Drosophilidae, Chloropidae, A n t h o m y i -
idae and m a n y o f the s m a l l e r f a m i l i e s as
well.
T h e study o f the Diptera does n o t have
to b e l i m i t e d to i n v e n t o r i e s o f t h e fauna.
In the field o f b i o l o g y m u c h w o r k is still t o
be d o n e . T h e available i n f o r m a t i o n o f t e n
derives from a small n u m b e r of observa
tions i n o n l y a f e w s p e c i e s . In particular,
the v a s t d i v e r s i t y w i t h regard t o e c o l o g y
and behaviour m a k e the Diptera an inter
e s t i n g t o p i c for r e s e a r c h suitable for t h e
amateur. For e x a m p l e , rearing p h y
t o p h a g o u s Diptera can in m a n y cases be
done w i t h o u t specialist training or means.
O n e n e e d n o t b e an e x p e r t t o d i s c o v e r
n e w facts.
Diptera belong to the h o l o m e t a b o l o u s
i n s e c t s , i.e., e a c h i n d i v i d u a l l e a d s t w o
' s e p a r a t e ' l i v e s : first as a l a r v a , t h e n as an
a d u l t . A s a r u l e , t h e l a r v a l s t a g e s are c o n
s i d e r e d n e c e s s a r y t o b u i l d u p a n d store
reserves to enable the adult stage to exist.
T h i s final s t a g e , t h e a d u l t d i p t e r o n , is
m a i n l y d i r e c t e d at p r o p a g a t i o n . It is
w o r t h w h i l e t o s t u d y t h e l a r v a e , as e v e n
t o d a y v e r y little or e v e n n o t h i n g at all is
k n o w n a b o u t t h i s stage o f life i n a n u m b e r
o f Dipterous families. Readers interested
i n t h i s s u b j e c t w i l l find m o r e i n f o r m a t i o n
i n Ferrar 1987; M a t i l e 1 9 9 5 ; S m i t h 1 9 8 9 ;
S t u b b s 8t C h a n d l e r 1 9 7 8 ; C o u r t n e y et al.
2000.
C o l l e c t i n g a n d p r e s e r v i n g i n a g r o u p as
d i v e r s e as t h e D i p t e r a r e q u i r e s v a r i o u s
t e c h n i q u e s . It is b e y o n d t h e s c o p e o f t h e
p r e s e n t b o o k t o deal w i t h t h i s . R e f e r e n c e s
o n t h i s s u b j e c t are G u l l a n 8t C r a n s t o n
2 0 0 5 ; M a t i l e 1993; S c h a u f f 2 0 0 2 ; S m i t h e r s
1982; S t u b b s 8t C h a n d l e r 1 9 7 8 ; U p t o n
1991.
7
CLASSIFICATION
T h e order Diptera includes t w o major
groups, viz., the Nematocera (mosqui
t o e s , m i d g e s , g n a t s , etc.) a n d the
B r a c h y c e r a (true flies). B o t h g r o u p s are
d i v i d e d i n t o a n u m b e r o f s u b g r o u p s (see
fig. i a n d C l a s s i f i c a t i o n t a b l e b e l o w ) .
T h e r e is s o m e g e n e r a l a g r e e m e n t o n t h e
divisions m e n t i o n e d here. T h e relation
s h i p s b e t w e e n t h e v a r i o u s g r o u p s (fig. 1),
h o w e v e r , are still a m a t t e r o f d e b a t e . T h i s
also applies to the question o f w h i c h
g r o u p s are to b e a c c o r d e d f a m i l y s t a t u s .
In t h e table b e l o w , g r o u p i n g s o f f a m i l i e s
for w h i c h n o c u r r e n t n a m e is available are
i n d i c a t e d as L o w e r M u s c o m o r p h a , L o w e r
Eremoneura, Lower Cyclorrhapha. The
t e r m s H i g h e r a n d L o w e r are u s e d i n t h e
literature o n s y s t e m a t i c s t o i n d i c a t e t h e
position o f a group w i t h i n the overall
o r d e r o f all g r o u p s .
T h e name Nematocera means 'thread
l i k e a n t e n n a e ' a n d refers t o t h e a n t e n n a e
i n this g r o u p t h a t are o f t e n l o n g a n d u s u
ally comprise many segments.
Conversely, the Brachycera have short
antennae, m o s t often consisting o f three
s e g m e n t s p l u s an arista. T h e m a x i m u m
n u m b e r o f s e g m e n t s i n t h e B r a c h y c e r a is
10, w i t h t h e e x c e p t i o n o f t h e R a c h i c e r i d a e
i n w h i c h this n u m b e r r e a c h e s a b o u t 4 0
due to secondary division o f the seg
ments.
T h e E u r o p e a n N e m a t o c e r a are split u p
into s e v e n infraorders and the European
B r a c h y c e r a i n t o f o u r infraorders. O n e o f
t h e latter, t h e infraorder M u s c o m o r p h a ,
- Ptychopteromorpha
-Culicomorpha
- Blephariceromorpha
- Bibionomorpha
- Psychodomorpha
-Tipulomorpha
Tabanomorpha
Xylophagomorpha
Stratiomyomorpha
Brachycera' Nemestrinoidea
Asiloidea
Muscomorpha' Empidoidea
Lower Cyclorrhapha
Eremoneura
Lg^Acalyptratae
Cyclorrhapha^ ^ { Z C a l t r a t a e
Schizophora
i n c l u d e s t h e v a s t m a j o r i t y o f f a m i l i e s and
the large g r o u p Cyclorrhapha. In
C y c l o r r h a p h a t h e l a r v a e p u p a t e inside a
usually barrel-shaped puparium made up
o f t h e s k i n o f t h e last larval instar. A d u l t
C y c l o r r h a p h o u s flies l e a v e this p u p a r i u m
t h r o u g h a m o r e o r less circular orifice. In
case o f all B r a c h y c e r a t h a t h a v e b e e n p o s i
t i o n e d earlier i n the s y s t e m a t i c order, t h e
a d u l t s e m e r g e t h r o u g h a l o n g i t u d i n a l or a
T - s h a p e d slit, g i v i n g rise to t h e n a m e o f
O r t h o r r a p h a t h a t is o f t e n u s e d to d e n o t e
these Diptera.
In t h e C y c l o r r h a p h a , t h e largest g r o u p
are t h e S c h i z o p h o r a , c o m p r i s i n g all
A c a l y p t r a t e and C a l y p t r a t e fly f a m i l i e s , all
of w h i c h possess a ptilinum (see
T e r m i n o l o g y ) . In t h e C a l y p t r a t a e t h e
l o w e r c a l y p t e r s are g e n e r a l l y w e l l d e v e l
o p e d , i n t h e A c a l y p t r a t e flies this struc
t u r e is s m a l l (see the identification key,
n o . 82). T h e latter g r o u p i n c l u d e s m a n y
f a m i l i e s . In t h e C a l y p t r a t e f a m i l i e s
H i p p o b o s c i d a e , N y c t e r i b i i d a e and S t r e b -
lidae, all parasites o f m a m m a l s and birds,
t h e l a r v a e d e v e l o p f u l l y inside the f e m a l e
w o m b a n d p u p a t e i m m e d i a t e l y after d e p
o s i t i o n . T h e s e f a m i l i e s are s o m e t i m e s
grouped under the name Pupipara.
A n o t h e r g r o u p w i t h i n t h e C a l y p t r a t e flies
are t h e b o t s a n d w a r b l e f l i e s , i n c l u d i n g the
three families o f Gasterophilidae, H y p o -
dermatidae and Oestridae. Their larval
s t a g e s l i v e as parasites i n m a m m a l s . T h e
a d u l t flies s h o w l a r g e l y r e d u c e d m o u t h -
parts a n d are u n a b l e t o feed.
Nematocera
(midges, gnats
mosquitoes, etc.)
Brachycera Outline of the relation
(true flies)
ships between the higher
categories of the
Diptera; after Yeates &
Wiegmann 1999
CLASSIFICATION TABLE

N a m e s o f h i g h e r c a t e g o r i e s h a v e specific e n d i n g s , i n d i c a t i n g t h e i r r a n k ; s o m e o f t h e e n d -
i n g s u s e d i n z o o l o g y are: I n f r a o r d e r : - m o r p h a ; S u p e r f a m i l y : - o i d e a ; F a m i l y : - i d a e ;
S u b f a m i l y : -inae; T r i b e : -ini.
B e h i n d the f a m i l y n a m e t h e n u m b e r o f g e n e r a a n d s p e c i e s in E u r o p e is g i v e n .

SUBORDER
INFRAORDER SUPERFAMILY FAMILY(i32)

N e m a t o c e r a ( M i d g e s and the like) 32 families

Tipulomorpha Tipuloidea TJgulidae: 1 0 , 4 7 0
L i m o n i i d a e : jo, 5 6 0
Pediciidae: 5 , 6 0
Cylindrotomidae: 4 , 6
Trichoceroidea Trichoceridae:j2 j;o !

Blephariceromorpha Blephariceroidea Blephariceridae^5_,^8

Axymyiomorpha jf^xymyioidea A x y m y i i d a e : 1,1
Bibionomorpha Pachyneuroidea P a c h y n e u r i d a e : 1,1
Bibionoidea Bibionidae: 2,47
Pleciidae: 1,2
H e s p e r i n i d a e : 1,1
Sciaroidea Mycew^tdlidae: 70, 945
B o l i t o p h i l i d a e : i,J(6^
D i a d o c i d i i d a e : 1, 5
D i t o m y i i d a e : 2, 2
Keroplatidae: 1 6 , 1 1 0
Sciaridae: 28, 620
Cecidomyiidae: 280,1640
Psychodomorpha Psychodoidea Pjychodidaejj.o, 500
Anisopodoidea Anisopodidae: 1,10
Mycetobiidae: 2 , 4
Scatopsoidea Scatopsidae: 22,100
S j m n e u r i d a e : 1,1
Carrthyjoscelidae: 1, 2
Ptychopteromorpha Ptychopteroidea Ptydbogteridae: 1,15
Culicomorpha Culicoidea Dixidae: 2,32
Chaoboridae: 3 ^
C u l i c i d a e : 8, i o 5 _
Chironomoidea T h a u m a l e i d a e : 2r>2a
Simuliidae: 8,230
Ceratogogonidae: 0,530^
Chironomidae: 180,1130

Brachycera (True Flies) loojamilies

Xylophagomorpha Xyloghagoidea _ X y J o p h a g i d a e : 1, 5
C o e n o m y i i d a e : 1,1
R a c h i c e r i d a e : 1,1
Stratiomyomorpha Stratiomyoidea Xjyjomyidae: 2 , 8
Sttatiomyidaejji/-, 1 4 Q _ _

9
 Tabanomorpha Tabanoidea Athericidae: 4 , 1 0
R h a g i o n i d a e : 6, 85^
T a b a n i d a e : 13,220
V e r m i l e o n i d a e : 2, 9
Muscomorpha
Lower Muscomorpha Nemestrinoidea Nemestrinidae: 6 i 3 _ _ t

A c r o c e r i d a e : 8, j
Asiloidea B o m b y l i i d a e : 52, 3 4 0
M y t h i c o m y i i d a e : j , 30
Hilarimorphidae: l, 2
Therevidae: ljVioo
Scenopinidae: 2,16
Mydidae: 4, j_
Asilidae^77,540
Eremoneura
Lower Eremoneura Empidoidea A t e l e s t i d a e : 3,JL
Hybotidae^jo, 4 4 0
Empididae: 23,810
B r a c h y s t o m a t i d a e : 5,ji3_
Microphoridae: 4 , 1 6
D o l i c h o p o d i d a e : bo/ZJIL^.
(Cyclorrhapha)
Lower Cyclorrhapha Platypezoidea Platypezidae: 12,45
O p e t i i d a e : 1,1

Lonchopteroidea
P horidae^s^Goj
L o n c h o p t e r i d a e : 2^13
Syrpjioidea Syrphidae: 90,830
P i p u n c u l i d a e n ^ 20 0
Schizophora
Acalyptratae Nerioidea Micropezidae: 5,22
P s e u d o p o m y z i d a e : 1,1
Diopsoidea T a n y p e z i d a e : 1,1
S t r o n g y l o p h t h a l m y i i d a e : 1,2
Psilidae: 6^30
M e g a m e r i n i d a e : 1,1
D i o p s i d a e : 1,1
Conopoidea Conopidae: 14,85
Tepjiritoidea Lonchaeidae: 8,100
Pallopteridae^jj, 23^
P i o p h i l i d a e : 1 6 , 30
U l i d i i d a e : IJ, i o _
Platystomatidae: 2,20
_jnej&ritidae220jj220
P y r g o t i d a e : 1,1
Lauxanioidea Lauxaniidae: 1 8 , 1 6 0
Chamaemyiidae^, n o
C r e m i f a n i i d a e : 1,2
Sciomyzoidea Coelopidae: 2,3
Dryomyzidae: 2,4
H e l c o m y z i d a e : 1, 2
H e t e r o c h e i l i d a e : 1,1
Sciomyzidae: 2 4 , 1 4 0

1 o
 Phaeomyjidae.^i,_ 3
Sepsidae: 3^50
OjDomyzoidea Clusiidae: 5 ^ 5
Acartophthalmidae: i,jj
Odiniidae: 3 , 1 4
Ajgromyzidaejjyugio
O p o r n y z i d a e : 3^33
Anthomyzidaerjj^o
Aulacigastridae: 1,4
S t e n o m i c r i d a e : 1,3
Periscelididae: 1 , 4
Xenasteiidae: 2,2 _
Asteiidae: 4 , 1 8
Carnoidea Milichiidae: 9 , 4 5
Carnidae: 3 , 4 0
C r y p t o c h e t i d a e : 1,3
B r a u l i d a e : 1,3
T e t h i n i d a e : 3, 35
Canacidae: V 4
Chloropidae: 6 5 , 3 3 5 ^
Sphaeroceroidea Heleomyzidae: 23,150
T r i x o s c e l i d i d a e : 1, 25
C h i r q p t e r o m y z i d a e : 2, 2
C n e m q s g a t h i d a e : 1,1
Chyrornyidae:^, 60
S g h a e r o c e r i d a e : 4 0 , 260
Ephydroidea Drosophilidae: 17,120
Campichoetidae: 2,7
D i a s t a t i d a e : 1, 9
C u r t o n o t i d a e : 1,1
C a m i l l i d a e : 1,8
E p h y d r i d a e : 6 0 , 335
Sjmerfamily u n k n o w n N a n n o d a s t i i d a e : 1,3
Calyptratae Hippoboscoidea H i p p o b o s c i d a e : n,jJo^
Nycteribiidae: 4 ^ 1 3
Streblidae: 1,1
Muscoidea Scathophagidae: 38,160
Anthomyiidae: 35,480
F a n n i i d a e : 3, 82
Muscidae: 4 5 , 5 7 5
Oestroidea Calliphoridae: 22,115
Rhinophoridae: 11,45
S a r c o p h a g i d a e : 30,310^
Tachinidae: 280,880
Oestridae: 4 , 8
Gasteroghilidae: 1,6
Hypodermatidae: 4 , 8

( H i g h e r Classification a c c o r d i n g to Y e a t e s 8t W i e g m a n n 1999)
TERMINOLOGY

People can be o v e r w h e l m e d b y the multitude o f terms used to describe the characters neces
sary for insect identification. T h e terms often vary b e t w e e n groups and are often used only b y
specialists w o r k i n g o n a given group. In this b o o k o n l y those terms are used that have been
internationally accepted b y Diptera workers. T h e y are mainly concerned w i t h the head and its
various bristles, the thorax, w i n g s , w i n g venation and legs. Terms that occur frequently and
those used in the text are enumerated here b y order o f b o d y parts and in alphabetical order.
For a more extensive explanation and a fuller s u r v e y o f terms, the reader is referred to Ziegler
2003 and the introductory chapters o f the Manuals o f Nearctic and Palaearctic Diptera,
respectively b y J.F. M c A l p i n e et al. (1981-1989) and Papp 8t Darvas (1997-2000).

GENERAL TERMS
A n t h r o p o g e n o u s or Synanthropic. Mainly occurring in habitats created b y man.
A p t e r o u s . W i t h o u t w i n g s (fig. 4 4 0 ) .
Brachypterous. H a v i n g short w i n g s (fig. 4 4 8 ) .
Brackish. H a v i n g a salinity b e t w e e n that of fresh w a t e r and seawater.
Bristle. See Macrotrichium (fig. 16).
Carnivorous. Feeding o n meat (see also Predator).
Cerci (singular: Cercus). U s e d in this b o o k to refer to the dorsal valves o f the ovipositor in
s o m e Nematocera (fig. 31). See Sexual d i m o r p h i s m .
C o m m e n s a l i s m (Commensal). T h e relation b e t w e e n t w o kind of organisms (commensals) in
w h i c h one obtains food, protection or other benefits from the other w i t h o u t damaging or
benefiting it.
C o p r o p h a g o u s . Feeding o n excrement.
C o s m o p o l i t a n . Distributed w o r l d - w i d e rather than regional, usually c o m m o n in m o s t parts
of the w o r l d and under varied ecological conditions. See also Ubiquitous.
Facultative. Optional; able to survive a n d / o r propagate itself under a variety o f circum
stances.
Female. See Sexual d i m o r p h i s m .
Filiform. H a v i n g the shape of a thread or filament.
Forensic e n t o m o l o g y . A branch o f applied e n t o m o l o g y largely dedicated to crime-solving and
coroner's inquests. U s e d , for example, to determine the date o f death of a deceased person.
Genitalia, male ( h y p o p y g i u m ) . See Sexual d i m o r p h i s m .
Habitat. Ecological range; the preferred e n v i r o n m e n t of a plant or animal species (e.g. marsh,
forest, alpine m e a d o w s , etc.).
Hairs. See Microtrichium.
Halophilous (Halobiont). Flourishing in a saline or salty habitat; such an organism is called a
halobiont.
Hemispherical. R e s e m b l i n g a hemisphere, the half of a sphere.
H y p o p y g i u m . Male genitalia. See Sexual d i m o r p h i s m .
Infuscated. Darkened w i t h a tinge, e.g. the w i n g m e m b r a n e can be entirely darkened b y a
b r o w n i s h tinge.
Inquiline. A n animal that lives in the nest or abode of s o m e other species.
Joint. See segment.
Larviparous. T h e eggs hatch inside the female w h i c h deposits larvae instead of eggs.
Macrotrichium. T h i s is a general t e r m for a bristle or seta. This can be recognised b y the rein
forcement ring at the base, n a m e d the alveolus (fig. 16). O v e r s i z e d macrotrichia are named
spurs or spines, e. g., the apical spurs or spines o n the tibiae (fig. 75). N o r m a l macrotrichia
are called bristles or setae, small ones are called setulae.
Male. See Sexual d i m o r p h i s m .
Microtrichium. T h i s t e r m is reserved for elements w i t h o u t an alveolus (fig. 16). Usually they
 are hair-like; hence, they are often called hairs w h i c h is, strictly speaking, erroneous since
real hairs occur only in m a m m a l s .
Mining. Living in, and eating its w a y through, plant tissue.
Mycophagous. Feeding o n m u s h r o o m s , (bracket) fungi, or m y c e l i u m .
Myiasis. Infection or disease caused b y Diptera larvae.
Necrophagous. Feeding o n dead animals.
Obligatory. Restricted to a specific w a y of life or m o d e of propagation.
Oviscape. N o n retractable basal segment of the ovipositor. See also Sexual d i m o r p h i s m .
Oviparous. Laying eggs.
Ovipositor. See Sexual d i m o r p h i s m .
Parasite. Feeding in (endoparasite) or o n (ectoparasite) another, usually m u c h larger animal,
in an intimate (subtle) manner, for the entire or a relatively l o n g portion o f the parasite's life
cycle, causing some damage b u t c o n s u m i n g a small proportion of the host, n o t killing it.
Parasitoid. Like a parasite feeding in or o n the h o s t in an intimate manner for a relatively l o n g
period, b u t unlike a parasite and like a predator in predictably killing the h o s t and c o n s u m
ing m o s t or all of its tissues. In some cases, several hosts are required for the d e v e l o p m e n t
of a single parasitoid, e.g. in some Sciomyzidae.
Parthenogenetic. Capable o f reproduction that involves d e v e l o p m e n t o f a female (or rarely a
male) gamete w i t h o u t fertilisation.
Phytophagous. Feeding o n plant materials.
Pilose. C o v e r e d w i t h hair, especially o f soft texture.
Pilosity. T h e state of being pilose.
Plumose. Provided w i t h rays of long hairs or hair-like bristles.
Predator (Predatory, Predacious). A t t a c k i n g prey and feeding u p o n living animal tissue.
Pruinose. C o v e r e d w i t h w h i t i s h dust or b l o o m .
Pruinosity. T h e state of being pruinose.
Pubescent. C o v e r e d w i t h fine soft short hairs.
Pupiparous. T h e larvae develop inside the female and pupate i m m e d i a t e l y after h a v i n g been
deposited.
Raptorial. Living on prey; adapted to seize prey.
Reniform. Resembling a kidney in shape.
Saline. Salty to a degree just b e l o w or above that o f seawater.
Saprophagous. Feeding on decaying organic matter.
Segment. In all Diptera, m a n y b o d y parts consist o f a series o f externally sclerotized m e m
bers, the segments. T h e y are separated b y joints. Segments are n u m b e r e d g o i n g from the
b o d y outward, starting from the head.
Seta(e). See Macrotrichium (fig. 16).
Setula(e). See Macrotrichium.
Sexual d i m o r p h i s m . A l t h o u g h the differences b e t w e e n male and female are generally limit
ed, there are m a n y cases in the Diptera w h e r e there are secondary sexual characters and
some w h e r e the sexes look quite different (e.g. Platypezidae, A n t h o m y i i d a e ) . Usually
males are smaller, s h o w better d e v e l o p m e n t of the antennae and a m o r e conspicuous colour
pattern. In several families the males have larger eyes w h i c h m e e t each other at a p o i n t or
along a line (holoptic). Differences in w i n g shape or venation sometimes occur.
T h e male can be recognised by the presence of the genitalia or h y p o p y g i u m . This consists
of a set of intricate structures the male needs to h o l d the posterior end of the female
abdomen during copulation. T h e h y p o p y g i u m can be largely visible, w h i c h is the case in
many Nematocera and Brachycera, b u t quite often it is s w u n g d o w n and protected b y a cov
ering plate (e.g. the epandrium) or it can be sunk into the abdomen proper to a large extent.
T h e female generally has an ovipositor. In the 'primitive' families this is n o t really a tube,
but a set o f paired dorsal and ventral appendages (cerci and h y p o g y n i a l valvae, together
named ovipositor). In some of the higher Diptera a tube has b e e n formed through fusion o f
several elements. In some instances, e.g. in the Tephritoidea, part of this tube, the oviscape,
is always visible as an elongate structure (fig. 331,350,351,353,357).

13
Spine. See Macrotrichium.
Spur. See Macrotrichium.
Synanthropic (Anthropogenous). Mainly occurring in habitats created b y m a n .
Ubiquitous. O c c u r r i n g e v e r y w h e r e , or in m a n y places in different ecosystems, but not nec
essarily throughout the w o r l d (= C o s m o p o l i t a n ) .

ORIENTATION AND SHAPE

N o t e that combinations o f orientation occur: Posteroventral means posterior and ventral at
the same time. T h e s e combinations are often necessary to describe a location more exactly.
A c u t e . Ending in, or w i t h a sharp angle.
Anterior. A t the head end or directed t o w a r d the head end.
Anterodorsal. Position intermediate b e t w e e n anterior and dorsal.
Anteroventral. Position intermediate b e t w e e n anterior and ventral.
A p i c a d (Distad). T o w a r d the apex. O p p o s i t e of basad.
Apical (Distal). A t or near the top; in the direction of the top, the apex (fig. 2a).
Basad (Proximad). T o w a r d the base. O p p o s i t e o f apicad.
Basal (Proximal). Near the base, the beginning; in the direction o f the base (fig. 2a).
Caudad. Backward, t o w a r d the rear. O p p o s i t e o f rostrad.
Caudal. A t or near the rear (fig. 2a).
Cephalic. Forward or near, at or o n the head (fig. 2a).
Concave. H o l l o w , c u r v i n g inward.
C o n t i n u o u s . N o t interrupted (costa).
C o n v e r g i n g bristles. Bristles that are directed t o w a r d each other, m e e t or cross each other; e.g.
fig. 200: P-bristles.
C o n v e x . Bulging, c u r v i n g o u t w a r d .
Distad. A n o t h e r t e r m for apicad.
Distal. A n o t h e r t e r m for apical (fig. 2a).
D i v e r g i n g brisdes. Bristles that are directed a w a y from each other; e.g. fig. 9: P-bristles.
Dorsal. O n the upper side; v i e w e d from above.
Dorsoventral. D o w n from the top.
Dorsoventrally compressed or flattened. C o n s p i c u o u s l y w i d e r than high. O p p o s i t e o f later
ally compressed.
Elongate. O b l o n g , (much) longer than w i d e .
Equidistant. A t the same distance.
Exclinate bristles. Bristles c u r v i n g o u t w a r d ; e.g. fig. 8: F-bristles.
Geniculate. W i t h a kink or sharp curve; a geniculate element consists of t w o (or more, if it is
t w i c e geniculate) parts w i t h different orientation.
Inclinate bristles. Bristles c u r v i n g inward, mediad; e.g. fig. 9: bristles 4 - 6 .
Interrupted. W i t h o n e or t w o breaks (costa).
Laterad. From the centre line t o w a r d the side. O p p o s i t e o f mediad.
Lateral. O f or o n the side, v i e w e d from the side.
Laterally compressed. C o n s p i c u o u s l y higher than w i d e . O p p o s i t e o f dorsoventrally c o m
pressed.
Lateroclinate bristles. Bristles directed o u t w a r d ; e.g. fig. 8: F-bristles.
Mediad. From the side(s) t o w a r d the middle; t o w a r d the plane o f s y m m e t r y . O p p o s i t e of lat
erad.
Medial. In the middle, in the plane o f s y m m e t r y (fig. 2a).
Parallel. W i t h the same orientation, equidistant at all points.
Posterior. A t the h i n d end or directed t o w a r d the hind end.
Posterodorsal. Position intermediate b e t w e e n posterior and dorsal.
Posteroventral. Position intermediate b e t w e e n posterior and ventral.
Preapical. Near, but not at the top or apex. A n example is the dorsal preapical bristle of the tibia
(fig. 15). T h i s is inserted o n the dorsal side o f the tibia, near but distincdy before the end
(apex).

14
Proclinate bristles. Bristles c u r v i n g forward, rostrad; e.g. fig. 245: l o w e r m o s t F-bristles.
Proximad. A n o t h e r t e r m for basad.
Proximal. A n o t h e r t e r m for basal (fig. 2a).
Pulvilliform. O f the same shape as a p u l v i l l u m (fig. 103). See Leg: Tarsus.
Reclinate bristles. Bristles c u r v i n g backward, posteriorly; e.g. fig. 200: F-bristles.
Rostrad. T o w a r d the front, forward. O p p o s i t e o f caudad.
Terminal. A t the end, at the top.
Ventral. O n , of, t o w a r d the l o w e r side; v i e w e d from b e l o w .

anterior

2a Tabanidae (Tabanus) 2b Brachystomatidae (Brachystoma)

2 Terms used in pointing out the orientation

HEAD
Antenna (fig. 3-7). In n a m i n g the various parts o f the antenna (fig 6 a-c), a distinction is made
:

b e t w e e n the t w o basal segments (first = scape, second = pedicel) and all other elements
together (= the flagellum). T h e elements o f the flagellum are n a m e d flagellomeres and
therefore the third antennal s e g m e n t is also the first flagellomere.
In the Nematocera the flagellum consists o f several (in general about 14) usually simple
and uniform segments (flagellomeres). T h e y often s h o w a basal ring o f l o n g hairs, n a m e d
the verticils (fig. 6a). In the l o w e r Brachycera the antennal segments can be u n i f o r m as w e l l ,
similar to w h a t is found in the Nematocera. Usually, h o w e v e r , the flagellomeres (some 8 or
less) are not that similar. T h e first flagellomere is often the largest, the consecutive ones are
conical and together they m a k e u p a style (fig. 6b) or, as in the higher Brachycera, they c o n
stitute the arista w h i c h can be similar to a bristle and w h i c h appears as an elongation of the
first flagellomere. Separate segments of the arista are n a m e d aristomeres (fig. 6c). T h e arista
can be bare (fig. 6c), pubescent (fig. 10), or p l u m o s e (fig. 7, 8).
Arista. See A n t e n n a .
Aristomere. See A n t e n n a .
Bristles of the head (fig. 8,9). T h e place of insertion and the orientation of the bristles on the
head are of crucial importance to the identification o f acalyptrate flies. T h e vertical bristles
are found on the upper margin of the head, the vertex. T h e s e bristles are divided into the P-
bristles (post-ocellar bristles) immediately b e h i n d the ocellar triangle (fig. 8, 9: P); the Iv-
bristles (innervertical bristles, fig. 8, 9: Iv) and Ov-bristles (outervertical bristles, fig. 8, 9:
O v ) . In b e t w e e n the eyes, usually along the eye margin, are the l o w e r frontorbital bristles
(fig. 9, bristles 4-6). More dorsally, the upper frontorbital bristles are located (fig. 9, bristles
1-3). In this b o o k , all frontorbital bristles are included under the t e r m F-bristles (fig. 8: F).

1 5
 occiput

ocell

compound eye

thorax

mouthparts

closed cell open cell

3 habitus brachycerous fly 7 head calyptrate fly

F = Frontorbital bristles
ocelli frons IV = Innervertical bristles
compound eye Oc = Ocellar bristles
frontorbital
Ov = Outervertical bristles
plate
P = Postocellar bristles
parafacialia
ptilinal suture
facial ridge
fronto-genal suture

fronto-clypeal
suture

4 head calyptrate fly; 5 eyes in contact 8 head calyptrate fly

eyes separated with each other
(dichoptic) (holoptic)

1st antennal segment (scapus)

2nd antennal segment (pedicel)

3rd antennal segment (1st flagellomere)

6a antenna Culicidae (Nematocera)

style 3rd antennal segment

(1st flagellomere)

flagellomere
2nd (pedicel) (scapus)

6b antenna Tabanidae 6c antenna calyptrate fly 9 head acalyptrate fly

(lower Brachycera) (higher Brachycera) 1-6 = F-bristles

16
 T h e Oc-bristles (ocellar bristles) are in the dorsomedial region o f the head, usually inside
the ocellar triangle, in rare cases just outside it. T h e area b e t w e e n the ocellar triangle, the
plates o n w h i c h the F-bristles are inserted and the ptilinal suture or bases o f the antennae
is n a m e d the frontal vitta or interfrontal area w h i c h may, or m a y not, s h o w characteristic
interfrontal bristles or setulae (fig. 8). B e l o w the eyes, w h e r e the genae m e e t the l o w e r mar
gin of the face, the vibrissae are inserted.
Apart from the presence o f bristles their orientation is an important character. T h e bris-
des can be c u r v i n g a w a y from each other (diverging), be parallel (equidistant), c u r v i n g
toward or e v e n touching or crossing each other (converging). For the F-brisdes it is i m p o r
tant to k n o w if they are c u r v i n g o u t w a r d (latero- or exclinate), b a c k w a r d (reclinate),
inward (inclinate), or forward (proclinate).
C l y p e u s . A distinct part o f the head b e t w e e n the fronto-clypeal suture and the mouthparts
(fig. 4). Usually the clypeus is small and inconspicuous, b u t it can be large and o b v i o u s (94,
n r
95> 3 > 3!4)' f example in families s h o w i n g w e l l developed piercing or sucking m o u t h -
parts.
Dichoptic. See Eye.
Eye bridge. See Eye.
Eye. A n insect eye is a so-called c o m p o u n d eye, made u p o f a multitude o f small lenses, the
facets or ommatidia. In the Diptera, the eyes usually o c c u p y a major part of the head. T h e y
are usually separate (dichoptic, fig. 3,4) b u t sometimes m e e t each other at a point or along
a line (holoptic, fig. 5). In m o s t cases the holoptic condition is limited to the males, b u t in
some families both sexes can be holoptic. W h o l e series o f facets can be larger in size than
those elsewhere, this difference in size can be so extensive a n d / o r p r o m i n e n t that it is p o s
sible to discriminate b e t w e e n t w o differently faceted parts o f the eye (e.g. an upper and a
lower part, as in fig. 55). In some Nematocera the eyes are joined through a n a r r o w bridge,
named the eye bridge, above the antennae. It consists o f several r o w s o f facets (fig. 58,73).
Many Diptera possess hairs in b e t w e e n the facets, n a m e d ommatrichia, w h i c h can be so
long or n u m e r o u s as to produce a v e l v e t y aspect.
F-bristles (Frontorbital bristles) (fig. 8: F; fig. 9: brisdes 1-6). See Brisdes of the head.
Face (Prefrons) (fig. 4). T h e plate dorsally delimited b y the antennal sockets, ventrally b y the
fronto -clypeal suture and laterally b y the fronto -genal suture or, if it is absent, b y the c o m
p o u n d eyes. T h e face m a y be small in b l o o d sucking (Culicidae, C e r a t o p o g o n i d a e ,
Tabanidae) or raptorial Diptera (Empididae, Hybotidae, fig. 96), b u t is w e l l - d e v e l o p e d in
m o s t Diptera, often w i t h grooves in w h i c h the antennae are situated, separated b y a m e d i
an ridge, the facial carina. O t h e r grooves that can be present are the fronto-genal sutures,
separating the medial part o f the face from the facial ridge, and the ptilinal suture, separat
ing the parafacialia from the remainder of the face (fig. 4 , 9 1 ) .
Facets. See Eye.
Facial carina. See Face.
Flagellomere. See A n t e n n a .
Flagellum. See A n t e n n a .
Frons (fig. 4 , 7 , 8 ) . Part of the head bordered b y the eyes, the vertex and the lunula or antennal
bases. T h e frons encompasses the frontorbital plates, the ocellar triangle and the inter
frontal area in b e t w e e n .
Frontal. Pertaining to the frons. N o t to be confused w i t h anterior or rostral.
Frontal vitta. Interfrontal area. See Frons and Bristles of the head.
Frontorbital plates (fig. 4). T h e sclerotized plates along the eyes o n w h i c h the frontorbital
bristles are inserted.
Frontorbital bristles (fig. 8: F; fig. 9: brisdes 1-6). See Brisdes of the head.
Gena (fig. 7, 8). T h e part o f the head b e l o w the l o w e r eye margin. In the Brachycera it often
possesses strong brisdes of w h i c h the anterior one is the vibrissa.
Holoptic. T h e eyes are touching in their dorsal region; sometimes also in their ventral region
( % 5).
Innervertical brisdes (fig. 8,9: Iv). See Bristles o f the head.
Interfrontal area (fig. 8). See Bristles of the head.
Interfrontal bristles (fig. 8). See Bristles o f the head.
Iv-bristles (Innervertical bristles) (fig. 8,9: Iv). See Bristles of the head.
Lunula. A s o m e w h a t triangular plate above the antennal insertion, just underneath the ptilinal
suture (fig. 4 , 5 , 9 1 ) . In almost all Schizophora it is w e l l developed. A plate resembling a lunula
also occurs in some groups that hold a lower position in the system, e.g. the Syrphidae.
Mouthparts (Proboscis) (fig. 4 , 7 ) . In the Diptera, functional mouthparts are always sucking or
piercing, never biting or c h e w i n g . H o w e v e r , b l o o d s u c k i n g flies are usually referred to in
English as "biting". Laterally the mouthparts are provided w i t h palps. T h e s e consist o f five
s e g m e n t s i n m o s t N e m a t o c e r a , t w o in the l o w e r Brachycera and o n e in m o s t
Muscomorpha.
Nasus. Prolongation at the anterior end of the rostrum (fig. 22).
Oc-bristles (Ocellar bristles) (fig. 8 , 9 : O c ) . See Bristles o f the head.
Occiput. T h e entire back o f the head (fig. 7).
Ocellar bristles (fig. 8 , 9 : O c ) . See Bristles o f the head.
Ocellar triangle. T h e plate o n w h i c h the ocelli are found. It is nearly always larger than the tri
angle m a d e u p b y the ocelli themselves.
Ocelli (singular: ocellus). Single lenses in the dorsal area of the head. If present there are near
ly a l w a y s three of t h e m , arranged in a triangle (fig. 3 , 4 , 7 ) .
Ocular. Pertaining to the eye(s), e.g. ocular margin.
O m m a t i d i a (singular: o m m a t i d i u m ) . T h e facets o f the insect eye.
O m m a t r i c h i a (singular: o m m a t r i c h i u m ) . T h e hairs b e t w e e n the facets of the insect eye.
Oral angle (fig. 187,188). See Vibrissal angle.
Oral margin (Peristoma) (fig. 308,311,314). T h e l o w e r margin of the face.
Outervertical bristles (fig. 8 , 9 : O v ) . See Bristles of the head.
Ov-bristles (Outervertical bristles) (fig. 8,9: O v ) . See Bristles of the head.
P-bristles (Postocellar bristles) (fig. 8,9: P). See Bristles of the head.
Palp. See Mouthparts.
Parafacialia. In the Schizophora the plate along the eye b e t w e e n the frontorbital plate and the
gena, separated from the remainder o f the face b y the ptilinal suture (fig. 4 , 9 1 ) .
Peristoma. A n o t h e r term for oral margin (fig. 308,311,314).
Postocellar bristles (fig. 8 , 9 : P). See Bristles of the head.
Prefrons. See Face.
Proboscis (fig. 4 , 7 ) . See Mouthparts.
Pseudopostocellar bristles. In s o m e cases n o real P-bristles are present, but other bristles,
placed m o r e laterally or caudally are present.
Pseudovibrissae. Vibrissa-like bristles that are n o t exactly inserted on the vibrissal angle but
m o r e caudally, ventrally or laterally.
Ptilinal fissure. O t h e r term for Ptilinal suture.
Ptilinal suture. A suture, fissure, or g r o o v e , present in the Schizophora, usually in a horse
shoe shape, r u n n i n g above the lunula d o w n o n either side of the antennae along the inner
margin o f the parafacialia (the plates bordering the eyes) (fig. 4 , 5 , 8 , 9 1 ) . See also Ptilinum.
Ptilinum (fig. 10). In Schizophora an inflatable bag in the head that is used w h e n the emerg
ing fly leaves the puparium. T h e p t i l i n u m emerges above the antennae and thereby pushes
o p e n the p u p a r i u m valve. In this position the face is s w u n g into an anteroventral position.
Subsequently, the bag is retracted and the face regains its normal position, separated from
the other elements o f the head b y the ptilinal suture.
PVT-bristles (Postvertical bristles). O t h e r term for P-bristles.
R o s t r u m . In particular in some Nematocera the l o w e r part o f the head can be transformed
into a l o n g snout, the rostrum that m a y carry a nasus o n its anterior end (fig. 22).
Style (Stylus). See A n t e n n a .
Vertex. T h e t o p o f the head, behind the ocellar triangle. It is b o u n d e d b y the eyes, the frons
and the occiput (fig. 4).
Verticils. Hair-like setae, usually in rings, at the base o f the flagellar segments (fig. 6a).

18
Vibrissa. A usually strong bristle at the vibrissal angle (fig. 4 , 7 ) .
Vibrissal angle (Oral angle) (fig. 187,188). T h e point b e l o w the eye w h e r e the l o w e r margin o f
the gena meets the l o w e r margin of the face (oral margin) and w h e r e the vibrissa is insert
ed (fig. 4 , 7 ) .

THORAX
Ac-bristles (Acrostichal brisdes) (fig. 11). See Scutal brisdes.
Acrostichal bristles (fig. 11). See Scutal bristles.
Air inlet. See Spiracle.
A m p u l l a . See Greater ampulla.
Anatergite. Part of the laterotergite (fig. 12). See M a i n parts.
A n e p i m e r o n (fig. 12). See Main parts.
A n e p i s t e r n u m (fig. 12). See Main parts.
Dc-brisdes (Dorsocentral brisdes) (fig. 11). See Scutal brisdes.
Dorsocentral brisdes (fig. 11). See Scutal bristles.
Greater ampulla (fig. 12). A r o u n d bulge o n the lateral side o f the thorax just b e l o w the front
end of the w i n g attachment. T h i s bulge occurs in all calyptrate families, b u t is also found in
some other families, e.g. Periscelididae, m a n y Syrphidae and s o m e Psilidae, S c i o m y z i d a e
and Tephritidae.
Haltere (plural: Halteres) (fig. 3 , 1 2 , 41). A remnant o f the h i n d w i n g that has b e e n trans
formed into an organ of balance. Consisting o f a stem (peduncle) and a k n o b .
Humeral callus. A n o t h e r t e r m for p o s t p r o n o t u m (fig. 11,12).
H u m e r u s . A n o t h e r term for p o s t p r o n o t u m (fig. 11,12).
H y p o p l e u r o n . A n o t h e r t e r m for m e r o n (fig. 12).
Intra-alar brisdes (fig. 11). See Scutal brisdes.
Katatergite. Part of the laterotergite (fig. 12). See M a i n parts.
Katepimeron (fig. 12). See M a i n parts.
Katepisternum (fig. 12). See M a i n parts.
Laterotergite (fig. 12). See Main parts.
Main parts (fig. 11,12). T h e main parts o f the dorsal side of the thorax are the p r o n o t u m (usu
ally reduced), s c u t u m , scutellum and mediotergite. T h e transverse suture divides the scu
t u m into the presutural and postsutural parts. T h e subscutellum is found in b e t w e e n the
mediotergite and scutellum. It can be strongly c o n v e x and p i l l o w - s h a p e d (fig. 374, 399).
Laterally o f the s c u t u m s o m e m i n o r elements are found. T h e y generally have bristles w h i c h
are important for identification purposes. In an anterior to posterior order, these elements
are: p o s t p r o n o t u m (fig. 11,12), notopleuron (fig. 11,12) and postalar callus (fig. 11).
The lateral side of the thorax is made u p of, in the order of anterior to posterior and dor
sal to ventral (fig. 12): p r o n o t u m (usually reduced), propleuron consisting o f proepister-
n u m and proepimeron, anepisternum, katepisternum, anepimeron, laterotergite ( w h i c h
may comprise t w o parts: anatergite and katatergite), katepimeron and m e r o n .
Mediotergite (fig. 1 2 , 2 5 , 4 1 ) . See Main parts.
Meral bristles. Bristles o n the m e r o n , e.g. as present in certain Calyptrate families (fig. 386,
387).
Meron (fig. 12). See Main parts.
M e s o n o t u m . T e r m for s c u t u m + scutellum (fig. 3,11,12).
Mesopleuron. A n o t h e r t e r m for anepisternum (fig. 12).
Metasternum. S o m e t i m e s used as the n a m e for the plate b e t w e e n and in front of the coxae o f
the hind legs.
N o t o p l e u r o n (fig. 12). See Main parts.
Peduncle. See Haltere.
Pleurotergite. A n o t h e r term for laterotergite (fig. 12).
Postalar callus (fig. 11). See M a i n parts.
Postpronotal lobe. A n o t h e r t e r m for p o s t p r o n o t u m (fig. 11)
Postpronotum (fig. 11). See Main parts.

19
 R2+3
Rs I _ Sc-R R-M

ptilinum

13 hypothetical ground-plan
10 head Heleomyzidae of the Diptera wing
with extruding ptilinum
postsutural scutum costal break costal break
presuturai scutum (humeral break) (subcostal break)
acrostichal bristles
H
dorsocentral bristles

- postpronotum
L^^-posthumeral bristles
' ( y > presuturai bristle

cup DM-Cu
alula CuA2 \ BM-Cu
anal vein
(A1+CuA2)

14 wing Calliphoridae
(Calliphora)

preaplcal bristle
I apical bristles
(spurs)

^ ^ z * tarsal
v
^Caiti segments
\^^-5th

15 Diptera leg
subscutellum
haltere
- mediotergite
laterotergite
greater ampulla
posterior spiracle bristle -
meral bristles
katepimeron
anepimeron
meron V
kate pi sternum coxa 3
16 bristle (macrotrichium)
with socket (alveolus)
12 thorax Calliphoridae
Postscutellum. A n o t h e r t e r m for subscutellum (fig. 12).
Postsutural s c u t u m (fig. 11,12). See Main parts.
Precoxal bridge (fig. 133,134,170). A precoxal bridge is present w h e n the plate o n the ventral
side of the thorax b e t w e e n the neck and the fore legs (the prosternum) is confluent w i t h the
lateral plates o n w h i c h it borders (the episterna).
Prescutum. A n o t h e r term for presutural s c u t u m (fig. 11).
Presutural s c u t u m (fig. 11). See Main parts.
Proepimeron (fig. 12). See Main parts.
Proepisternum (fig. 12). See Main parts.
Pronotum. See Main parts.
Propleuron. See Main parts.
Prosternum. Plate o n the ventral side of the thorax b e t w e e n the neck and the fore legs (fig. 133,
134,171). See Precoxal bridge.
Pteropleuron. A n o t h e r term for anepimeron (fig. 12).
Scutal bristles. Several longitudinal r o w s o f brisdes are found o n the s c u t u m , in particular in
the higher Diptera. T h e y are important for identification, especially o n the generic and spe
cific level. G o i n g laterad from the middle the f o l l o w i n g r o w s o f brisdes occur (fig. 11): acros-
tichal bristles, dorsocentral brisdes and intra-alar bristles. In all series the distinction o f
insertion in front versus b e h i n d the transverse suture is made. T h e t e r m 'acrostichals 1+3'
means that one acrostichal bristle is found in front and three are situated b e h i n d the trans
verse suture.
Scutellum. See Main parts.
S c u t u m (fig. 3,11,12). See Main parts.
Spiracle (fig. 12). T h e air inlet. T h e lateral side of the thorax has t w o spiracles. T h e anterior spira
cle is behind the p r o n o t u m or proepisternum, the posterior spiracle is b e t w e e n the haltere
and the meron and can be partially or entirely covered. T h e w a y in w h i c h the spiracle is cov
ered is a character used to distinguish b e t w e e n some o f the Calyptrate families (fig. 402-404).
Sternopleuron. A n o t h e r t e r m for katepisternum (fig. 12).
Stigma. A n o t h e r t e r m for spiracle (fig. 12).
Subscutellum (fig. 12). See Main parts.
Thoracic suture. A n o t h e r term for transverse suture (fig. 11,12).
Transverse suture (fig. 11,12). T h e transverse suture extends o n the dorsal side o f the thorax,
anterior to the w i n g insertions, from one side to the other. In several families o f the
Nematocera this suture is V - s h a p e (fig. 25); in m a n y groups the m i d part o f the suture is less
distinct or e v e n absent.

WING
For orientation purposes, the w i n g s are considered to be fully extended s i d e w a y s .
Abbreviations of veins and crossveins are given in capitals (e.g. v e i n R ) , those of the w i n g cells
in l o w e r case (e.g. cell d).
W i n g cells are n a m e d after the v e i n o n their anterior side.
A w i n g cell is closed w h e n it is n o w h e r e b o u n d e d b y the w i n g margin (fig. 1 4 : cell 4+5 is
open, cell d m is closed).
A-vein(s) (Anal vein(s)) (fig. 13: A i , A 2 ; fig. 14: A1+CUA2). See Longitudinal veins.
Alula (fig. 14). A lobe-like extension o f the basal h i n d part o f the w i n g , situated in b e t w e e n
the anal lobe (where that is present) and the upper calypter, and separated from t h e m b y
notches or narrower parts of the w i n g .
A n a l angle. T h e angle of the basal part o f the hind margin o f the w i n g (fig. 79).
A n a l cell. A n o t h e r term for cell cup.
A n a l lobe. T h e lobe at the basal h i n d part o f the w i n g (fig. 13,79).
A n a l v e i n (fig. 13: A i , A 2 ; fig. 14: A1+CUA2). See Longitudinal veins.
Basal cells (fig. 13,14). Cells br, b m and cup, situated in the basal part of the w i n g .
Bifurcation of a v e i n . See Fork.

21
C - v e i n (Costa) (fig. 1 3 , 1 4 : C). See Longitudinal veins.
Calypter (Squama) (fig. 14). T h e t w o (upper and lower, i.e., relatively apical and basal) lobes at
the basal hind margin of the w i n g n e x t to the thorax.
Cell b m . O n e o f the basal w i n g cells (fig. 13,14), abbreviated from base of the Media.
Cell br. O n e o f the basal w i n g cells (fig. 13,14), abbreviated from base of the Radius.
Cell cup (Anal cell) (fig. 1 3 , 1 4 ) . O n e o f the basal w i n g cells, name not related to the words
" c u p " or "cup shaped" b u t abbreviated from posterior branch of the Cubitus.
Cell d m (Discal medial cell) (fig. 14). See Discal cell.
Cell gb (Greater basal cell). T h e cell resulting from fusion of cells br and b m (fig. 72,76-82).
Costa (fig. 1 3 , 1 4 : C). See Longitudinal veins.
Costal break (fig. 14: arrows). T h e costa can be interrupted once, t w i c e or not at all. This is an
important character, in particular in the identification o f Acalyptrate families. This charac
ter, h o w e v e r , is n o t always clear. Hence it has b e e n used as little as possible in the key. It is
a g o o d idea to l o o k at the w i n g from an oblique position and to light it from b e l o w instead
o f from above.
Crossvein B M - C u (fig. 14). Crossvein found in the higher Diptera in b e t w e e n cells b m and dm.
Crossvein D M - C u (fig. 14). Closing v e i n of cell d m .
Crossvein H (Humeral vein) (fig. 13,14). Crossvein in b e t w e e n the costa and subcosta in the
basal part o f the w i n g .
Crossvein M - C u (fig. 13). Crossvein found in the more primitive Diptera, in b e t w e e n cell b m
and the m o s t basal/posterior m cell.
Crossvein M - M (fig. 13). C r o s s v e i n found in the m o r e primitive Diptera, closing cell d poste
riorly.
Crossvein R - M (fig. 13,14). Crossvein in b e t w e e n cell br and the m o s t basal/posterior r cell.
C u - v e i n (Cubitus). See Longitudinal v e i n s .
C u b i t u s (Cu). See Longitudinal v e i n s .
Discal cell (cell d); discal-medial cell (cell d m ) (fig. 13: d; fig. 14: d m ) . T h e discal cell or discal-
medial cell is a closed cell in the central or posterior part of the w i n g . A discal cell is present
w h e n the cell is n o w h e r e bordered b y v e i n C u A i ; if the cell is bordered b y this vein, then it
is n a m e d discal-medial cell. T h e o u t e r m o s t closing v e i n of the discal cell is crossvein M - M
(fig. 13), in case o f the discal-medial cell it is crossvein D M - C u (fig. 14). T h e definition of the
discal cell can be applied to m o s t Nematocera. H o w e v e r , in the Nematocera as w e l l as in the
L o w e r Brachycera (the L o w e r M u s c o m o r p h a in particular) there are several families w i t h
genera for w h i c h the discal cell definition obtains, alongside other genera w h e r e that of the
discal-medial cell applies.
In this b o o k the t e r m discal cell is u s e d for all Nematocera and Brachycera excluding the
Eremoneura, w h e r e a s the t e r m cell d m is used for all families o f the Eremoneura (i.e., all
families from Atelestidae to, and including, the Hypodermatidae in the classification table
given above).
Discal-medial cell (cell dm) (fig. 14: d m ) . See Discal cell.
Fork. Point o f bifurcation of a vein, e.g. fig. 13: v e i n R2+3 bifurcates into R2 and R3 at the fork.
Greater basal cell. See Cell gb.
H u m e r a l break. A n interruption o f the costa just b e y o n d crossvein H (fig. 14).
H u m e r a l v e i n . See Crossvein H .
Longitudinal veins. T h e six longitudinal veins in the w i n g are, from the front margin to the hind
margin: Costa (fig. 1 3 , 1 4 : C) w h i c h occupies the front (anterior) margin; in some primitive
Diptera it occupies the entire w i n g margin, including the hind (posterior) margin. Subcosta
(fig. 13,14: Sc) is a weaker vein b e t w e e n costa and Ri. Radius (fig. 13,14: R) in its primitive lay
out s h o w s five branches reaching the margin, four of w h i c h have a c o m m o n stem, the radial
sector (vein Rs). Media (fig. 13,14: M) in its primitive layout has three branches reaching the
margin. Cubitus, in its primitive layout has t w o branches, the anterior o f w h i c h ramifies into
C u A i and Q 1 A 2 . A n a l vein, in the more primitive Diptera this is vein A i , in the higher Diptera
this is vein A1+C11A2. In the Tipulomorpha, a second anal v e i n is found, named A2; this vein

22
 is absent, or less developed, in m o s t other Diptera. T h e spurious vein (vena spuria) is not con
nected to any other vein and crosses R - M at a right angle (fig. 135).
Lower Calypter (fig. 14). A basal appendage at the hind margin o f the w i n g , close to the tho
rax and n e x t to the upper calypter.
M-vein(s) (Media) (fig. 13,14: M). See Longitudinal veins.
Media (fig. 13,14: M). See Longitudinal veins.
Petiolate (Petiole). H a v i n g a section of a v e i n (the petiole) b e t w e e n a closed cell and the w i n g
margin (fig. 109,389).
Pterostigma. See Stigma.
R-vein(s) (Radius) (fig. 13,14: R). See Longitudinal veins.
Radial sector (Rs-vein). Basal part or stem of the R veins in b e t w e e n v e i n R i and the other R-
veins (fig. 13: Rs).
Radius (fig. 13,14: R). See Longitudinal veins.
Rs-vein (fig. 13: Rs). See Radial sector.
Sc-vein (Subcosta) (fig. 13,14: Sc). See Longitudinal veins.
Spurious v e i n (Vena spuria), (fig. 135). See Longitudinal v e i n s .
Squama. A n o t h e r term for calypter (fig. 14).
Stem. Part of the v e i n before the ramification or fork.
Stigma (Pterostigma). Characteristic coloured area o f the w i n g surface alongside the costa
near the end of vein(s) Sc a n d / o r R i (fig. 71,165-168).
Subcosta (fig. 13,14: Sc). See Longitudinal v e i n s .
Subcostal break. Interruption of the costa, just before the point w h e r e the subcosta or v e i n R i
meets or (if reduced or merged) should m e e t the costa (fig. 14).
U p p e r Calypter (fig. 14). A basal appendage at the hind margin o f the w i n g , close to the tho
rax and n e x t to the L o w e r Calypter.
Vena spuria (Spurious vein) (fig. 135). See Longitudinal v e i n s .
W i n g tip. In case of fully extended w i n g , the p o i n t furthest a w a y from the body.

LEG
The leg consists o f the following major parts (fig. 15), g o i n g o u t w a r d from the thorax: coxa
(plural: coxae), trochanter, femur (plural: femora), tibia (plural: tibiae), tarsus (plural: tarsi),
claw, e m p o d i u m and pulvilli w h e n developed.
Basitarsus. See Tarsus.
Claw. See Tarsus.
Distitarsus. See Tarsus.
E m p o d i u m . See Tarsus.
Metatarsus. See Tarsus.
Pulvilli. See Tarsus.
Tarsomeres. See Tarsus.
Tarsus. T h e Tarsus is made u p of five segments (tarsomeres or tarsal segments) (fig. 3,15); the
first (basal) segment is also n a m e d metatarsus or basitarsus; the last (apical) one is also
called distitarsus. T h e fifth tarsomere includes a small terminal sclerite (acropod or post-
tarsus) bearing the t w o claws, and, w h e n developed, three appendages: t w o pulvilli w i t h
an e m p o d i u m in b e t w e e n , the latter can be bristle-like or o f the same shape as the pulvilli
(pulvilliform) (fig. 103,104).
Tibia. T h e tibia may, in some cases, have apical a n d / o r preapical brisdes, spurs, or spine(s)
(fig. 1 5 , 6 3 , 7 5 , 1 8 5 ) .
17 antenna Culicidae (Aedes) (female)

23 habitus Tipulidae
(Tpula)

18 antenna Culicidae (Aedes) (male)

CuA1
A1 CuA2
24 wing Mycetobiidae (Mycetobia)

19 antenna Tipulidae
(Dictenidia) (male)
style 3rd antennal segment
(1st flagellomere)
26 thorax
scutellum
mediotergite Cylindrotomidae
flagellomere
2nd (pedicel) (scapus)

20 antenna Tabanidae 25 thorax Tipulidae

27 wing Trichoceridae (Trichocera)

palp
(one seg
ment)

ments)
22 head Tipulidae
21 head calyptrate fly (Tipula) 28 wing Trichoceridae (Diazosma)
DENTIFICAT10N KEY

T h i s k e y i s a s o - c a l l e d d i c h o t o m o u s k e y : e v e r y c o u p l e t c o n t a i n s t w o a l t e r n a t i v e s . It i s
r e c o m m e n d e d t o read b o t h a l t e r n a t i v e s b e f o r e c o m i n g t o a d e c i s i o n a n d t o h a v e a l o o k
at t h e f a m i l y d e s c r i p t i o n s for a d d i t i o n a l d i a g n o s t i c characters. In order t o b e able t o
i d e n t i f y s p e c i m e n s i t is o f g r e a t i m p o r t a n c e t h a t t h e y are i n g o o d c o n d i t i o n . T h i s is
e s p e c i a l l y t r u e for t h e h e a d , w i n g s , l e g s a n d t h e b r i s t l e s o f t h e h e a d a n d l e g s (bristles
w h i c h are b r o k e n o f f still r e v e a l f o r m e r p r e s e n c e b y their s o c k e t , t h e a l v e o l u s , w h i c h
stays b e h i n d , fig. 16). For s p e c i m e n s larger t h a n 6 m m , a h a n d l e n s w i t h 2 0 X m a g n i f i c a
t i o n w i l l g e n e r a l l y b e sufficient. For s m a l l e r s p e c i m e n s a s t e r e o m i c r o s c o p e u p t o 40X
m a g n i f i c a t i o n is r e q u i r e d .

> Indicates a n o t e a b o u t (additional) c h a r a c t e r s , classification, o r s o m e t h i n g

t o b e a w a r e of.
( ) T h e n u m b e r s b e t w e e n b r a c k e t s refer t o p r e v i o u s c o u p l e t s so t h e p a t h b a c k
t h r o u g h t h e k e y c a n b e retraced easily.

1 B o d y n o t s t r o n g l y d o r s o v e n t r a l l y flattened. W i n g w e l l d e v e l o p e d , n o t
s n a p p e d off n e a r t h e b a s e , n o t s t r o n g l y r e d u c e d o r a b s e n t . 2
- B o d y s t r o n g l y d o r s o v e n t r a l l y flattened (fig. 4 4 1 - 4 4 3 ) a n d / o r w i n g s t r o n g
l y r e d u c e d , a b s e n t , o r s n a p p e d o f f n e a r t h e b a s e (fig. 4 2 7 - 4 5 1 ) . 191
2(1) A n t e n n a filiform o r r e s e m b l i n g a series o f b e a d s , w i t h m o r e t h a n 6 s e g
m e n t s ; all s e g m e n t s o r t h o s e b e y o n d t h e s e c o n d m o r e o r less s i m i l a r (fig.
17); a n t e n n a l s e g m e n t s s o m e t i m e s w i t h l o n g v e r t i c i l s (fig. 18) o r w i t h
a p p e n d a g e s (fig. 1 9 , 85). P a l p i n N e m a t o c e r a o f t e n w i t h 3-5 s e g m e n t s (fig.
22, 2 9 , 4 8 ) .
Nematocera (3-38) a n d Brachycera with a primitive antenna (39-43) - 3
A n t e n n a n o t filiform or r e s e m b l i n g a series o f b e a d s , w i t h l e s s t h a n 10 s e g
m e n t s ; t h e t h i r d s e g m e n t n e a r l y a l w a y s larger t h a n t h e s u b s e q u e n t s e g
m e n t s w h i c h , i n t h a t case, s t r o n g l y differ f r o m t h e t h i r d s e g m e n t , o r
t o g e t h e r c o n s t i t u t e a n ( u n ) s e g m e n t e d c o n t i n u a t i o n : a s t y l e (fig. 20) o r
arista (fig. 21). Palp u s u a l l y w i t h 1-2 s e g m e n t s (fig. 21). Brachycera - 4 4
3(2) W i n g w i t h 2 anal v e i n s ( v e i n s A i a n d A 2 ) r e a c h i n g t h e w i n g m a r g i n (fig.
23, w i n g figures 27-35). U p p e r side o f t h o r a x w i t h a V - s h a p e d t r a n s v e r s e
s u t u r e (fig. 25; s u t u r e i n C y l i n d r o t o m i d a e l e s s d i s t i n c t laterally, fig. 26). 4
O n l y 1 (or n o ) anal v e i n ( v e i n A i ) r e a c h i n g t h e w i n g m a r g i n (fig. 2 4 ) .
T r a n s v e r s e s u t u r e a b s e n t , unclear, i n t e r r u p t e d or, i f c o m p l e t e l y p r e s e n t
a n d V - s h a p e d , haltere w i t h a basal a p p e n d a g e (fig. 4 1 ) . 8
4(3) O c e l l i p r e s e n t . A n a l v e i n A 2 s h o r t a n d c u r v e d (fig. 27), l o n g e s t i n
Diazosma (fig. 28). Trichoceridae
O c e l l i a b s e n t (in v e r y f e w c a s e s , 2 r u d i m e n t a r y ocelli p r e s e n t , fig. 29). A n a l
v e i n A 2 a l m o s t a l w a y s at least h a l f as l o n g as v e i n A i (fig. 23, 30-35). 5
32 wing Limoniidae (Elliptera)

34 habitus Cylindrotomidae
(Cylindrotoma)

26
5(4) E y e p u b e s c e n t , w i t h o m m a t r i c h i a i n b e t w e e n t h e facets. C r o s s v e i n S c - R
l o c a t e d b e f o r e t h e o r i g i n o f v e i n R s ; Sc l o n g , p a s t t h e fork o f R s (fig. 30).
Pediciidae
- E y e bare, w i t h o u t o m m a t r i c h i a . C r o s s v e i n S c - R u s u a l l y l o c a t e d p a s t t h e
o r i g i n o f v e i n R s (fig. 3 1 , 3 3 , 3 5 ) , i f l o c a t e d b e f o r e t h e o r i g i n o f v e i n R s , t h e n
v e i n Sc n o t d i s t i n c t l y p a s t t h e fork o f R s (fig. 32). 6
6(5) Last (5th) s e g m e n t o f t h e p a l p e l o n g a t e , d i s t i n c t l y l o n g e r t h a n t h e p r e c e d
i n g s e g m e n t s ; r o s t r u m u s u a l l y w e l l d e v e l o p e d , o f t e n w i t h a n a s u s (fig. 22,
29). V e i n Sc u s u a l l y d o e s n o t e n d i n t h e c o s t a (fig. 33). A n t e n n a u s u a l l y
w i t h 13 s e g m e n t s or, i n a f e w c a s e s , m o r e . Tipulidae
Last palpal s e g m e n t s h o r t , i f s l i g h t l y e l o n g a t e t h e n c o m b i n a t i o n o f charac
ters n o t as g i v e n a b o v e . R o s t r u m u s u a l l y s h o r t and w i t h o u t n a s u s (fig. 3 4 ) ,
r o s t r u m s o m e t i m e s e l o n g a t e (fig. 31). V e i n S c u s u a l l y e n d s i n t h e c o s t a (fig.
31, 32; b u t n o t so i n fig. 35). A n t e n n a u s u a l l y w i t h 1 4 o r m o r e s e g m e n t s , i n
a f e w cases less t h a n 13. 7
7(6) A p i c a l p a r t o f v e i n Q 1 A 2 s t r o n g l y c u r v i n g t o w a r d t h e w i n g m a r g i n (fig.
35). V - s h a p e d t r a n s v e r s e s u t u r e o n u p p e r side o f t h o r a x less d i s t i n c t at t h e
sides (fig. 26). M a l e : apical p a r t o f t h e d i s t i p h a l l u s c o n s i s t s o f 2 or 3 separate
s m a l l t u b e s ( w e l l v i s i b l e f r o m o u t s i d e ) (fig. 36). F e m a l e : u p p e r v a l v e s o f
t h e o v i p o s i t o r (cerci) s h o r t a n d w i d e (fig. 37). Cylindrotomidae
Apical part o f vein C u A 2 usually n o t strongly c u r v i n g toward the w i n g
m a r g i n (fig. 32). V - s h a p e d t r a n s v e r s e s u t u r e o n u p p e r side o f t h o r a x d i s
t i n c t t h r o u g h o u t (fig. 25). M a l e : apical p a r t o f t h e d i s t i p h a l l u s w i t h n o
m o r e t h a n 2 o r 3 ( u s u a l l y o ) separate o p e n i n g s w h i c h are h a r d l y o r n o t at
all v i s i b l e f r o m o u t s i d e . F e m a l e : u p p e r v a l v e s o f t h e o v i p o s i t o r (cerci) u s u
ally l o n g , s l e n d e r a n d a c u t e (fig. 31). Limoniidae
8(3) D i s c a l cell o r cell d m a b s e n t ( w i n g figures 39-83). 9
- D i s c a l cell o r cell d m p r e s e n t (fig. 38). 38
9(8) O c e l l i absent. 10
O c e l l i p r e s e n t (fig. 5 8 , 7 3 ) . 19
10(9) Nine or more longitudinal veins or longitudinal v e i n branches reaching
t h e w i n g m a r g i n , i n c l u d i n g t h e anal v e i n ( A i ) ( w i n g figures 3 9 - 4 5 ) . 11
- N o more than eight longitudinal veins or longitudinal vein branches
r e a c h i n g t h e w i n g m a r g i n ( w i n g figures 4 7 - 5 3 ) . 15

^7
 spurious vein

fold CuA2
40 wing Ptychopteridae (Ptychoptera)

45 habitus Culicidae
(Aedes) (female)
mediotergite
basal appendage

41 haltere Ptychopteridae
(Ptychoptera)

46 head Chaoboridae
(Chaoborus) (male)

42a wing Psychodidae / f i i n R1 R2+3

(Trichomyia)
42 habitus Psychodidae
(Tinearia) R 2 + 3 f o r k

R2+3
Rs

47 wing Thaumaleidae (Thaumalea)

48 head Thaumaleidae
(Thaumalea)

43 habitus Dixidae (Dixa)

2nd antennal segment
1 st antennal segment

44 antenna Dixidae 49 habitus Cecidomyiidae

(Dixella) (Cecidomyia)

28
II(IO) H a l t e r e w i t h basal a p p e n d a g e (fig. 4 1 ) . First a n t e n n a l s e g m e n t larger t h a n
s e c o n d . U p p e r side o f t h o r a x w i t h U - o r V - s h a p e d t r a n s v e r s e s u t u r e .
W i n g v e n a t i o n as i n fig. 4 0 , w i t h a s p u r i o u s v e i n b e t w e e n t h e R a n d M
veins. Ptychopteridae
(In E u r o p e 1 g e n u s , Ptychoptera, w i t h a b o u t 15 s p e c i e s . )
H a l t e r e w i t h o u t basal a p p e n d a g e . First a n t e n n a l s e g m e n t u s u a l l y r i n g -
shaped, s e c o n d s e g m e n t largest a n d spherical (fig. 6a, 4 4 , 4 6 ) .
C o m b i n a t i o n o f o t h e r c h a r a c t e r s n o t as g i v e n a b o v e . 12
12(11) C r o s s v e i n s near t h e w i n g b a s e o r c r o s s v e i n s (partly) a b s e n t (fig. 4 2 , 4 2 a ) ;
w i n g o f t e n s t r o n g l y h a i r y a n d a c u t e (fig. 4 2 ) , s o m e t i m e s r o u n d e d (fig.
42a); c o s t a s u r r o u n d i n g t h e w i n g c o m p l e t e l y [ n o t clear f r o m t h e d r a w i n g
but it should b e from the insect]. Psychodidae
- C r o s s v e i n s n o t l i m i t e d t o n e a r t h e b a s e ; w i n g n o t a c u t e (fig. 4 3 , 4 5 ) ; c o s t a
surrounding the w i n g completely or not. 13
13(12) V e i n R2+3 n o t i n l i n e w i t h v e i n R s , b u t c u r v e s t o w a r d t h e fork; w i n g w i t h
p u b e s c e n c e o n t h e v e i n s (fig. 4 3 ) . A n t e n n a l s e g m e n t s w i t h o u t l o n g v e r t i
cils near their b a s e s (fig. 4 4 ) . Dixidae
- V e i n R2+3 i n l i n e w i t h v e i n R s (fig. 3 9 , 4 5 ) ; w i n g o n t h e v e i n s a n d e s p e c i a l
ly along the h i n d m a r g i n c o n s p i c u o u s l y hairy or set w i t h scales. A n t e n n a l
s e g m e n t s w i t h l o n g b a s a l v e r t i c i l s (fig. 4 5 , 4 6 ) . 14
14(13) Scales present o n t h e w i n g m a r g i n and v e i n s , t h e legs and usually o n t h e
a b d o m e n . M o u t h p a r t s c o n s p i c u o u s l y e l o n g a t e (fig. 4 5 ) , f e m a l e b l o o d s u c k
ing. Culicidae
> [In s p e c i m e n s p r e s e r v e d i n l i q u i d t h e scales are e a s i l y lost.]
- S c a l e s v i r t u a l l y e x c l u s i v e l y o n t h e w i n g m a r g i n , t h e scales o n t h e v e i n s
n a r r o w o r r e s e m b l i n g h a i r - l i k e b r i s t l e s ; t h e rest o f t h e w i n g w i t h p u b e s
c e n c e o n l y . M o u t h p a r t s n o t c o n s p i c u o u s l y e l o n g a t e (fig. 4 6 ) , f e m a l e n o t
bloodsucking. Chaoboridae
15(10) W i n g v e n a t i o n as i n fig. 4 7 : c r o s s v e i n s n e a r t h e w i n g b a s e a n d w i t h a
c r o s s v e i n b e t w e e n v e i n s R i a n d R2+3; v e i n R2+3 c u r v i n g ; c o s t a s u r r o u n d
i n g t h e w i n g c o m p l e t e l y [not clear f r o m t h e d r a w i n g b u t i t s h o u l d b e f r o m
t h e i n s e c t ] . E y e s h o l o p t i c i n b o t h s e x e s . A n t e n n a as i nfig.4 8 : first a n d s e c
ond antennal segments w i d e n e d , t h e other segments narrower.
Thaumaleidae
- C o m b i n a t i o n o f characters n o t as g i v e n a b o v e . 16
16 (15) Legs w i t h first tarsal s e g m e n t m u c h s h o r t e r t h a n s e c o n d , o r w i t h at m o s t 4
tarsal s e g m e n t s (fig. 4 9 ) . O n l y a l i m i t e d n u m b e r o f l o n g i t u d i n a l v e i n s
( u s u a l l y 2 t o 4 , at m o s t 6) r e a c h o r g o i n t h e d i r e c t i o n o f t h e w i n g m a r g i n .
M a r g i n g e n e r a l l y w i t h a b r e a k p a s t v e i n R 4 + 5 (fig. 57). U s u a l l y s m a l l , d e l i
cate i n s e c t s w i t h c o n s p i c u o u s l y l o n g a n t e n n a .
Cecidomyiidae (except Lestremiinae)
- First tarsal s e g m e n t n o t c o n s p i c u o u s l y s h o r t e r t h a n t h e s e c o n d a n d t a r s u s
w i t h 5 segments. Usually 6 or m o r e longitudinal veins reach or g o in the
direction of the w i n g margin. 17

29
52 wing Chironomidae (spec.) 57 wing Cecidomyiidae (Groveriella)

54 wing Blephariceridae (Liponeura) 59 habitus Scatopsidae

(Scatopse)

jo
17(16) V e i n s C u A i a n d C11A2 are separate f r o m e a c h o t h e r r i g h t f r o m t h e w i n g
base; w i n g w i d e ; costa and R-veins m o r e c o n s p i c u o u s than t h e other
v e i n s ; v e i n Q 1 A 2 u s u a l l y c u r v i n g l i k e a n e l o n g a t e , i n v e r t e d S (fig. 50); v e i n
R s n o t f o r k e d (fig. 50) o r f o r k e d i n R2+3 a n d R 4 + 5 . A n t e n n a s h o r t , w i d e ,
t a p e r i n g t o w a r d t h e a p e x , w i t h 11 (rarely 9) s e g m e n t s (fig. 51). Simuliidae
V e n a t i o n and a n t e n n a different. V e i n s C u A i a n d C u A 2 h a v e a l o n g j o i n t
s t e m s t a r t i n g f r o m t h e w i n g b a s e (fig. 52, 53; s t e m m a y b e t r a n s p a r e n t o r
vestigial, especially i n small species). 18
18(17) V e i n M c o n t i n u i n g singly, n o t f o r k e d ; R - v e i n s n o t s h o r t e n e d w i t h R 4 + 5
e n d i n g c l o s e t o the w i n g tip (fig. 52). N o p i e r c i n g m o u t h p a r t s .
Chironomidae
V e i n M forked into M i a n d M 2 a n d / o r R - v e i n s shortened; usually b o t h
characters p r e s e n t t o g e t h e r (fig. 53); v e i n M 2 o f t e n r e d u c e d or p o o r l y v i s i
ble near t h e fork; i f r e d u c e d t h r o u g h o u t , t h e n R - v e i n s s h o r t e n e d ;
c r o s s v e i n R - M a b s e n t i n Leptoconops. F e m a l e w i t h p i e r c i n g m o u t h p a r t s .
Ceratopogonidae
19(9) W i n g w i t h a g r i d o f s e c o n d a r y v e i n s a n d a c o n s p i c u o u s anal l o b e ; 1 o r 2 o f
the M - v e i n s reach the w i n g margin; if M2 present t h e n n o t connected to
M i (fig. 5 4 ) . E y e o f t e n s h o w t w o separate, differently faceted p a r t s (fig. 55).
Legs long. Blephariceridae
- W i n g w i t h o u t s e c o n d a r y v e i n s a n d c o m b i n a t i o n o f o t h e r characters n o t as
given above. 20
20(19) T i b i a e o f m i d a n d h i n d l e g s w i t h o u t apical b r i s t l e s o r s p u r s (fig. 5 9 , 62). 21
T i b i a e o f m i d and h i n d l e g s w i t h apical b r i s t l e s o r s p u r s (fig. 6 3 , 7 5 , 7 8 ) . 25
21(20) W i n g w i t h 4 R - v e i n s o f w h i c h R 2 is s h o r t a n d e n d s near R i (fig. 56).
Axymyiidae
(In E u r o p e 1 s p e c i e s , Mesaxymyia kerteszi ( D u d a ) . )
- N o m o r e t h a n 3 R - v e i n s (fig. 5 7 , 5 9 - 6 1 ) . 22
22(21) C o s t a s u r r o u n d i n g t h e w i n g c o m p l e t e l y [not clear f r o m t h e d r a w i n g b u t i t
s h o u l d b e f r o m t h e insect] and u s u a l l y i n t e r r u p t e d j u s t b e y o n d v e i n R 4 + 5 ;
o n l y a s m a l l n u m b e r o f v e i n s (6 or less) r e a c h i n g t h e w i n g m a r g i n (fig. 57).
S m a l l , delicate i n s e c t s u s u a l l y w i t h s t r i k i n g l y l o n g a n t e n n a .
Cecidomyiidae: Lestremiinae
- C o s t a n o t s u r r o u n d i n g all o f t h e w i n g b u t e n d i n g at or b e f o r e t h e w i n g t i p
(fig. 59). C o m b i n a t i o n o f o t h e r characters n o t as g i v e n a b o v e . 23
23(22) Palp w i t h 1 s e g m e n t o n l y (fig. 58). W i n g s h o r t , w i d e a n d w i t h anal l o b e ;
c o s t a e n d s w e l l b e f o r e t h e w i n g tip, n e a r l y a l w a y s n e a r t h e p o i n t w h e r e
R 4 + 5 r e a c h e s t h e w i n g m a r g i n (fig. 59). Scatopsidae

31
 R1-3

I absent , . . . & S S * M2 66 wing Hesperinidae (Hesperinus)

CuA2 CuA1
60 wing Synneuridae (Synneuron) fork Rs
Rs BM-u R;M So R \ R 1 R 2 + 3

R1 R2+3

A1 :
;i..--**^CuA2

CuA2 CuA1 67 wing Pleciidae (Plecia)

61 wing Canthyloscelidae
(Hyperoscelis)

62 hind leg
Canthyloscelidae
(Hyperoscelis)
-R4+5

68 habitus Pleciidae
(Penthetria funebris)
(male)
69 head Sciaridae (Pnyxia)

stem fork M

64 tibia fore leg 70 wing Sciaridae (Pnyxia)

Bibionidae (Dilophus)
63 habitus Bibionidae (Bibio)
R 1 stigma

65 head Hesperinidae
(Hesperinus)

71 wing Pachyneuridae (Pachyneura)

3 2
 Palp w i t h 2 o r m o r e s e g m e n t s . W i n g n a r r o w e r , w i t h o u t anal l o b e ; c o s t a
e n d s closer t o t h e w i n g t i p (fig. 6 0 , 61). 24
24(23) A n a l v e i n ( A i ) a b s e n t o r p r e s e n t as a basal r e m n a n t o n l y ; v e i n s R i a n d R s
c o n t i g u o u s o v e r a s h o r t stretch; M - f o r k i n c o m p l e t e , i.e., v e i n M 2 i s n o t
c o n n e c t e d t o v e i n M i (fig. 6 0 ) . F e m u r a n d tibia o f h i n d l e g slender.
A n t e n n a w i t h 12 s e g m e n t s . Synneuridae
(In E u r o p e 1 s p e c i e s : Synneuron annulipes Lundstrdm.)
A n a l v e i n ( A i ) d i s t i n c t l y v i s i b l e ; v e i n s R i a n d R s separate; M - f o r k u s u a l l y
c o m p l e t e (fig. 61). H i n d l e g w i t h apical h a l f o f f e m u r s w o l l e n a n d tibia
c u r v e d (fig. 62). A n t e n n a w i t h 16 s e g m e n t s . Canthyloscelidae
(In E u r o p e 1 g e n u s , Hyperoscelis, w i t h 2 species.)
25(20) A n t e n n a i n s e r t e d at l o w e r p a r t o f t h e h e a d , u s u a l l y j u s t a b o v e t h e l o w e r
m a r g i n o f the face (fig. 6 3 , 6 5 , 6 9 ) . 26
- A n t e n n a i n s e r t e d h a l f w a y u p t h e e y e , o r h i g h e r (fig. 7 3 , 7 8 ) . 29
26 (25) T h i r d a n t e n n a l s e g m e n t c o n s p i c u o u s l y e l o n g a t e , as l o n g as o r e v e n l o n g e r
t h a n t h e t w o f o l l o w i n g s e g m e n t s a n d as l o n g as (female) o r 1,5 t o 2 t i m e s
as l o n g as t h e h e a d (male) (fig. 65). W i n g v e n a t i o n as i n fig. 6 6 , m o r e o r less
similar t o t h a t o f t h e Pleciidae (fig. 67), b u t w i t h v e i n R 4 + 5 m o r e s t r o n g l y
curved. Hesperinidae
(In E u r o p e 1 s p e c i e s , Hesperinus imbecillus ( L o e w ) . )
T h i r d a n t e n n a l s e g m e n t a n d w i n g v e n a t i o n n o t as a b o v e (fig. 6 3 , 6 7 - 7 0 ) .
27
27(26) V e i n R f o r k e d i n t o R2+3 a n d R 4 + 5 w i t h v e i n R 4 + 5 c o n t i n u i n g m o r e o r l e s s
straight b e y o n d t h e fork (fig. 67) (in Penthetriafunebris M e i g e n , t h e w i n g
is d a r k e n e d and, i n t h e m a l e , s h o r t e r a n d n a r r o w e r , fig. 68). Pleciidae
(In E u r o p e 1 g e n u s , Penthetria, w i t h 2 species.)
V e i n R n o t f o r k e d ( v e i n R2+3 absent) (fig. 6 3 , 7 0 ) . 28
28(27) T i b i a o f fore l e g o f t e n s w o l l e n , i n Bibio w i t h a l o n g , s t o u t s p i n e a n d a n api
cal p r o j e c t i o n (fig. 63); i n Dilophus w i t h s e v e r a l r o w s o f s h o r t , s t r o n g
s p i n e s (fig. 6 4 ) . i Bibionidae
T i b i a o f fore l e g s i m p l e , w i t h o u t s p i n e s o r p r o j e c t i o n s . W i n g w i t h a l i m i t
e d n u m b e r o f l o n g i t u d i n a l v e i n s ; s t e m o f v e i n M is a b o u t as l o n g as p a r t
b e y o n d t h e M fork (fig. 7 0 ) . Sciaridae
> [ M o s t Sciaridae h a v e t h e a n t e n n a i n s e r t e d h a l f w a y u p t h e e y e w i t h a n e y e
b r i d g e a b o v e t h e a n t e n n a e (fig. 73; c o u p l e t 31); i n s o m e s p e c i e s t h e a n t e n
na is i n s e r t e d m u c h l o w e r a n d / o r t h e e y e b r i d g e is a b s e n t (fig. 69).]
2 2
9( 5) 4 R - v e i n s r e a c h i n g t h e w i n g m a r g i n (fig. 71). Pachyneuridae
(In E u r o p e 1 s p e c i e s , Pachynema fasciata Zetterstedt.)

33
72 wing Mycetobiidae (Mycetobia) " 'i""cuA2
A

76 wing Ditomyiidae (Ditomyia)

73 head Sciaridae (Sciara) 77 wing Keroplatidae (Keroplatinae) (Keroplatus)

stem M fork M

34
 i t o 3 R - v e i n s r e a c h i n g the w i n g m a r g i n ( w i n g figures 72-83). 30
30(29) W i n g w i t h a large basal cell g b (fig. 72: cells b r a n d b m f u s e d i n t o a s i n g l e
cell) f r o m w h i c h 6 v e i n s o r i g i n a t e ; R - f o r k b e f o r e o r at t h e s a m e l e v e l as
c r o s s v e i n R - M (fig. 72). C o x a e e l o n g a t e . Mycetobiidae
C e l l b m n o t c l o s e d , i.e., o p e n t o t h e w i n g m a r g i n (fig. 7 4 , 7 5 ) or, i f cell g b is
p r e s e n t , t h e n at m o s t 5 v e i n s o r i g i n a t e f r o m this cell (fig. 7 6 - 8 2 : cell g b ) ; R-
fork, i f p r e s e n t , b e y o n d c r o s s v e i n R - M (fig. 7 6 - 8 1 ) . C o x a e e l o n g a t e o r n o t .
3i
31(30) E y e s w i t h a n a r r o w e y e b r i d g e a b o v e t h e a n t e n n a e (fig. 73). S t e m o f v e i n M
m o r e o r less o f t h e s a m e l e n g t h as M i a n d M 2 ; cell m f r e q u e n t l y b e l l -
s h a p e d (fig. 74). C o x a e n o t c o n s p i c u o u s l y e l o n g a t e (fig. 7 5 , 7 8 ) . Sciaridae
E y e s separate. S t e m o f v e i n M s h o r t e r t h a n M i a n d M 2 ; cell m n e a r l y
a l w a y s n a r r o w , e l o n g a t e , n o t b e l l - s h a p e d (fig. 75-83). C o x a e u s u a l l y c o n
spicuously elongate. 32
32(31) V e i n C u A i a n d s t e m o f M n o t c o n n e c t e d o r c o n n e c t e d at t h e l e v e l o f
c r o s s v e i n H ; v e i n A i n o t r e a c h i n g w i n g m a r g i n (fig. 75). Mycetophilidae
V e i n C u A i c o n n e c t e d t o s t e m o f M t h r o u g h c r o s s v e i n M - C u (fig. 7 6 - 8 3 ) ;
this c o n n e c t i o n is a l w a y s m u c h m o r e apical t h a n t h e l o c a t i o n o f c r o s s v e i n
H ; v e i n A i i n m o s t c a s e s r e a c h i n g w i n g m a r g i n , at least as a d i s t i n c t fold
(fig. 7 6 - 8 2 ) . 33
2
33(3 ) V e i n R2+3 m o r e t h a n h a l f as l o n g as R 4 + 5 ; o r i g i n o f v e i n Sc s t r o n g , t h e rest
p r e s e n t o n l y as a fold a n d e n d i n g free (fig. 7 6 ) . Ditomyiidae
V e i n R2+3 less t h a n h a l f as l o n g as R 4 + 5 (fig. 77-81) o r a b s e n t (fig. 82, 83);
v e i n Sc n e a r l y a l w a y s l o n g , e n d i n g i n c o s t a o r i n v e i n R i (fig. 77-81). 34
34(33) S t e m o f v e i n M o r i g i n a t e s f r o m v e i n R s o r R (fig. 7 7 - 7 9 ) . 35
S t e m o f v e i n M and R s are c o n n e c t e d b y c r o s s v e i n R - M (fig. 81-83). 36
35(34) A n t e n n a relatively s h o r t , a t m o s t as l o n g as h e a d a n d t h o r a x t o g e t h e r (fig.
78). A n a l l o b e n o t e n l a r g e d (fig. 7 7 , 7 8 ) .
Keroplatidae, subfamily Keroplatinae
A n t e n n a m o r e t h a n t w o t h i r d s b o d y l e n g t h (fig. 80). A n a l l o b e large, w i t h
a (nearly) r i g h t anal angle (fig. 7 9 , 80).
Keroplatidae, subfamily Macrocerinae

35
 88 habitus
Stratiomyidae
(Beris)

36
V e i n R2+3 p r e s e n t (fig. 81). Bolitophilidae
(In E u r o p e 1 g e n u s , Bolitophila, w i t h a b o u t 36 species.)
V e i n R2+3 a b s e n t (fig. 82, 83). 37
V e i n R 4 + 5 i n a l i g n m e n t w i t h v e i n R s ; c r o s s v e i n s R - M a n d M - C u i n line
w h i c h each o t h e r (fig. 82). Diadocidiidae
(In E u r o p e 1 g e n u s , Diadocidia, w i t h 5 species.)
V e i n R s m o r e or less vertical w i t h respect t o v e i n R4+5; crossvein R - M
m o r e i n line w i t h v e i n R 4 + 5 (fig. 83). Heterotricha a n d Sciarosoma
[ T h e g e n e r a Heterotricha ( w i t h o n l y 1 E u r o p e a n s p e c i e s , H. takkae
C h a n d l e r , k n o w n f r o m S w i t z e r l a n d , Italy a n d G r e e c e ) , a n d Sciarosoma
(also w i t h o n l y 1 E u r o p e a n s p e c i e s , S. borealis C h a n d l e r , k n o w n f r o m
Fennoscandia, Northwestern Russia, G e r m a n y and the C z e c h Republic),
are 2 p r i m i t i v e S c i a r o i d e a g e n e r a n o t c u r r e n t l y a s s i g n e d t o a f a m i l y ; f o r
details see C h a n d l e r 2 0 0 2 , K a l l w e i t bk. J a s c h h o f 2 0 0 4 , J a s c h h o f et al. 2006.]
V e i n R 4 + 5 n o t forked; cell c u p w i d e l y o p e n t o w a r d w i n g m a r g i n (fig. 8 4 ) .
A n t e n n a w i t h o v e r 1 4 s e g m e n t s . Palp w i t h 4 s e g m e n t s . Anisopodidae
(In E u r o p e 1 g e n u s , Sylvicola, w i t h a b o u t 1 0 species.)
V e i n R 4 + 5 nearly a l w a y s forked; i f n o t s o , t h e n a n t e n n a w i t h less t h a n 1 4
s e g m e n t s ; cell c u p s t r o n g l y n a r r o w i n g t o w a r d o r c l o s e d b e f o r e w i n g m a r
g i n (fig. 8 6 - 9 0 ) . Palp w i t h 1 t o 3 s e g m e n t s . 39
M o r e t h a n 11 a n t e n n a l s e g m e n t s p r e s e n t (fig. 85) ( t h e o n l y E u r o p e a n
s p e c i e s h a s o v e r 30 a n t e n n a l s e g m e n t s ) . W i n g v e n a t i o n m o r e o r less as i n
fig. 86: cell m 3 c l o s e d , v e i n R 5 e n d s n e a r t h e w i n g tip. Rachiceridae
(In E u r o p e 1 s p e c i e s , Rachicerus tristis ( L o e w ) . )
N o m o r e t h a n 11 a n t e n n a l s e g m e n t s . 40
C e l l m 3 , t h e cell b e l o w t h e discal cell, c l o s e d ( v e i n s M 3 a n d C u A i m e e t
b e f o r e , fig. 8 6 , or at t h e w i n g m a r g i n , fig. 87). 41
C e l l 1T13 o p e n (fig. 8 8 - 9 0 ) . 42
V e i n R 5 e n d i n g near t h e w i n g tip; alula n o r m a l l y d e v e l o p e d (fig. 86); c o s t a
d o e s n o t s u r r o u n d all o f t h e w i n g b u t r e a c h e s at m o s t u p t o v e i n M i .
Xylomyidae
V e i n R 5 e n d i n g d i s t i n c t l y b e y o n d t h e w i n g tip; alula s m a l l o r a b s e n t (fig.
87); costa s u r r o u n d i n g t h e w i n g c o m p l e t e l y [ n o t clear f r o m t h e d r a w i n g
b u t it should be from the insect]. Vermileonidae
R - v e i n s i n t h e front p a r t o f t h e w i n g w i t h v e i n R 5 a l m o s t a l w a y s r e a c h i n g
t h e w i n g m a r g i n b e f o r e t h e w i n g tip; discal cell d i a m o n d s h a p e d a n d u s u
ally s t r i k i n g l y s m a l l (fig. 88); costa d o e s n o t s u r r o u n d all o f t h e w i n g b u t
c o n t i n u e s u p t o j u s t b e y o n d t h e w i n g tip. Stratiomyidae

37
 94 head
Rhagionk
(Arthrocei

lower cali
92 head
Tabanidae
(Haematopota)

93 head Asilidae
38 98 habitus Acroceridae (Ogcod
(Stichopogon)
 V e n a t i o n different; v e i n R 5 e n d i n g at o r d i s t i n c t l y b e y o n d t h e w i n g t i p ;
discal cell e l o n g a t e (fig. 8 9 , 9 0 ) ; c o s t a s u r r o u n d i n g t h e w i n g c o m p l e t e l y
[not clear f r o m t h e d r a w i n g b u t i t s h o u l d b e f r o m t h e i n s e c t ] . 43
43(42) S c u t e l l u m w i t h 2 s p i n y o u t g r o w t h s (fig. 89a). A l u l a p r e s e n t ; cell c u p
s t r o n g l y n a r r o w i n g t o w a r d w i n g m a r g i n , b u t n e a r l y a l w a y s o p e n (fig. 8 9 ) .
Coenomyiidae
(In E u r o p e 1 w i d e s p r e a d s p e c i e s , Coenomyiaferruginea (Scopoli).)
S c u t e l l u m w i t h o u t o u t g r o w t h s . A l u l a s m a l l o r absent; cell c u p c l o s e d
( v e i n s A i a n d C11A2 m e e t b e f o r e o r at t h e w i n g m a r g i n ) (fig. 9 0 ) .
Xylophagidae
(In E u r o p e 1 g e n u s , Xylophagus, w i t h 5 species.)
44(2) It is p o s s i b l e t o s k i p c o u p l e t 4 4 a n d t o c o n t i n u e w i t h c o u p l e t 4 5 , i n w h i c h
case t h e d e c i s i o n a b o u t t h e p t i l i n a l s u t u r e i s p o s t p o n e d t o c o u p l e t 7 1 . A l l
f a m i l i e s w i t h a m o r e c o m p l e t e w i n g v e n a t i o n (e.g. fig. 9 9 , 1 0 0 ) o r w i t h a
l o n g s p u r i o u s v e i n (fig. 135) k e y o u t b e f o r e c o u p l e t 7 1 , t h e large m a j o r i t y o f
t h e f a m i l i e s w i t h a m o r e s i m p l i f i e d w i n g v e n a t i o n (e.g. fig. 1 0 1 , 1 0 2 ) k e y
o u t b e y o n d c o u p l e t 71.
- Ptilinal s u t u r e p r e s e n t (fig. 9 1 ; a u s u a l l y h o r s e s h o e s h a p e d g r o o v e o r
s u t u r e r u n n i n g a b o v e t h e l u n u l a d o w n o n e i t h e r side o f t h e a n t e n n a e , s e p
arating t h e parafacialia f r o m t h e r e m a i n d e r o f t h e face). W i n g v e n a t i o n
w i t h at m o s t 7 v e i n s r e a c h i n g t h e w i n g m a r g i n ( w i n g figures 1 0 2 , 1 7 9 - 4 2 0 ) ,
v e n a t i o n u s u a l l y s i m p l e , v e i n R 4 + 5 n e v e r f o r k e d , cell b m a l w a y s s h o r t (fig.
102) o r f u s e d w i t h cell d m (fig. 207).
Acalypttate and Calyptrate families - 81
Ptilinal s u t u r e a b s e n t ( a l t h o u g h a c l o s e d s u t u r e m i g h t b e a p p a r e n t a b o v e
t h e a n t e n n a e t h i s d o e s n o t e x t e n d b e l o w t h e a n t e n n a e (fig. 9 2 - 9 7 ) , i f a
s u t u r e is p r e s e n t t h e n t h i s s u r r o u n d s t h e c l y p e u s (fig. 9 4 , 9 5 ) a n d d o e s n o t
extend to above t h e antennae). Venation sometimes simple (e.g.
D o l i c h o p o d i d a e , fig. 162) b u t u s u a l l y v e r y m u c h different, as s h o w n i n
w i n g figures 9 8 - 1 0 1 , 1 0 5 - 1 6 8 ) , o f t e n cells br, b m a n d c u p l o n g a n d t h e r e
fore t h e discal cell, i f p r e s e n t , i n distal p a r t o f t h e w i n g , v e i n R 4 + 5 o f t e n
f o r k e d (fig. 1 0 0 ) . 45
45(44) T h o r a x h u m p b a c k e d a n d a b d o m e n u s u a l l y g l o b o s e (fig. 9 8 , 9 9 ) . H e a d
small, situated (somewhat) b e l o w the highly vaulted thorax; eyes h o l o p -
tic; a n t e n n a w i t h 3 s e g m e n t s o f w h i c h t h e first c a n b e v e r y s m a l l . W i n g
v e n a t i o n v a r i e s f r o m r e d u c e d (fig. 9 8 ) t o c o m p l e t e (fig. 9 9 ) ; l o w e r c a l y p t e r
c o n s p i c u o u s , large, e a r - s h a p e d . E m p o d i u m p u l v i l l i f o r m (last tarsal s e g
m e n t s w i t h 3 c u s h i o n s ; fig. 103). Acroceridae
C o m b i n a t i o n o f characters n o t as g i v e n a b o v e . 46
46(45) A t t h e s a m e t i m e v e i n R 4 + 5 forked i n t o R 4 a n d R 5 a n d cell c u p v e r y e l o n
g a t e (this cell c l o s e d at (fig. 1 0 0 ) o r n o t far b e f o r e (fig. 1 0 5 , 1 3 1 ) t h e w i n g
m a r g i n , o r o p e n t o w a r d t h e w i n g m a r g i n (fig. 1 1 3 , 1 2 9 ) ) , o r w i n g v e n a t i o n
aberrant as s h o w n i n fig. 1 0 7 - 1 0 9 . 47
N o t a t t h e s a m e t i m e v e i n R 4 + 5 f o r k e d i n t o R 4 a n d R 5 a n d cell c u p v e r y
e l o n g a t e (e.g. fig. 1 0 1 , 1 3 9 : R 4 + 5 n o t f o r k e d a n d cell c u p e l o n g a t e ; fig. 102:
R 4 + 5 n o t forked a n d cell c u p s h o r t ; fig. 133: R 4 + 5 f o r k e d a n d cell c u p s h o r t ;
fig. 228: R 4 + 5 n o t f o r k e d a n d cell c u p absent). 62

39
 cell m3
cell cup 1 CuA2

105 wing Athericidae (Ibisia)

106 antenna
Athericidae
99 habitus Acroceridae (Cyrtus) (Atherix)
diagonal vein R 1

cell br long cell bm long R4+5

107 wing Nemestrinidae (Fallenia

cell cup long (and closed)

100 wing Therevidae

(Thereva)

R4+5

-cell cup long (and open)

101 wing Mythicomyiidae (Empidideicus)

108 habitus Nemestrinidae (Nemestrinus)

102 wing Oestridae (Oestrus)
diagonal vein

petioles

109 wing Nemestrinidae (Trichopsidea)

2 1,
2 . ,L

a: Heptatoma pellucens
b: Haematopota pluvialis

103 5th tarsomere ^style 2 1

104 5th tarsomere
Stratiomyidae Muscidae
(Inopus) (Musca) c: Haematopota italica
d: Tabanus autumnalis
40
110 a-d antennae Tabanidae
 E m p o d i u m p u l v i l l i f o r m (last tarsal s e g m e n t s w i t h 3 c u s h i o n s ; fig. 103). 4 8
[In Lampromyia ( V e r m i l e o n i d a e ) p u l v i l l i a n d / o r e m p o d i u m sometimes
extremely reduced; such species can b e recognised b y the v e r y long
m o u t h p a r t s i n c o m b i n a t i o n w i t h t h e w i n g v e n a t i o n (fig. 114).]
E m p o d i u m b r i s t l e - l i k e or a b s e n t (last tarsal s e g m e n t s w i t h at m o s t 2 c u s h
ions) (fig. 1 0 4 ) . 56
[A k e y for m o s t o f t h e f a m i l i e s a n d g e n e r a w h i c h k e y o u t i n c o u p l e t s 4 8 - 5 5
is g i v e n b y N a g a t o m i 8t D i n g Y a n g 1998.]
V e i n s R i a n d R2+3 reach t h e c o s t a at o r a l m o s t at t h e s a m e p o i n t (fig. 105).
T h i r d a n t e n n a l s e g m e n t k i d n e y - s h a p e d (fig. 9 5 , 1 0 6 ) . Athericidae
V e i n s R i and R2+3 reach t h e c o s t a at a clear d i s t a n c e f r o m e a c h o t h e r ( m o s t
o f the w i n g figures 107-119) a n d / o r w i n g v e n a t i o n v e r y different (fig. 1 0 7 -
109,116); third antennal s e g m e n t virtually never kidney-shaped. 49
R- a n d M - v e i n s m o r e o r less c o n v e r g i n g t o w a r d c o s t a a n d w i n g tip, M -
v e i n s e n d i n g b e f o r e o r j u s t b e y o n d t h e w i n g tip; apical p a r t o f w i n g w i t h
several, u s u a l l y l o n g i t u d i n a l l y a l i g n e d cells t h a t m a y i n t u r n b e d i v i d e d
i n t o s m a l l e r cells; a s o - c a l l e d d i a g o n a l v e i n p r e s e n t (fig. 1 0 7 - 1 0 9 ) . H e a d
large, u s u a l l y w i t h e l o n g a t e m o u t h p a r t s . T i b i a e w i t h o u t apical bristles (fig.
108). Nemestrinidae
C o m b i n a t i o n o f characters n o t as g i v e n a b o v e . 50
L o w e r c a l y p t e r c o n s p i c u o u s l y large; v e i n R 5 r e a c h i n g t h e w i n g m a r g i n
d i s t i n c t l y b e y o n d t h e w i n g tip a n d fork o f v e i n s R 4 a n d R 5 u s u a l l y strik
i n g l y d i v e r g i n g (fig. 111). A n t e n n a w i t h large t h i r d s e g m e n t (fig. 110 a-d)
and u s u a l l y a t a p e r i n g s t y l e (fig. 110 d). Tabanidae
L o w e r c a l y p t e r s m a l l . O t h e r characters as s p e c i f i e d a b o v e or n o t , b u t rarely
in t h e a b o v e c o m b i n a t i o n . 51
2
A l u l a s m a l l o r a b s e n t (fig. 112-114). 5
A l u l a n o r m a l l y d e v e l o p e d (fig. 115-119). 53
C e l l c u p c l o s e d ( v e i n s A i a n d C u A 2 m e e t b e f o r e o r at t h e w i n g m a r g i n ) ;
v e i n R 5 e n d s near t h e w i n g t i p (fig. 112). A l l a n t e n n a l s e g m e n t s p a s t t h e
s e c o n d m o r e o r less similar (fig. 112 a). Xylophagidae
(In E u r o p e 1 g e n u s , Xylophagus, w i t h 5 species.)

R2+3 R4
R1

wingtip

112 habitus Xylophagidae

habitus Tabanidae (Tabanus) (Xylophagus)
(female)

41
115 habitus
Xylomyidae
(Xylomya)
C e l l c u p o p e n (fig. 113), or, i f c l o s e d , t h e n v e i n R 5 e n d i n g d i s t i n c t l y b e y o n d
the w i n g tip (fig. 114). A n t e n n a l s e g m e n t s p a s t t h e s e c o n d n o t similar, b u t
forming a style. Vermileonidae
C e l l m 3 , t h e cell b e l o w t h e discal cell, i s c l o s e d ( v e i n s M 3 a n d C u A i m e e t
b e f o r e o r at t h e w i n g m a r g i n , fig. 115). Xylomyidae
C e l l m 3 o p e n (fig. 116-119). 54
R - v e i n s s i t u a t e d i n front p a r t o f t h e w i n g w i t h R 5 v i r t u a l l y a l w a y s e n d i n g
w e l l b e f o r e t h e w i n g t i p ; discal cell d i a m o n d - s h a p e d a n d s m a l l , o f t e n
s t r i k i n g l y s m a l l (fig. 116); c o s t a n o t s u r r o u n d i n g all o f t h e w i n g b u t r e a c h
i n g at m o s t t o j u s t b e y o n d t h e w i n g tip. Stratiomyidae
W i n g v e n a t i o n different; v e i n R 5 e n d i n g at o r w e l l b e y o n d t h e w i n g tip;
discal cell e l o n g a t e (fig. 117-119); c o s t a s u r r o u n d i n g t h e w i n g c o m p l e t e l y
[not clear f r o m t h e d r a w i n g b u t i t s h o u l d b e f r o m t h e i n s e c t ] . 55
S c u t e l l u m w i t h 2 s p i n y o u t g r o w t h s (fig. 117a). W i n g v e n a t i o n as i n fig. 117:
cell c u p s t r o n g l y n a r r o w i n g t o w a r d t h e w i n g m a r g i n b u t nearly a l w a y s
o p e n ; 4 v e i n s o r i g i n a t i n g f r o m t h e discal cell reach t h e w i n g m a r g i n .
Coenomyiidae
(In E u r o p e 1 w i d e s p r e a d s p e c i e s , Coenomyiaferruginea (Scopoli).)
S c u t e l l u m w i t h o u t s p i n y o u t g r o w t h s . W i n g v e n a t i o n w i t h cell c u p o p e n
(fig. 118) o r c l o s e d (fig. 119) a n d u s u a l l y 3 v e i n s o r i g i n a t i n g f r o m t h e d i s c a l
cell r e a c h i n g t h e w i n g m a r g i n . Rhagionidae
V e i n s R 4 , R 5 a n d M c u r v i n g f o r w a r d s o t h a t at least v e i n R 4 e n d s i n v e i n
R i a n d w i t h at m o s t 2 b u t u s u a l l y j u s t 1 v e i n r e a c h i n g t h e h i n d m a r g i n o f
t h e w i n g (fig. 120). Last a n t e n n a l s e g m e n t c o n s p i c u o u s l y s w o l l e n (fig. 121)
or aberrant. Mydidae
W i n g v e n a t i o n a n d a n t e n n a different. 57
 M1
M2
discal cell absent

122 wing Hilarimorphidae

(Hilarimorpha)

mystax

123 head Asilidae

(Dioctria)

128 habitus Bomb'

(Bombylella) '
fork
Rs Rs Sc
124 head Asilidae (Proagonistes)

125 head
Scenopinidae cell cup open
(Scenopinus) discal cell cell below discal cell
(cell cual) broadly open

129 wing Bombyliidae

(Amphicosmus)

126 habitus Scenopinidae

(Scenopinus)

130 habitus Bombyliidae

(Apolysis)

cell below discal cell

(cell m3) narrowing

127 habitus Therevidae

(Thereva) 131 Brachystomj
(Brachystom

44
57(56) V e i n s R 4 + 5 a n d M1+2 are f o r k e d i n ( a l m o s t ) s i m i l a r m a n n e r a n d discal cell
a b s e n t (fig. 122). Hilarimorphidae
(In E u r o p e 1 g e n u s , Hilarimorpha, w i t h 2 species.)
C o m b i n a t i o n o f characters n o t as g i v e n a b o v e . 58

58(57) U p p e r part o f head, b e t w e e n the eyes, w i t h a saddle-shaped depression;

ocelli o n a separate e l e v a t i o n w h i c h , h o w e v e r , d o e s n o t p r o j e c t a b o v e t h e
u p p e r m a r g i n o f t h e e y e s (fig. 123, a r r o w ) ; face r e l a t i v e l y l o n g a n d n e a r l y
a l w a y s w i t h l o n g bristles, f o r m i n g t h e m y s t a x or b e a r d ; p o w e r f u l , p i e r c i n g
m o u t h p a r t s p r e s e n t (fig. 1 2 3 , 1 2 4 ) . Asilidae
C o m b i n a t i o n o f characters n o t as g i v e n a b o v e . 59
59(5) A n t e n n a w i t h 3 s e g m e n t s o f w h i c h t h e t h i r d i s t h e largest; t h i s s e g m e n t
w i t h a m i n u t e s t y l e a n d / o r forked at t h e a p e x (fig. 125). V e i n M i c u r v i n g
f o r w a r d , e n d i n g b e f o r e t h e w i n g t i p o r i n v e i n R 5 (fig. 126).
Scenopinidae
C o m b i n a t i o n o f characters n o t as g i v e n a b o v e . 60
60(59) T h e cell b e l o w t h e discal cell c l o s e d o r b e c o m i n g n a r r o w e r t o w a r d t h e
h i n d m a r g i n o f the w i n g (fig. 127). Therevidae
T h e cell b e l o w t h e discal cell o p e n a n d w i d e n i n g t o w a r d t h e h i n d m a r g i n
o f t h e w i n g (fig. 1 2 8 , 1 2 9 ) , o r discal cell a b s e n t (fig. 130). 61
6i(6o) V e i n S c e n d s i n t h e c o s t a b e y o n d t h e fork o f R s ; v e i n s R2+3 a n d R 4 o f t e n
s t r o n g l y c u r v i n g f o r w a r d ; cell c u p u s u a l l y o p e n t o t h e w i n g m a r g i n (fig.
1 2 8 , 1 2 9 ) ; i f c l o s e d , t h e n e n d i n g i n a n a c u t e a n g l e (fig. 130); w i n g o f t e n w i t h
conspicuous markings. Mouthparts m a y or m a y n o t constitute a long
sucking proboscis. Bombyliidae
V e i n S c e n d s i n t h e c o s t a b e f o r e t h e fork o f R s ; V e i n s R2+3 a n d R 4 n o t
s t r o n g l y c u r v e d ; cell c u p c l o s e d , e n d i n g o b t u s e l y ; w i n g w i t h o u t c o n s p i c
u o u s m a r k i n g s (fig. 131). B r a c h y s t o m a t i d a e : Brachystoma
62(46) V e i n R 4 + 5 forked (fig. 132-134). 63
V e i n R 4 + 5 n o t forked (all w i n g figures b e y o n d 134). 64

132 wing Scenopinidae

(Scenopinus)

style'

style -

precoxal bridge

134 habitus Empididae

precoxal bridge (Clinocera)

133 habitus Empididae

(Chelifera)
137 wing Syrphidae (Psilota)
cell cup

spurious vein 141 wing Mythicomyiidae

(Empidideicus)
R4+5

138 wing Conopidae (Conops)

142 habitus
Mythicomyiidae
(Cyrtisiopsis)

4 6
63(62) A n t e n n a w i t h 3 s e g m e n t s o f w h i c h t h e t h i r d is t h e largest; this s e g m e n t
s o m e t i m e s forked at t h e a p e x a n d / o r w i t h a m i n u t e s t y l e (fig. 125). 2 v e i n s
o r i g i n a t i n g f r o m cell d m r u n i n t h e d i r e c t i o n o f t h e w i n g m a r g i n , M i a n d
C u A i (fig. 132), v e i n M i e n d s i n v e i n R 5 or i n t h e w i n g m a r g i n c l o s e t o v e i n
R5. Scenopinidae
S t y l e v i r t u a l l y a l w a y s w e l l d e v e l o p e d (fig. 133, 134). 3 v e i n s o r i g i n a t i n g
f r o m cell d m reach t h e w i n g m a r g i n : M i , M 2 a n d C u A i (fig. 1 3 3 , 1 3 4 ) .
Empididae
> [Including the brachystomatid genera with vein R4+5 forked:
Brachystoma, Gloma, Pseudoheleodromia a n d Trichopeza .]
64(62) A l o n g i t u d i n a l s p u r i o u s v e i n ( v e n a spuria) p r e s e n t o n b o t h sides o f
c r o s s v e i n R - M ; v e i n M i d o e s n o t reach t h e w i n g m a r g i n b u t c u r v e s for
w a r d a n d e n d s i n v e i n R 4 + 5 before t h a t v e i n reaches t h e m a r g i n (cell H + 5
c l o s e d ) ; cell d m p r e s e n t ; cell c u p l o n g a n d e n d i n g a c u t e l y j u s t b e f o r e t h e
w i n g m a r g i n (fig. 135). Syrphidae
W i n g v e n a t i o n different; v e n a s p u r i a u s u a l l y a b s e n t , b u t i f p r e s e n t ( s o m e
C o n o p i d a e ) , t h e n restricted t o cell r4+5 (fig. 138). 65
> [In t h e S y r p h i d a e g e n e r a Eristalinus a n d Psilota t h e v e n a spuria is h a r d l y
i n d i c a t e d or absent, a n d c o n f u s i o n is p o s s i b l e w i t h s o m e C o n o p i d a e g e n
era t h a t also p o s s e s s a c l o s e d cell r4+5 (fig. 1 3 8 , 1 4 4 ) . F u r t h e r m o r e , s o m e
C o n o p i d a e d o p o s s e s s a v e n a spuria b u t this is a l w a y s r e s t r i c t e d t o cell
r4+5 (fig. 138). D i s c r i m i n a t i n g characters are: cell n c l o s e d a n d v e i n R 4 + 5
s t r o n g l y c u r v e d (fig. 136): S y r p h i d a e : Eristalinus; v e i n M i m o r e or less rec
tangular w i t h r e s p e c t t o v e i n R 4 + 5 (fig. 137): S y r p h i d a e : Psilota; v e i n C u A i
reaches t h e w i n g m a r g i n (fig. 1 3 8 , 1 4 4 ) ; v e n a spuria, i f p r e s e n t , r e s t r i c t e d
t o cell r4+5 (fig. 138): C o n o p i d a e . ]
65(64) H e a d large, h e m i s p h e r i c a l t o n e a r l y s p h e r i c a l , t h e e y e s o c c u p y i n g a l m o s t
all o f the h e a d . W i n g v e n a t i o n as i n fig. 139 or 1 4 0 : cell c u p l o n g a n d e n d
i n g a c u t e l y j u s t before t h e w i n g m a r g i n ; cell T4+5 n a r r o w i n g t o w a r d w i n g
m a r g i n b u t o p e n ; s o m e t i m e s v e i n M i a n d cell d m a b s e n t (fig. 1 4 0 ) .
Mouthparts small. Pipunculidae
- C o m b i n a t i o n o f characters n o t as g i v e n a b o v e ; i f cell c u p e l o n g a t e a n d
acute a n d cell r4+5 n a r r o w i n g , t h e n m o u t h p a r t s a n d / o r a n t e n n a e l o n g a t e
(fig. 1 4 3 - 1 4 4 ) - 66
66(65) C e l l c u p o p e n a n d c o n s p i c u o u s l y l o n g (fig. 1 4 1 , 1 4 2 ) . Mythicomyiidae
C e l l c u p different or absent. 67
 149 head Phorii
(Megaseliai

discal cell R4+5

145 habitus Conopidae (Dalmannia)

150 wing Stratiomyiic

(Oplodontha)

anal vein

146 wing Lonchopteridae

(Lonchoptera) (male)
parafaclalia

151 head
calyptr
fly

anal vein

147 wing Lonchopteridae

(Lonchoptera) (female)

3rd antennal segment

(1st flagellomere)

152 antenna calyptrate

153 wing Oestric

(Oestrus)

148 habitus Phoridae (spec

4 8
67(66) C e l l r4+5 n a r r o w i n g t o w a r d w i n g tip (fig. 1 4 3 , 1 4 5 ) o r c l o s e d (fig. 1 4 4 ) ; cell
c u p u s u a l l y e l o n g a t e , e n d i n g a c u t e l y , s o m e t i m e s s h o r t (fig. 1 4 5 ) . S e c o n d
a n t e n n a l s e g m e n t e l o n g a t e . M o u t h p a r t s u s u a l l y l o n g , slender, g e n i c u l a t e
at t h e b a s e , s o m e t i m e s a g a i n g e n i c u l a t e i n their m i d r e g i o n (fig. 1 4 3 , 1 4 5 ) .
If m o u t h p a r t s s h o r t , t h e n arista t e r m i n a l (fig. 1 4 4 ) . Conopidae
C o m b i n a t i o n o f characters n o t as g i v e n a b o v e . 68
68(67) W i n g l a n c e t - s h a p e d w i t h characteristic v e n a t i o n : v e i n R i s h o r t , cell r2+3
n a r r o w i n g t o w a r d w i n g tip, v e i n M f o r k e d i n t o M i a n d M 2 , c r o s s v e i n s
p r e s e n t o n l y i n basal p a r t o f t h e w i n g ; l o n g i t u d i n a l v e i n s w i t h b l a c k b r i s
tles o n dorsal side (fig. 1 4 6 , 1 4 7 ) . Lonchopteridae
W i n g different. 69
69(68) W i n g v e n a t i o n characteristic, r e d u c e d : t h e s t r o n g R - v e i n s e n d i n t h e
c o s t a , a b o u t h a l f w a y u p the w i n g ; t h e o t h e r v e i n s are w e a k e r a n d u s u a l l y
cross t h e w i n g i n a n o b l i q u e d i r e c t i o n , m o r e o r less parallel t o e a c h o t h e r
(fig. 1 4 8 ) . H e a d , p a l p a n d l e g s u s u a l l y w i t h s t r o n g , d e n t a t e d o r feathered
bristles (fig. 1 4 9 ) . S m a l l a n d h u m p - b a c k e d , u s u a l l y dark flies. Phoridae
C o m b i n a t i o n o f characters n o t as g i v e n a b o v e . 70
70(69) W i n g v e n a t i o n as i n fig. 150, w i t h a s m a l l , d i a m o n d - s h a p e d d i s c a l cell.
Legs w i t h o u t bristles; e m p o d i u m p u l v i l l i f o r m (fig. 103). Stratiomyidae
- C o m b i n a t i o n o f characters n o t as g i v e n a b o v e . 71
71(70) Ptilinal s u t u r e p r e s e n t (fig. 151; a u s u a l l y h o r s e s h o e s h a p e d g r o o v e o r
s u t u r e r u n n i n g a b o v e t h e l u n u l a d o w n o n either side o f t h e a n t e n n a e , s e p
arating the parafacialia f r o m t h e r e m a i n d e r o f t h e face). A r i s t a w i t h a f e w
e x c e p t i o n s s i t u a t e d o n dorsal side o f t h e 3rd a n t e n n a l s e g m e n t , near t h e
2nd s e g m e n t (fig. 152, h e a d figures 1 8 2 - 4 1 6 ) , arista rarely t e r m i n a l o r
absent. Acalyptrate and Calyptrate families - 81
- Ptilinal s u t u r e a b s e n t ( a l t h o u g h a c l o s e d s u t u r e is o f t e n a p p a r e n t a b o v e t h e
a n t e n n a e t h i s d o e s n o t e x t e n d b e l o w t h e a n t e n n a e , t h u s t h e r e are n o v e r
tical s u t u r e s b e t w e e n t h e e y e s b e l o w t h e a n t e n n a e ; fig. 1 5 4 , 1 5 5 ) . A r i s t a fre
q u e n t l y t e r m i n a l , s i t u a t e d at t h e e n d o f t h e 3rd a n t e n n a l s e g m e n t (fig. 1 5 4 ,
1 5 6 , 1 5 9 - 1 6 1 ) , s o m e t i m e s s i t u a t e d m o r e d o r s a l l y (fig. 155). 72

155 head Dolichopodidae

(Hercostomus)

49
yz(yi) O r i g i n o f v e i n R s b e f o r e or o p p o s i t e c r o s s v e i n H , o r at a d i s t a n c e b e y o n d
c r o s s v e i n H less t h a n t h e l e n g t h o f t h a t c r o s s v e i n (fig. 1 5 7 - 1 6 4 ) . 73
O r i g i n o f v e i n R s farther b e y o n d c r o s s v e i n H , at a d i s t a n c e greater t h a n t h e
l e n g t h o f t h e c r o s s v e i n (fig. 1 6 5 , 1 6 8 ) . 78
2
73(7 ) W i n g v e n a t i o n as i n fig. 157 or 158: v e i n Sc c o m p l e t e , e n d i n g i n t h e c o s t a ;
c r o s s v e i n D M - C u absent; v e i n s M i a n d M 2 l o n g ; cell c u p s m a l l , n a r r o w i n g
t o w a r d its a c u t e apical a n g l e ; i n m a l e : anal l o b e large a n d anal v e i n n o t
r e a c h i n g w i n g m a r g i n ; in f e m a l e : anal l o b e v i r t u a l l y a b s e n t and anal v e i n
reaching w i n g margin. Opetiidae
(In E u r o p e 1 w i d e s p r e a d s p e c i e s , Opetia nigra M e i g e n . )
W i n g v e n a t i o n different. 74
74(73) V e i n Sc c o m p l e t e , e n d i n g i n c o s t a ; anal v e i n r e a c h i n g w i n g m a r g i n ; cell
c u p e n d i n g i n an a c u t e a n g l e ; alula a n d w e l l d e v e l o p e d anal l o b e p r e s e n t
(fig. 1 5 9 , 1 6 0 ) . Platypezidae
> [ M o s t P l a t y p e z i d a e s h o w t h e f o l l o w i n g a d d i t i o n a l characters: t e r m i n a l
arista; v e i n M f o r k e d a n d c r o s s v e i n D M - C u p r e s e n t ; first tarsal s e g m e n t o f
h i n d l e g in t h e s u b f a m i l y C a l l o m y i i n a e l o n g a n d c y l i n d r i c a l , u s u a l l y
s w o l l e n i n m a l e ( a r r o w fig. 159), s l e n d e r i n f e m a l e ; t h e s u b s e q u e n t tarsal
s e g m e n t s c y l i n d r i c a l . In t h e s u b f a m i l y P l a t y p e z i n a e the first tarsal s e g
m e n t o f h i n d l e g s h o r t , laterally c o m p r e s s e d , s u b s e q u e n t tarsal s e g m e n t s
laterally c o m p r e s s e d as w e l l , in particular i n t h e f e m a l e ( a r r o w fig. 1 6 0 ) .
C r o s s v e i n D M - C u a b s e n t in Microsania.]
- C o m b i n a t i o n o f characters n o t as g i v e n a b o v e ( n o t at t h e s a m e t i m e t h e
f o l l o w i n g four characters: v e i n Sc r e a c h i n g w i n g m a r g i n , anal l o b e w e l l
d e v e l o p e d , cell c u p e n d i n g i n an a c u t e a n g l e , anal v e i n r e a c h i n g w i n g m a r
gin). 75
> [A k e y for t h e f a m i l i e s w h i c h k e y o u t i n c o u p l e t s 7 5 - 8 0 is g i v e n b y Sinclair
& C u m m i n g 2 0 0 6 , i n c l u d i n g u n p l a c e d g e n e r a and a s o m e w h a t different
concept o f the families Empididae and Dolichopodidae.]
75(74) C o s t a s u r r o u n d i n g the w i n g c o m p l e t e l y [not clear f r o m t h e d r a w i n g b u t it
s h o u l d b e f r o m the i n s e c t ] ; v e i n S c e n d i n g i n t h e c o s t a ; anal v e i n s h o r t or
absent; cell d m p r e s e n t ; 3 v e i n s o r i g i n a t i n g f r o m this cell r e a c h t h e w i n g
m a r g i n (fig. 161). Microphoridae
C o s t a e n d i n g near the w i n g tip (fig. 1 6 3 , 1 6 5 , 1 6 6 b ) ; v e i n Sc u s u a l l y e n d i n g
i n R i (fig. 162) or i n c o m p l e t e a n d n o t r e a c h i n g the c o s t a (fig. 1 6 5 , 1 6 6 b ) ;
anal v e i n and cell d m variable. 76
76(75) C e l l s br, b m a n d c u p s m a l l ; c r o s s v e i n B M - C u a b s e n t (cells b m a n d d m are
fused) (fig. 1 6 2 - 1 6 4 ) ; 2 v e i n s f r o m c r o s s v e i n D M - C u t o w a r d t h e w i n g m a r
g i n , t h e anterior o f w h i c h m a y b e c u r v e d (fig. 163) or f o r k e d i n t o M i a n d
M 2 (fig. 1 6 4 ) . B o d y u s u a l l y w i t h a g r e e n i s h m e t a l l i c lustre, i n a f e w s p e c i e s
d u l l , y e l l o w , b r o w n or b l a c k . Dolichopodidae

51
166a habitus Atelestidae
(Meghyperus) 170 front view thorax 171 front view thorax
Empididae (Empis) Hybotidae (Hybc

1st antennal segme

172 antenna Empididae (Emp

il cup ' .
B M C j anal vein cell dm

166b wing Atelestidae

co I bm
(Atelestus) cell cup
1st antennal segme

alula absent

173 antenna Hybotidae (Hybc

167 habitus
Hybotidae
stigma
(Anthalia)
mouthparts

eye
z^cell dm
cell bm
:
= * ^ c e l l cup

168 habitus Hybotidae

(Hybos)

174a ventral view head a

thorax Hippoboscid
(Melophagus)

169 habitus Hybotidae (Drapetis)

5^
 C e l l s b r a n d b m large; cell c u p v a r i a b l e b u t u s u a l l y large; c r o s s v e i n B M - C u
c e l
p r e s e n t (fig. 1668-167); l d m m a y b e p r e s e n t (fig. 166a) o r n o t (fig. 1 6 6 b ) .
C o m b i n a t i o n o f o t h e r characters n o t as g i v e n a b o v e . 77
W i n g w i t h alula; cell c u p large a n d a t l e a s t as l o n g as cells b r a n d b m , u s u
ally l o n g e r ; either cell d m p r e s e n t a n d v e i n M f o r k e d i n t o M i a n d M 2 (fig.
166a); o r cell d m a b s e n t a n d v e i n M n o t f o r k e d (fig. 1 6 6 b ) .
Atelestidae: Atelestinae
W i n g w i t h o u t alula; cell c u p u s u a l l y s h o r t e r t h a n cells b r a n d b m (fig. 167)
or cell c u p a b s e n t . In case o f cell c u p b e i n g as l o n g as o r l o n g e r t h a n cells b r
a n d b m , t h e n cell d m p r e s e n t a n d v e i n M n o t f o r k e d (fig. i 6 8 ) . H y b o t i d a e
E i t h e r a p r e c o x a l b r i d g e p r e s e n t ( p r o s t e r n a l sclerite o n t h e v e n t r a l side o f
t h e t h o r a x , b e t w e e n t h e n e c k a n d t h e fore l e g s , large a n d f u s e d w i t h t h e
lateral plates b o r d e r i n g it) (fig. 1 3 3 , 1 3 4 , 1 7 0 ) , a n d / o r t h e c o s t a s u r r o u n d i n g
all o f t h e w i n g . First a n t e n n a l s e g m e n t u s u a l l y d i s t i n c t l y b r i s t l e d , b u t i n
a n y case w i t h s o m e b r i s t l e s o n its dorsal a n d / o r v e n t r a l side (fig. 172).
Empididae
[Including the o n e brachystomatid g e n u s w i t h v e i n R4+5 n o t forked,
Heleodromia.]
N o p r e c o x a l b r i d g e p r e s e n t ( p r o s t e r n u m s m a l l , s e p a r a t e d f r o m lateral
plates b y a m e m b r a n e ) (fig. 171). C o s t a e n d s near t h e w i n g tip. First a n t e n
nal s e g m e n t w i t h o u t b r i s t l e s (fig. 173). 79
C r o s s v e i n s B M - C u a n d D M - C u a b s e n t ; cell c u p l o n g a n d a l m o s t r e c t a n g u
lar; anal v e i n a n d v e i n C u A i a r c h e d t o w i n g m a r g i n (fig. 1 6 5 ) .
Atelestidae: Nemedininae
(In C e n t r a l E u r o p e 1 s p e c i e s , Nemedina alamirabilis C h a n d l e r . )
C o m b i n a t i o n o f w i n g characters n o t as g i v e n a b o v e (fig. 1 6 6 - 1 6 9 ) . 80
W i n g w i t h alula; cell c u p large a n d at l e a s t as l o n g as cells b r a n d b m , u s u
ally l o n g e r ; e i t h e r cell d m p r e s e n t a n d v e i n M f o r k e d i n t o M i a n d M 2 (fig.
166a); or cell d m a b s e n t a n d v e i n M n o t f o r k e d (fig. 1 6 6 b ) .
Atelestidae: Atelestinae
W i n g w i t h o u t alula; cell c u p u s u a l l y s h o r t e r t h a n cells b r a n d b m (fig. 167)
or cell c u p a b s e n t (fig. 1 6 9 ) . In case o f cell c u p b e i n g as l o n g as o r l o n g e r
t h a n cells b r a n d b m , t h e n cell d m p r e s e n t a n d v e i n M n o t f o r k e d (fig. 168).
Hybotidae

'70 C o x a e o f m i d l e g s n o t far apart. Tarsal c l a w s n o r m a l , n o t s t r o n g l y c u r v e d .

Fly n o t e c t o p a r a s i t i c o n b i r d s o r m a m m a l s . 82
- C o x a e o f m i d legs, and u s u a l l y o f the other legs as w e l l , far apart (fig.
174a). Tarsal c l a w s strongly c u r v e d (fig. 174b) o r last tarsal s e g m e n t
w i d e n e d and e y e s reduced or absent (fig. 175). Fly l i v i n g as a n ectopar
asite o n birds or m a m m a l s . Calyptratae: H i p p o b o s c o i d e a - 1 6 9

175 head and fore

legs Streblidae
eyes rudimentary (Brachytarsina)
or absent

174b habitus Hippoboscidae 53

(Ornithomyia)
 one pair of Iv
F-bristles
'Ov

182 head
Lonchaeidae
(Dasiops)

176 thorax
calyptrate fly

antennal ,1st
seam \ _Xv- antennal
L

yy J segment

~" 2nd

177 antenna
calyptrate fly

178 head Muscidae (Lispe)

17 b
( \ Se R1 R-M c
183 habitus Lonchaeidae
I I J- i. / R2 3
+
(Setisquamalonchaea)

179 wing
Calliphoridae
(Calliphora)
BM-Cu absen

181 dorsal view

180 dorsal view
thorax
thorax
acalyptrate fly 184 habitus Sphaeroceridae
calyptrate fly
(Limosina)
54
G r e a t e r a m p u l l a p r e s e n t as a m a j o r b u l g e i m m e d i a t e l y b e l o w t h e anterior
p a r t o f t h e w i n g i n s e r t i o n (fig. 1 7 6 ) . S e c o n d a n t e n n a l s e g m e n t w i t h a d i s
t i n c t s e a m o n its u p p e r side (fig. 177). U s u a l l y 1 or m o r e s t r o n g v i b r i s s a e
p r e s e n t , as w e l l as i n c u r v e d l o w e r F - b r i s t l e s (fig. 178). L o w e r c a l y p t e r u s u
ally s t r o n g l y d e v e l o p e d a n d v e i n Sc m o r e or less c o m p l e t e , separate f r o m
v e i n R i (fig. 1 7 9 , 396). U p p e r side o f t h o r a x u s u a l l y w i t h a d i s t i n c t a n d
c o m p l e t e t r a n s v e r s e s u t u r e (fig. 1 7 6 , 1 8 0 ; n o t c o m p l e t e i n m o s t
S c a t h o p h a g i d a e a n d s o m e M u s c i d a e , A n t h o m y i i d a e or F a n n i i d a e ) ; p o s t a -
lar callus u s u a l l y d i s t i n c t l y s e p a r a t e d (fig. 180).
O t h e r C a l y p t r a t e families - 1 7 0
Greater ampulla inconspicuous, if h o w e v e r w e l l developed, then vibrissae
a b s e n t a n d o t h e r characters different ( s e c o n d a n t e n n a l s e g m e n t u s u a l l y
w i t h o u t a d i s t i n c t s e a m ; l o w e r c a l y p t e r s m a l l or a b s e n t ; central p a r t o f
t r a n s v e r s e s u t u r e u s u a l l y less d i s t i n c t or a b s e n t (fig. 181); p o s t a l a r c a l l u s
n o t d i s t i n c t l y s e p a r a t e d or a b s e n t (fig. 181)). A c a l y p t r a t e families - 83
H a l t e r e dark b r o w n t o b l a c k . H e a d large, w i d e a n d h i g h ( h e m i s p h e r i c a l ) ;
o n l y 1 pair o f F - b r i s t l e s , s i t u a t e d at t h e l e v e l o f t h e ocellar triangle a n d
c u r v i n g b a c k w a r d (fig. 182). H i n d m a r g i n o f a n e p i s t e r n u m w i t h a r o w o f
bristles. W i n g a l w a y s w i t h o u t m a r k i n g s . C e l l c w i d e ; v e i n Sc c o m p l e t e ,
e n d s i n t h e costa separate f r o m v e i n R i (fig. 183). Lonchaeidae
[ U s u a l l y c o m p a c t , 3 to 6 m m l o n g dark flies w i t h or w i t h o u t lustre a n d t h e
f o l l o w i n g characters: L u n u l a large; d i v e r g i n g P-bristles w e a k l y d e v e l o p e d .
C o s t a w i t h s u b c o s t a l b r e a k . T i b i a e u s u a l l y w i t h o u t dorsal preapical b r i s t l e ,
s o m e t i m e s present o n m i d leg only. A b d o m e n short, w i d e , compressed.
F e m a l e w i t h a l a n c e o l a t e , i n c o m p l e t e l y retractable o v i p o s i t o r . ]
H a l t e r e n o t dark b r o w n to b l a c k or o t h e r characters different. If haltere
dark b r o w n , t h e n h e a d n o t h e m i s p h e r i c a l a n d / o r m o r e t h a n 1 pair o f F -
bristles p r e s e n t a n d / o r a n e p i s t e r n u m w i t h o u t a r o w o f b r i s t l e s a l o n g its
hind margin, a n d / o r w i n g w i t h a pattern of markings. 84
V i b r i s s a e (fig. 1 8 4 , 1 8 5 , 1 8 9 ) or v i b r i s s a - l i k e bristles (fig. 186) p r e s e n t at o r
near t h e v i b r i s s a l a n g l e . 85
N o v i b r i s s a e or v i b r i s s a - l i k e b r i s t l e s (fig. 1 8 7 , 1 8 8 ) . 134
First tarsal s e g m e n t o f h i n d l e g m u c h s w o l l e n , a n d u s u a l l y s h o r t e r t h a n t h e
s e c o n d s e g m e n t (fig. 1 8 4 , 1 8 5 ) . T h i r d a n t e n n a l s e g m e n t s h o r t , arista l o n g .
W i n g v e n a t i o n r a n g i n g f r o m s o m e w h a t r e d u c e d (fig. 1 8 4 : c r o s s v e i n B M -
C u a b s e n t , cell c u p o p e n , v e i n s b e y o n d c r o s s v e i n D M - C u l e s s d i s t i n c t or
l a r g e l y absent) t o m o r e or less c o m p l e t e (fig. 185). Sphaeroceridae
First tarsal s e g m e n t o f h i n d l e g n o t s w o l l e n , u s u a l l y l o n g a n d slender.
O t h e r characters v a r i a b l e . 86

/
/
>M1

S DM-Cu

*BM-Cu
""cell cup

preapical
bristle
"\
1 st tarsal
segment

185 habitus
56
86(8 )
5
D o r s a l p r e a p i c a l bristle at l e a s t p r e s e n t o n tibia o f m i d l e g , b u t u s u a l l y o n
all tibiae (fig. 1 9 0 , 1 9 3 , 1 9 8 , 201, 204). 87
A l l tibiae w i t h o u t dorsal preapical bristle. 101
87(86) V e i n S c c o m p l e t e , r e a c h i n g t h e c o s t a separate f r o m v e i n R i (fig. 1 9 0 , 1 9 2 ,
193). 88
V e i n S c i n c o m p l e t e , d o e s n o t r e a c h t h e costa separately: S c r e d u c e d o r
s h o r t e n e d (fig. 191), o r m e r g i n g w i t h v e i n R i b e f o r e t h e c o s t a (fig. 215b).
94
88(87) S e c o n d a n t e n n a l s e g m e n t triangular a t o u t e r side; t h i r d a n t e n n a l s e g m e n t
s h o r t ; P-bristles d i v e r g i n g o r a b s e n t ; a s i n g l e p a i r o f interfrontal bristles,
c u r v i n g f o r w a r d (fig. 189). C o s t a w i t h a s u b c o s t a l b r e a k . V e i n Sc c o m p l e t e
a n d separate f r o m v e i n R i (fig. 1 9 0 ) . S l e n d e r , d a r k - b r o w n , dark b r o w n i s h -
y e l l o w o r y e l l o w flies. Clusiidae
S e c o n d a n t e n n a l s e g m e n t n o t t r i a n g u l a r at o u t e r side. O t h e r c h a r a c t e r s as
s p e c i f i e d a b o v e or n o t , b u t rarely i n t h e a b o v e c o m b i n a t i o n . 89
89(88) V e i n s Sc a n d R i d i s t i n c t l y separate a n d r e a c h i n g t h e c o s t a at a c o n s i d e r a b l e
d i s t a n c e f r o m e a c h o t h e r (fig. 1 9 3 , 1 9 8 , 201, 203). 90
V e i n s Sc a n d R i are parallel a n d r e a c h t h e c o s t a c l o s e t o g e t h e r (fig. 192). 93
90(89) A n a l v e i n s h o r t , n o t r e a c h i n g t h e w i n g m a r g i n a n d at t h e s a m e t i m e c o s t a
w i t h o u t s t r o n g b r i s t l e s (fig. 1 9 3 , 1 9 8 ) . 91
A n a l v e i n l o n g , r e a c h i n g t h e w i n g m a r g i n o r e n d i n g j u s t s h o r t o f i t (fig.
201, 203); costa o f t e n w i t h s t r o n g bristles (fig. 201); i f anal v e i n n o t c o n
spicuously long, then costa w i t h strong bristles. 92
91(90) P-bristles d i v e r g i n g o r a b s e n t ; arista bare (fig. 1 9 4 , 1 9 5 ) . P o s t e r i o r spiracle
w i t h 1 o r m o r e bristles at t h e p o s t e r o v e n t r a l m a r g i n (fig. 1 9 6 , 1 9 7 ) .
Sepsidae
[ U s u a l l y g l o s s y - b l a c k a n t - l i k e flies w i t h r e l a t i v e l y f e w b r i s t l e s o r p u b e s
cence, a round head and a basally constricted a b d o m e n ; w i n g often w i t h a
dark m a r k i n g at t h e w i n g tip (Sepsis, fig. 198); i n t h e m a l e , f e m u r a n d tibia
o f fore l e g s h o w i n g p r o j e c t i o n s , s p i n e s , t e e t h , e t c . (fig. 2 4 4 ) . E x c e p t i o n :
Orygma: b r o w n , s o m e w h a t flattened, r o b u s t f l y p r o v i d e d w i t h b r i s t l e s ,
fore l e g s i m p l e , h e a d as i n fig. 1 9 4 , 1 9 9 , i n coastal habitats.]
P-bristles a l m o s t a l w a y s c o n v e r g i n g (fig. 2 0 0 ) , v e r y rarely parallel.
Posterior spiracle w i t h o u t b r i s t l e s at t h e p o s t e r o v e n t r a l m a r g i n .
Lauxaniidae
[Small t o m e d i u m - s i z e d (2.5 t o 6 m m ) r o b u s t flies, n o t a n t - l i k e ; t h i r d
a n t e n n a l s e g m e n t c a n b e e l o n g a t e (fig. 1 8 7 , 1 9 3 ) ; arista bare t o l o n g p u b e s
c e n t ; w i n g u s u a l l y clear, i n a n u m b e r o f s p e c i e s w i t h d a r k e n i n g a l o n g s i d e
t h e v e i n s o r w i t h m a r k i n g s all o v e r (fig. 303).]

haltere,-
posterior
spiracle J[

abdomen

196 posterior spiracle

Sepsidae (Orygma)

197 posterior spiracle

Sepsidae (Nemopoda)
57
201 habitus Heleomyzidae (Orbellia) 205 head Trixoscelididae
o (Trixoscelis)
 92(90) C o s t a w i t h b o t h larger a n d s m a l l e r b r i s t l e s (fig. 201). H e a d n o t as i n fig.
202a ( l o w e r p a r t o f face n o t s t r o n g l y p r o t r u d i n g ) . S c u t u m n o t d o r s o v e n -
trally flattened (fig. 201). F i f t h tarsal s e g m e n t n o t e n l a r g e d a n d triangular.
Heleomyzidae
C o s t a w i t h s m a l l e r b r i s t l e s o n l y (fig. 203). H e a d w i t h l o w e r p a r t o f face
s t r o n g l y p r o t r u d i n g b e l o w circular t h i r d a n t e n n a l s e g m e n t (fig. 202a).
Fifth tarsal s e g m e n t large, triangular, w i d e r t h a n o t h e r tarsal s e g m e n t s ;
first tarsal s e g m e n t o f m i d a n d h i n d l e g u s u a l l y w i t h s p i n e s o n v e n t r a l side
(fig. 202b). In Coelopa s c u t u m s o m e w h a t flattened a n d l e g s d e n s e l y s e t
w i t h fine b r i s t l e s . [Flies o f c o a s t a l habitats.] Coelopidae
93(89) O c - b r i s t l e s i n s e r t e d o n ocellar t r i a n g l e ; arista l o n g p l u m o s e ; plates o n
w h i c h t h e F - b r i s t l e s are i n s e r t e d are separate f r o m t h e m a r g i n s o f t h e e y e s ;
t h e l o w e r m o s t F - b r i s t l e s c u r v i n g f o r w a r d (fig. 2 0 4 ) . C r o s s v e i n B M - C u
a b s e n t (cells b m a n d d m f u s e d ) . Curtonotidae
(In E u r o p e 1 w i d e s p r e a d s p e c i e s , Curtonotum anus ( M e i g e n ) . )
- O c - b r i s t l e s i n s e r t e d n e x t t o t h e ocellar triangle at t h e s a m e l e v e l as t h e
anterior o c e l l u s o r l o w e r ; arista u s u a l l y s h o r t p u b e s c e n t , rarely s h o r t
p l u m o s e ; plates o n w h i c h t h e F - b r i s t l e s are i n s e r t e d b o r d e r i n g t h e m a r
g i n s o f the e y e s ; F - b r i s t l e s c u r v i n g b a c k w a r d (fig. 205). C r o s s v e i n B M - C u
p r e s e n t (cells b m a n d d m separate) (fig. 206). Trixoscelididae
(In E u r o p e 1 g e n u s , Trixoscelis, w i t h a b o u t 25 species.)
94(87) C e l l c u p o p e n ; c r o s s v e i n B M - C u a b s e n t (cells b m a n d d m f u s e d ) ; anal v e i n
a b s e n t (fig. 207). A t m o s t tibia o f t h e m i d l e g w i t h dorsal p r e a p i c a l b r i s t l e .
95
- C e l l c u p c l o s e d ; v e i n B M - C u p r e s e n t (cells b m a n d d m separate, fig. 208),
or v e i n B M - C u a b s e n t (cells b m a n d d m f u s e d , fig. 219). A n a l v e i n u s u a l l y
present (short in Diastatidae a n d C a m p i c h o e t i d a e , s o m e t i m e s absent in
D r o s o p h i l i d a e ) . U s u a l l y all tibiae w i t h dorsal p r e a p i c a l bristle (fig. 215a,
222), t h o u g h s o m e t i m e s t h i n a n d d e l i c a t e i n O d i n i i d a e a n d
Chiropteromyzidae. 96

cell bm J

cell dm

B M
" C u
cell dm
208 wing Campichoetidae
206 wing Trixoscelididae (Trixoscelis) (Euthychaeta)

210 femur
inai vein absent ; fore leg
BM-Cu absent T.. Camillidae
cell dm (Camilla)
-spine
207 wing Camillidae 209 head Camillidae
(Camilla) (Camilla)

59
211 habitus Ephydridae
(Coenia)

215a habitus
Cnemospathidae
(Prosopantrum)

215b detail wing base 7r.,\\

Cnemospathidae ci,i'\ v ,H
212 head Ephydridae transverse
(Prosopantrum) suture
(Limnellia)

215c thorax
Cnemospathidae
(Prosopantrum)

213 head Ephydridae

(Thinoscatella)

214 head Odiniidae (Odinia)

6 o
216b habitus Chiropteromyzidae
(Chiropteromyza)
 95(94) Arista p l u m o s e w i t h l o n g rays a b o v e and v e r y short rays b e l o w ; P-bristles
c o n v e r g i n g (fig. 209). F e m u r o f fore l e g w i t h a s p i n e o n its v e n t r a l s i d e ,
b e y o n d t h e m i d d l e (fig. 210). W i n g v e n a t i o n as i n fig. 207. Camillidae
(In E u r o p e 1 g e n u s , Camilla, w i t h a b o u t 8 s p e c i e s . )
A r i s t a u s u a l l y w i t h l o n g r a y s , l i m i t e d to t h e u p p e r side (fig. 211), arista
s o m e t i m e s bare (fig. 212) or s h o r t p u b e s c e n t ; n o real P - b r i s t l e s , b u t s o m e
t i m e s d i v e r g i n g p s e u d o p o s t o c e l l a r b r i s t l e s are p r e s e n t (fig. 213); face u s u
ally s w o l l e n , c o n v e x (fig. 211, 212). F e m u r o f fore l e g w i t h or w i t h o u t v e n
tral s p i n e s . Ephydridae
96(94) 3 pairs o f F-bristles p r e s e n t ; t h e l o w e s t pair c u r v i n g i n w a r d , the u p p e r a n d
m i d d l e pairs c u r v i n g b a c k w a r d ; P - b r i s t l e s d i v e r g i n g (fig. 214). L e g s o f t e n
w i t h alternating b r o w n i s h and y e l l o w i s h bands. Odiniidae

2 or 3 pairs o f F - b r i s t l e s p r e s e n t : 1 pair c u r v i n g f o r w a r d , o b l i q u e l y f o r w a r d
or o u t w a r d , 1 or 2 pair(s) c u r v i n g b a c k w a r d ; P-bristles c o n v e r g i n g , s o m e
times a b s e n t (fig. 215-218, 220-224). 97
97(96) 2 pairs o f F-bristles p r e s e n t , t h e l o w e r pair c u r v i n g o u t w a r d (fig. 216a);
arista bare t o p u b e s c e n t (fig. 215a, 2 1 6 b ) . 98
F-bristles n o t c u r v i n g o u t w a r d (fig. 217, 218, 220-224); arista u s u a l l y
p l u m o s e (fig. 217, 218) b u t in s o m e c a s e s p u b e s c e n t (fig. 223-224). 99

98(97) T h o r a x w i t h 3 pairs o f d o r s o c e n t r a l b r i s t l e s o f w h i c h t h e first pair is l o c a t

e d near or j u s t b e f o r e t h e t r a n s v e r s e s u t u r e (fig. 215c: d c ) ; k a t e p i s t e r n u m
w i t h 2 s t r o n g b r i s t l e s (fig. 215a). A l l tibiae w i t h s t r o n g dorsal p r e a p i c a l b r i s
tle. Cnemospathidae
(In E u r o p e 1 s p e c i e s , Prosopantrumflavifrons (Tonnoir & Malloch).)
T h o r a x w i t h 4 pairs o f d o r s o c e n t r a l b r i s t l e s , o f w h i c h 1 pair i n front, a n d 3
pairs b e h i n d t h e t r a n s v e r s e s u t u r e ; k a t e p i s t e r n u m w i t h 1 s t r o n g b r i s t l e s
w i t h a f e w w e a k , s m a l l b r i s t l e s i n f r o n t (fig. 216b). D o r s a l p r e a p i c a l bristle
p r e s e n t , s o m e t i m e s s m a l l e r o n tibiae o f fore a n d h i n d l e g .
Chiropteromyzidae

218 head Drosophilidae

(Chymomyza)

anepisternum

219 wing Drosophilidae (Drosophila)

17 habitus Drosophilidae 6 1
(Drosophila)
 224 head Campichoetidae
(Campichoeta)

preapical
bristle

222 habitus Diastatidae

(Diastata)

229 head Asteiidae

(Asteia)
 99(97) T h e F-bristle t h a t c u r v e s f o r w a r d or o b l i q u e l y f o r w a r d is i n s e r t e d o n the
s a m e l e v e l or further a w a y f r o m t h e e y e m a r g i n t h a n t h e n e a r e s t bristle
t h a t c u r v e s b a c k w a r d (fig. 218, 220, 271); t h i s n e a r e s t b a c k w a r d d i r e c t e d
bristle c a n b e v e r y s m a l l (fig. 220b). A n e p i s t e r n u m bare (fig. 217). W i n g
v e n a t i o n as in fig. 219; c r o s s v e i n B M - C u u s u a l l y a b s e n t b u t s o m e t i m e s
present. Drosophilidae
T h e F-bristle t h a t c u r v e s f o r w a r d is i n s e r t e d c l o s e r to t h e e y e m a r g i n t h a n
the n e a r e s t bristle t h a t c u r v e s b a c k w a r d (fig. 221-224). A n e p i s t e r n u m bare
or w i t h b r i s t l e s (fig. 222). 100

ioo(99) A r i s t a s h o r t t o m o d e r a t e l y l o n g p l u m o s e ; 1 s t r o n g F - b r i s t l e c u r v i n g for
w a r d , 1 s t r o n g F-bristle c u r v i n g b a c k w a r d (fig. 221). A n e p i s t e r n u m w i t h
s h o r t bristles o n u p p e r a n d p o s t e r i o r part, s t r o n g e s t n e a r h i n d m a r g i n .
C o s t a w i t h b o t h h u m e r a l a n d s u b c o s t a l b r e a k s (fig. 222). Diastatidae
(In E u r o p e 1 g e n u s , Diastata, w i t h a b o u t 9 species.)
A r i s t a p u b e s c e n t (fig. 223, 224); e i t h e r 1 s t r o n g F - b r i s t l e c u r v i n g f o r w a r d
a n d 1 s t r o n g F-bristle c u r v i n g b a c k w a r d (fig. 223) o r w i t h y e t a n o t h e r ,
w e a k e r bristle c u r v i n g b a c k w a r d (fig. 224). A n e p i s t e r n u m bare. C o s t a
w i t h s u b c o s t a l b r e a k o n l y (fig. 225). Campichoetidae
101(86) V e i n R2+3 e n d s in t h e c o s t a a b o u t h a l f w a y u p t h e w i n g ; v e i n M i a p i c a l l y
pallid, n o t r e a c h i n g t h e w i n g m a r g i n (fig. 227). Xenasteiidae
> [Very s m a l l (1.3 t o 1.7 m m ) dark c o l o u r e d flies w i t h a p u b e s c e n t arista (fig.
226); c r o s s v e i n s B M - C u a n d D M - C u a b s e n t ; cell c u p c l o s e d ; anal v e i n
a b s e n t ; alula w i t h l o n g hairs.]
[ N a n n o d a s t i i d a e c o u l d k e y o u t h e r e i f t h e anterior w e a k b r i s t l e s a l o n g t h e
l o w e r m a r g i n o f t h e g e n a e h a v e b e e n i n t e r p r e t e d as v i b r i s s a e .
D i s t i n g u i s h i n g c h a r a c t e r s are: all F - b r i s t l e s c u r v i n g b a c k w a r d (in
X e n a s t e i i d a e l o w e r pair c u r v i n g i n w a r d ) ; c r o s s v e i n D M - C u s o m e t i m e s
p r e s e n t (fig. 363), alula a b s e n t (fig. 3 6 4 ) ; see also c o u p l e t 167 a n d t h e f a m i
l y descriptions.]
V e i n R2+3 is s t r i k i n g l y s h o r t (fig. 228) or e n d s in t h e c o s t a d i s t i n c t l y
b e y o n d t h e m i d d l e (fig. 230, 2 3 4 , 235), or i f v e i n R2+3 e n d s in t h e c o s t a
a b o u t h a l f w a y u p the w i n g , t h e n v e i n M i r e a c h i n g t h e w i n g m a r g i n (fig.
240). 102
102(101) W i n g b a s e n a r r o w , alula a n d anal v e i n s m a l l o r a b s e n t , cell c u p s o m e t i m e s
a b s e n t as w e l l (fig. 228, 230, 234-237); v e i n Sc i n c o m p l e t e , s h o r t e n e d or
m e r g i n g w i t h v e i n R i at s o m e d i s t a n c e f r o m t h e c o s t a (in Geomyza, fig.
236, apical p a r t s o m e t i m e s v i s i b l e as a t h i n l i n e r e a c h i n g t h e c o s t a ) . P - b r i s -
tles s m a l l , h a i r - l i k e or a b s e n t (fig. 229, 232). 103
W i n g b a s e n o t n a r r o w , alula w e l l d e v e l o p e d . O t h e r c h a r a c t e r s as s p e c i f i e d
a b o v e or n o t , b u t rarely i n the a b o v e c o m b i n a t i o n . 106
103(102) C e l l r4+5 n a r r o w i n g b e c a u s e v e i n s R 4 + 5 a n d M i c o n v e r g i n g t o w a r d t h e
w i n g tip; c r o s s v e i n B M - C u a b s e n t (cells b m a n d d m f u s e d ) ; cell c u p o p e n
or a b s e n t (fig. 228, 230). O c - b r i s t l e s p r e s e n t or a b s e n t . 104

alula
absent

230 wing Stenomicridae

(Stenomicra) (Stenomicra) 63
 pseudo- Iv
P-bristle-

R4+5

^ffiwn^ cell dm
M
237 habitus Opomyzidae (Geomyza) 242 wing Agromyzidae (Melanagromyza)

64
- Cell r4+5 n o t n a r r o w i n g ; crossvein B M - C u present (cells b m and d m separate);
cell c u p closed (fig. 234,236), rarely o p e n (fig. 235). Oc-bristles present. 105
104(103) V e i n R2+3 c o n s p i c u o u s l y s h o r t , e n d i n g i n t h e basal h a l f o f t h e w i n g ;
c r o s s v e i n D M - C u a b s e n t (fig. 228). A r i s t a s o m e t i m e s bare or p u b e s c e n t
b u t u s u a l l y w i t h a characteristic z i g - z a g o f l o n g e r o r s h o r t e r dorsal a n d
v e n t r a l r a y s (fig. 229) or, i n o n e s p e c i e s , arista absent; ocellar triangle s i t u
ated at t h e n o r m a l p o s i t i o n , i n t h e u p p e r p a r t o f t h e frons. Asteiidae
- V e i n R2+3 n o t s t r i k i n g l y s h o r t ; c r o s s v e i n D M - C u a b s e n t (fig. 230) o r p r e s
e n t . A r i s t a w i t h l o n g r a y s (fig. 231); ocellar triangle s i t u a t e d m o r e o r l e s s i n
t h e m i d d l e o f the frons. Stenomicridae
(In E u r o p e 1 g e n u s , Stenomicra, w i t h 3 species.)
105(103) 2 t o 3 pairs o f F - b r i s t l e s , o f t e n w i t h 1 o r m o r e w e a k e r F - b r i s t l e s i n front. P -
bristles s m a l l , c o n v e r g i n g , rarely a b s e n t (fig. 2 3 2 , 2 6 9 ) . F e m u r o f fore l e g i n
several g e n e r a w i t h a s h o r t s p i n e o n t h e v e n t r a l side, b e y o n d t h e m i d d l e
(fig. 233). W i n g v e n a t i o n as i n fig. 2 3 4 , 235; cell c u p s o m e t i m e s o p e n ,
crossvein D M - C u s o m e t i m e s absent. Anthomyzidae
- O n l y 1 pair o f F - b r i s t l e s p r e s e n t (fig. 237); P-bristles a b s e n t . F e m u r o f fore
l e g w i t h o u t s h o r t s p i n e o n v e n t r a l side. W i n g v e n a t i o n as i n fig. 236, 237.
O p o m y z i d a e : Geomyza
106(102) G e n a w i t h 1 o r m o r e s t r o n g u p c u r v e d bristle(s); t h e 2 t o 5 F - b r i s t l e s c u r v
i n g o u t w a r d o v e r t h e m a r g i n o f t h e e y e ; n o real P-bristles, b u t o f t e n
replaced b y d i v e r g i n g p s e u d o p o s t o c e l l a r b r i s t l e s (fig. 238a). V e i n S c paral
lel t o , a n d j u s t b e f o r e t h e c o s t a , t o u c h i n g o r m e r g i n g w i t h v e i n R i ;
c r o s s v e i n B M - C u p r e s e n t (cells b m a n d d m separate); cell c u p c l o s e d (fig.
238b). [Flies o f coastal habitats.] Canacidae
G e n a w i t h o u t s t r o n g u p c u r v e d bristles. C o m b i n a t i o n o f o t h e r characters
n o t as g i v e n a b o v e . 107
> [ O f t h e f a m i l i e s k e y i n g o u t h e r e (vibrissae p r e s e n t , preapical b r i s t l e s
absent) o u t w a r d c u r v i n g F - b r i s t l e s c a n b e f o u n d i n : P i o p h i l i d a e a n d
H e l e o m y z i d a e (Oldenbergiella a n d t h e B o r b o r o p s i n a e ) (anal v e i n l o n g ,
u s u a l l y r e a c h i n g t h e w i n g m a r g i n , fig. 2 5 4 ) , T e t h i n i d a e ( l u s t r o u s b u l g e
b e t w e e n antenna and vibrissa-like bristle, fig. 255), Carnidae a n d
M i l i c h i i d a e ( l o w e r F - b r i s t l e s c u r v i n g i n w a r d , fig. 2728-274) a n d
C h l o r o p i d a e ( c r o s s v e i n B M - C u a b s e n t ; cell c u p o p e n o r a b s e n t , fig. 240).]
0
107(106) V e i n S c a b r u p t l y b e n t f o r w a r d t o w a r d t h e c o s t a at n e a r l y 9 0 ; cell c u p
c l o s e d b y a g e n i c u l a t e v e i n Q 1 A 2 a n d w i t h a n a c u t e apical e n d (fig. 239a).
Tephritidae
> [ T e p h r i t i d a e are w i t h o u t t r u e v i b r i s s a e b u t s o m e g e n e r a ( e . g .
Chaetostomella, Chetostomd) h a v e s t r o n g bristles near t h e vibrissal angle
(fig. 239b); their w i n g p a t t e r n is u s u a l l y as i n fig. 239a. S e e also c o u p l e t 150.]
V e i n Sc a n d cell c u p n o t as g i v e n a b o v e . 108
108(107) V e i n Sc c o m p l e t e , u s u a l l y r e a c h i n g t h e c o s t a separate f r o m v e i n R i ( w i n g
figures 243-2578), s o m e t i m e s Sc a n d R i e n d i n g c l o s e t o g e t h e r (fig. 256), o r
v e i n Sc m e r g e s w i t h R i j u s t b e f o r e t h e c o s t a (e.g. A g r o m y z i n a e , fig. 2 4 2 ;
T e t h i n i d a e , fig. 251; C h a m a e m y i i d a e , fig. 257a). C e l l c u p c l o s e d . 109
- V e i n Sc absent, or short and n o t reaching the costa, or m e r g i n g w i t h v e i n
R i d i s t i n c t l y b e f o r e t h e c o s t a (fig. 2 4 0 , w i n g figures 258-262, 267-298), o r
r e d u c e d t o a fold t h a t m a y o r m a y n o t r e a c h t h e c o s t a ( P h y t o m y z i n a e , fig.
263, 264). C e l l c u p c l o s e d , o p e n o r a b s e n t . 120
> [In m o s t o f t h e s p e c i e s b e l o n g i n g t o t h e 21 f a m i l i e s k e y i n g o u t h e r e ( v i b r i s -

65
 sae a n d alula p r e s e n t , preapicals a b s e n t ) , v e i n Sc c o m p l e t e o r i n c o m p l e t e is
clearly v i s i b l e . B u t t h e r e are s o m e b o r d e r cases a n d therefore s o m e f a m i l i e s
h a v e b e e n k e y e d o u t t w i c e . N e v e r t h e l e s s , i n case o f d o u b t it is r e c o m
m e n d e d to follow b o t h alternatives and to consult the family descriptions
before c o m i n g t o a decision.]
109(108) P-bristles d i v e r g i n g (fig. 2 4 1 , 2 4 5 ) , parallel, r e d u c e d o r a b s e n t , i n t h e latter
case v e i n s S c a n d R i e n d i n t h e c o s t a at a c o n s i d e r a b l e d i s t a n c e f r o m e a c h
o t h e r (fig. 2 4 3 , 2 4 4 ) . 110
P-bristles c o n v e r g i n g o r a b s e n t , i n t h e latter case v e i n s S c a n d R i m e r g e
o v e r a s h o r t d i s t a n c e ( ( f i g . 250, 257a), h a v e a j o i n t e n d i n g i n t h e c o s t a (fig.
251), o r e n d i n t h e c o s t a c l o s e t o e a c h o t h e r (fig. 256). 115
110(109) V e i n Sc m e r g i n g w i t h R i j u s t b e f o r e t h e c o s t a ; v e i n R i c u r v i n g t o w a r d t h e
costa f r o m t h i s p o i n t ; c o s t a w i t h s u b c o s t a l b r e a k a n d w i t h a d i s t i n c t , d e e p
n o t c h (fig. 242). U s u a l l y , at l e a s t t h e l o w e r 2 pairs o f t h e F - b r i s t l e s c u r v e d
i n w a r d ; i n t e r f r o n t a l b r i s t l e s a b s e n t (fig. 2 4 1 ) . F e m a l e w i t h a n o n -
retractable o v i p o s i t o r , v i s i b l e as a c o n s p i c u o u s c o n i c a l s t r u c t u r e (fig. 266).
Agromyzidae: Agromyzinae
> [A d e e p n o t c h i n t h e c o s t a is a l s o f o u n d i n s o m e s p e c i e s o f M i l i c h i i d a e a n d
C a r n i d a e ; i n b o t h f a m i l i e s t h e i n t e r f r o n t a l b r i s t l e s are p r e s e n t (fig. 273,
V 4 . 287).]
V e i n s Sc a n d R i different; c o s t a w i t h o r w i t h o u t s u b c o s t a l b r e a k b u t i n a n y
case w i t h o u t a d i s t i n c t n o t c h t h e r e (fig. 2 4 3 , 2 4 4 ) . F - b r i s t l e s a n d inter
frontal b r i s t l e s v a r i a b l e , u s u a l l y n o t s i m i l a r t o w h a t is d e s c r i b e d a b o v e (fig.
243,244)- i "
111(110) V e i n s Sc a n d R i e n d i n t h e c o s t a at a c o n s i d e r a b l e d i s t a n c e f r o m e a c h other,
t h i s d i s t a n c e b e i n g at least o n e - t h i r d o f t h e d i s t a n c e b e t w e e n c r o s s v e i n H
a n d t h e e n d o f v e i n S c (fig. 2 4 3 , 2 4 4 ) . 112
- V e i n s S c a n d R i e n d i n t h e c o s t a n o t far apart, at a d i s t a n c e w h i c h is l e s s
than one-fourth o f the distance b e t w e e n crossvein H and the end o f v e i n
S c (fig. 2 4 6 , 2 4 8 ) . 114
112(111) W i n g d a r k e n e d w i t h n u m e r o u s s m a l l s p o t s (fig. 3 0 6 ) , o r w i t h a d i s t i n c t
w i n g pattern. Platystomatidae
- W i n g clear o r i n f u s c a t e d , b u t w i t h o u t a p a t t e r n o f m a n y s p o t s o r m a r k
i n g s , at m o s t w i t h a dark s p o t at t h e w i n g tip. 113
113(112) 3 pairs o f d o r s o c e n t r a l b r i s t l e s ; k a t e p i s t e r n u m w i t h a s t r o n g bristle (fig.
243). C o s t a w i t h a h u m e r a l b r e a k (fig. 243). P o s t e r i o r spiracle w i t h o u t b r i s
tles at t h e p o s t e r o v e n t r a l m a r g i n . Acartophthalmidae
(In E u r o p e 1 g e n u s , Acartophthalmus, w i t h 3 species.)
1 o r 2 pair(s) o f d o r s o c e n t r a l bristles; k a t e p i s t e r n u m w i t h o u t a bristle (fig.
2 4 4 ) . C o s t a c o n t i n u o u s . P o s t e r i o r spiracle w i t h 1 o r m o r e b r i s t l e s at t h e
p o s t e r o v e n t r a l m a r g i n (fig. 1 9 6 , 1 9 7 ) . Sepsidae
114(111) T h e l o w e r m o s t o f the 3 pairs o f F - b r i s t l e s c u r v i n g i n w a r d o r f o r w a r d , t h e
o t h e r F-bristles c u r v i n g b a c k w a r d ; v i b r i s s a l a n g l e w i t h 1 s t r o n g v i b r i s s a
(fig. 2 4 5 ) . A n a l v e i n n o t r e a c h i n g t h e w i n g m a r g i n (fig. 2 4 6 ) . Clusiidae
F - b r i s t l e s a b s e n t , o r p r e s e n t i n 1-4 pairs, u s u a l l y c u r v i n g o u t w a r d o r b a c k
w a r d , i f t h e l o w e r 1 o r 2 pair(s) c u r v i n g i n w a r d , t h e n t h e o t h e r F - b r i s t l e s
c u r v i n g o u t w a r d a n d / o r v i b r i s s a l a n g l e w i t h 2 s t r o n g v i b r i s s a e (fig. 2 4 7 ) .
A n a l v e i n l o n g (fig. 2 4 8 ) , u s u a l l y r e a c h i n g t h e w i n g m a r g i n . P i o p h i l i d a e
> [Pallopteridae c o u l d k e y o u t h e r e i f t h e anterior s e t u l a e a l o n g t h e l o w e r

67
251 wing Tethinidae
(Pelomyiinae)

255 head Tethinidae

(Tethininae)

BM-Cu
256 wing Chyromyidae
(Chyromya)

V J \ -
257a wing Chamaemyiidae
(Parochthiphila)
P-bristles absent

252 head Tethinidae (Pelomyiinae)

Oc-bristles absent

F-bristles absent

257b head
Chamaemyiidae
(Leucopis)

253 head and thorax BM-Cu and

Heleomyzidae Ov
cell cup absent

(Borboropsinae)
(Borboropsis) 258 wing Asteiidae (Leiomyza)
it " / x

A \. / % . f \ 259 head Periscelididae

anal vein , ( $\ celiS)
( P e r i s

BM-Cu c e || d m

jf 254 habitus Heleomyzidae

(Borboropsinae) (Nidomyia) 260 wing Periscelididae (Periscelis)

6 8
 m a r g i n o f t h e g e n a e h a v e b e e n i n t e r p r e t e d as v i b r i s s a e . D i s t i n g u i s h i n g
characters are t h a t P i o p h i l i d a e h a v e s t r o n g l y d e v e l o p e d v i b r i s s a e a n d t h e
c l y p e u s s m a l l a n d w i t h d r a w n , i n P a l l o p t e r i d a e t h e c l y p e u s is e x p o s e d (fig.
3ii).]
115(109) 2 pairs o f F - b r i s t l e s , l o w e r pair c u r v i n g f o r w a r d a n d s o m e w h a t i n w a r d ,
u p p e r pair c u r v i n g b a c k w a r d , l o w e r pair i n s e r t e d c l o s e r to e y e m a r g i n
t h a n u p p e r pair. O c - b r i s t l e s r e d u c e d or a b s e n t (fig. 249). V e i n s Sc a n d R i
r e a c h i n g the c o s t a separately, b u t j u s t b e f o r e t h e y m e r g e for a s h o r t d i s
t a n c e or t o u c h e a c h o t h e r ; c r o s s v e i n B M - C u a b s e n t (cells b m a n d d m
fused) (fig. 250). Aulacigastridae
(In E u r o p e 1 g e n u s , Aulacigaster, w i t h 4 s p e c i e s . )
F-bristles a r r a n g e d differently o r absent. O t h e r c h a r a c t e r s as s p e c i f i e d
a b o v e or n o t , b u t n o t i n t h e a b o v e c o m b i n a t i o n . 116
116(115) C r o s s v e i n B M - C u a b s e n t (cells b m a n d d m f u s e d ) (fig. 251) or v e r y w e a k l y
i n d i c a t e d ; v e i n Sc m e r g i n g w i t h R i j u s t b e f o r e or at t h e c o s t a . 1 o r 2 pair(s)
o f F - b r i s t l e s , c u r v i n g b a c k w a r d (fig. 252). Tethinidae: Pelomyiinae
- C r o s s v e i n B M - C u distinctly present (cells b m and d m separate) (fig. 254,256,
257). F-bristles u s u a l l y different, o n l y i n a f e w species as g i v e n a b o v e . 117
117(116) A n a l v e i n l o n g , r e a c h i n g t h e w i n g m a r g i n (fig. 254). 1 or 2 pair(s) o f F - b r i s
tles, c u r v i n g o u t w a r d , s o m e t i m e s c u r v i n g s o m e w h a t o b l i q u e l y b a c k w a r d s
as w e l l (fig. 253). Heleomyzidae
Anal vein not reaching the w i n g margin (fig. 256,257a). F -bristles absent (fig. 257b),
or not curving outward, or more than 2 pairs of F-brisdes present (fig. 255). 118
118(117) A s m a l l , l u s t r o u s b u l g e p r e s e n t i n b e t w e e n a n t e n n a a n d m o s t anterior v i b -
rissa-like bristle; this b u l g e s l i g h t l y larger t h a n a n o c e l l u s a n d m o r e o r less
o f t h e s a m e c o l o u r (fig. 255); interfrontal b r i s t l e s p r e s e n t ; 3 t o 5 pairs o f F -
bristles p r e s e n t . Tethinidae: Tethininae
Lustrous bulge absent. Combination of other characters not as given above. 119
119(118) U s u a l l y y e l l o w or p a l e flies w i t h l i g h t - c o l o u r e d b r i s t l e s a n d p u b e s c e n c e ;
s o m e t i m e s w i t h b r o w n , g r e y o r b l a c k m a r k i n g s o n h e a d , t h o r a x and
a b d o m e n . V e i n s Sc a n d R i m o r e or less parallel; c o s t a w i t h a s u b c o s t a l
b r e a k (fig. 256). Chyromyidae
U s u a l l y s i l v e r y g r e y flies w i t h dark b r i s t l e s a n d p u b e s c e n c e , i n s o m e c a s e s
w i t h b r o w n , g r e y or b l a c k m a r k i n g s o n h e a d , t h o r a x and a b d o m e n , or flies
b l a c k a n d l u s t r o u s . In m o s t s p e c i e s , v e i n s Sc a n d R i a p p r o a c h e a c h o t h e r
b e f o r e t h e y s e p a r a t e l y reach t h e c o s t a ; c o s t a w i t h o u t b r e a k s (fig. 257a).
E s p e c i a l l y i n t h e large g e n u s Leucopis dorsal b r i s t l e s o f t h e h e a d l a r g e l y
a b s e n t (fig. 257b). Chamaemyiidae
120(108) C r o s s v e i n B M - C u p r e s e n t (cells b m a n d d m separate); cell c u p u s u a l l y
closed ( w i n g figures 260-272^ o p e n in s o m e Periscelididae and
Drosophilidae). 121
> [In s e v e r a l s p e c i e s o f M i l i c h i i d a e , cells b m a n d c u p are rather i n c o n s p i c u
o u s a n d t h e c l a r i t y o f c r o s s v e i n B M - C u is v a r i a b l e . A p a r t f r o m t h e
i n c u r v e d F-bristles (fig. 273, 274), t h e s e s p e c i e s s h o w o n e o r m o r e o f t h e
f o l l o w i n g characters: w i n g v e n a t i o n as i n fig. 277 (basal e n d o f cell d m is
acute); b o d y colour black, lustrous; m o u t h p a r t s conspicuously elongate
and f o l d e d b a c k (fig. 276), a d i s t i n c t d e e p n o t c h at t h e p o i n t w h e r e v e i n s Sc
and R i reach t h e c o s t a (fig. 278).]

69
 cell cup

261 wing Odiniidae

(Odinia)

266 habitus female Agromyzidae

(Agromyza)
262 wing Odiniidae
(Turanodinia)
cell cup

BM-Cu

267 wing Anthomyzidae

cell c u p \ J (Paranthomyza)
I \ BM-Cu

V
263 wing Agromyzidae (Liriomyza) J ^ x v 268 femur fore leg
Xj^ Anthomyzidae
(Anthomyza)

Interfrontal
setulae

264 wing Agromyzidae (Phytomyza) 269 head

Anthomyzidae
(Amygdalops)

270 head
Chyromyidae
(Aphaniosoma)
lower F-brlstle
curved forward

271 head
Drosophilidae
265 head Agromyzidae (Cacoxenus)
(Napomyza)
7
C r o s s v e i n B M - C u a b s e n t (cells b m a n d d m f u s e d ) ; cell c u p o p e n or a b s e n t
(fig. 258), c l o s e d i n P s e u d o p o m y z i d a e (fig. 2 9 6 ) , c l o s e d or o p e n i n
Drosophilidae. 128
O n l y 1 pair o f F - b r i s t l e s , c u r v i n g b a c k w a r d ; s e c o n d a n t e n n a l s e g m e n t c o v
e r i n g t h e t h i r d s e g m e n t like a h o o d ; arista w i t h l o n g rays (fig. 259). C e l l
c u p o p e n or c l o s e d (fig. 260), c r o s s v e i n D M - C u s o m e t i m e s w e a k e n e d and
i n 1 s p e c i e s absent. Periscelididae
(In E u r o p e 1 g e n u s , Periscelis, w i t h 4 s p e c i e s . )
M o r e t h a n 1 pair o f F-bristles p r e s e n t (head figures 2 6 5 - 2 7 6 ) . C e l l c u p
c l o s e d (fig. 261-264). O t h e r characters v a r i a b l e . 122
W i n g v e n a t i o n as i n fig. 261 ( c r o s s v e i n s a n d e n d o f v e i n R i w i t h dark
spots) or fig. 262 ( c r o s s v e i n D M - C u i n c o m p l e t e ) . 3 pairs o f F-bristles p r e s
ent, o f w h i c h the l o w e r pair c u r v i n g i n w a r d (fig. 214). Odiniidae
C o m b i n a t i o n o f characters n o t as g i v e n a b o v e . N o F-bristles c u r v i n g
i n w a r d (fig. 2 6 9 , 270, 271) or m o r e t h a n 1 pair c u r v i n g i n w a r d (fig. 265,
272a, 273, 274); i f o n l y 1 pair o f i n c u r v e d F - b r i s t l e s is p r e s e n t ( s o m e
A g r o m y z i d a e ) , t h e n c r o s s v e i n D M - C u i n basal h a l f o f t h e w i n g (fig. 263) or
a b s e n t a l t o g e t h e r (fig. 2 6 4 ) . 123
P-bristles d i v e r g i n g ; at least t h e l o w e r m o s t F-bristles c u r v i n g i n w a r d ;
interfrontal bristles a b s e n t , at m o s t s o m e s e t u l a e p r e s e n t (fig. 265). W i n g
n o t m a r k e d ; c o s t a w i t h a s u b c o s t a l b r e a k ; v e i n b o r d e r i n g cell b m o n t h e
u p p e r side o f t e n p a l l i d or a b s e n t (fig. 263, 2 6 4 ) ; c r o s s v e i n D M - C u o f t e n
a b s e n t (fig. 2 6 4 ) . F e m a l e w i t h a n o n - r e t r a c t a b l e o v i p o s i t o r , v i s i b l e as a
c o n s p i c u o u s c o n i c a l s t r u c t u r e (fig. 266). A g r o m y z i d a e : P h y t o m y z i n a e
[In the g e n u s Xeniomyza, w i t h o n e s p e c i e s in K a z a k h s t a n and a n o t h e r
s p e c i e s i n S p a i n , w h i c h is o n e o f the s m a l l e s t E u r o p e a n a g r o m y z i d s p e c i e s ,
m e a s u r i n g 0.5 m m , t h e r e are n o P - b r i s t l e s . A l l o t h e r characters o b t a i n ,
i n c l u d i n g the w i n g v e n a t i o n characters as i n fig. 264.]
C o m b i n a t i o n o f characters n o t as g i v e n a b o v e . If t h e l o w e r m o s t F - b r i s t l e s
are c u r v i n g i n w a r d , t h e n interfrontal b r i s d e s p r e s e n t . F e m a l e o v i p o s i t o r
retractable, n o c o n s p i c u o u s e x t e r n a l c o n i c a l s t r u c t u r e p r e s e n t . 124
C o s t a w i t h s u b c o s t a l b r e a k o n l y (fig. 2 5 6 , 2 6 7 ) . 2 t o 6 pairs o f F-bristles p r e s
e n t (rarely F-bristles absent), u s u a l l y c u r v i n g b a c k w a r d or s l i g h d y o u t w a r d
as w e l l , in s o m e cases t h e l o w e r pair c u r v i n g i n w a r d (fig. 2 6 9 , 270). 125
C o s t a w i t h h u m e r a l and s u b c o s t a l b r e a k s (fig. 272b, 277, 278). A l m o s t
a l w a y s 1 or 2 pair(s) o f l o w e r F-bristles c u r v i n g f o r w a r d (fig. 271) or i n w a r d
(fig. 272a, 273, 274). 126

,P ocellar
/ \ triangle

H I I R1

272b wing Carnidae (Hemeromyia)

272a head Carnidae (Hemeromyia)

 cell cup

cell dm
lower
F-bristles
incurved
277 wing Milichiidae (Madiza)
deep notch
273 head Milichiidae (Desmometopa)

cell cup

DM-Cu
BM-Cu sometimes vague or absent

278 wing Milichiidae (Milichia)

274 head Milichiidae (Phyllomyza)

BM-Cu a b s e n t ^

279 wing Asteiidae (Leiomyza)

cell cup absent ^ \

275 head Milichiidae (Phyllomyza)
BM-Cu absent
DM-Cu absent

276 head Milichiidae (Madiza) 281 head Asteiidae (Asteia)

7*
 125(124) 2 to 3 pairs o f F-bristles c u r v i n g b a c k w a r d , o f t e n w i t h 1 or m o r e pairs o f
w e a k e r b r i s t l e s i n front; i n t e r f r o n t a l area bare or w i t h f e w s e t u l a e (fig.
269). F e m u r o f fore l e g i n s e v e r a l g e n e r a w i t h a s h o r t s p i n e o n t h e v e n t r a l
side, b e y o n d t h e m i d d l e (fig. 268). B r i s t l e s a n d p u b e s c e n c e n o t y e l l o w .
W i n g v e n a t i o n as i n fig. 267. Anthomyzidae
- 2 t o 6 pairs o f F - b r i s t l e s p r e s e n t (rarely F - b r i s d e s a b s e n t ) , u s u a l l y c u r v i n g
b a c k w a r d or s l i g h t l y o u t w a r d as w e l l , i n s o m e c a s e s t h e l o w e r pair c u r v i n g
i n w a r d ; i n t e r f r o n t a l area w i t h m a n y s e t u l a e (fig. 270). F e m u r o f fore l e g
w i t h o u t s p i n e o n v e n t r a l side. U s u a l l y y e l l o w or pallid flies, their b r i s d e s
a n d p u b e s c e n c e also o f a pale c o l o u r , i n s o m e c a s e s w i t h b r o w n , g r e y or
b l a c k m a r k i n g s o n h e a d , t h o r a x a n d a b d o m e n . W i n g v e n a t i o n as i n fig.
256, b u t v e i n Sc apically pallid. Chyromyidae
126(124) Interfrontal bristles a b s e n t , h e a d as i n fig 271. D r o s o p h i l i d a e : Cacoxenus
- Interfrontal b r i s d e s p r e s e n t , h e a d as i n figures 2728-276. 127
127(126) O n l y 1 pair o f i n t e r f r o n t a l b r i s t l e s , i n s e r t e d j u s t a b o v e t h e a n t e n n a e ; o c e l -
lar triangle large, o b v i o u s ; P - b r i s t l e s s o m e w h a t d i v e r g i n g t o parallel (fig.
272a). A n a l v e i n l o n g (fig. 272b). Carnidae: Hemeromyia
M o r e t h a n a s i n g l e pair o f i n t e r f r o n t a l b r i s t l e s , a r r a n g e d i n r o w s (fig. 273)
o r m o r e irregular; ocellar triangle s m a l l e r t h a n s h o w n i n fig. 272a; P - b r i s
tles parallel or c o n v e r g i n g (fig. 273, 274); t h i r d a n t e n n a l s e g m e n t a n d
m o u t h p a r t s i n s o m e c a s e s c o n s p i c u o u s l y large or e l o n g a t e (fig. 275, 276).
A n a l v e i n s h o r t or a b s e n t (fig. 277, 278). Milichiidae
128(120) W i n g w i t h a c o n s p i c u o u s n o t c h w h e r e v e i n R i e n d s in t h e c o s t a ; c r o s s v e i n
D M - C u i n the apical p a r t o f t h e w i n g (fig. 278). T h e 2 l o w e r pairs o f F - b r i s
tles c u r v i n g i n w a r d ; m o r e t h a n 1 pair o f i n t e r f r o n t a l b r i s t l e s , a r r a n g e d i n
r o w s or m o r e irregular; P - b r i s t l e s parallel o r c o n v e r g i n g (fig. 273, 274).
Milichiidae
W i n g a n d b r i s t l e s o f t h e h e a d n o t as d e s c r i b e d a b o v e . If a n o t c h p r e s e n t
w h e r e v e i n R i e n d s i n the c o s t a a n d 2 l o w e r pairs o f F - b r i s t l e s c u r v i n g
i n w a r d , t h e n c r o s s v e i n D M - C u e i t h e r a b s e n t (fig. 288) or i n t h e b a s a l h a l f
o f the w i n g (fig. 289) a n d o n l y 1 pair o f i n t e r f r o n t a l b r i s t l e s p r e s e n t , i n s e r t
e d j u s t a b o v e t h e a n t e n n a e (fig. 286, 287). 129
129 (128) C e l l r4+5 e n d s near the w i n g tip a n d is s l i g h t l y n a r r o w i n g since v e i n s R 4 + 5
a n d M i are s o m e w h a t c o n v e r g i n g ; c o s t a c o n t i n u o u s (fig. 2 7 9 , 280); i n s e v
eral g e n e r a , v e i n R2+3 s t r i k i n g l y s h o r t , e n d i n g i n t h e basal h a l f o f t h e w i n g
(fig. 280). A r i s t a s o m e t i m e s bare or p u b e s c e n t b u t u s u a l l y w i t h a c h a r a c
teristic z i g - z a g o f l o n g e r or s h o r t e r d o r s a l a n d v e n t r a l r a y s (fig. 281) or, i n
o n e s p e c i e s , arista a b s e n t . Asteiidae

a Trachysiphonella b Gampsocera c Eurina d Chlorops

282 a-d head Chloropidae

73
 sharp carina

proepisternum

interfrontal bristles-

283 thorax Chloropidae 287 head Carnidae (Meoneura)

(Chlorops)

DM-Cu absent ' %

288 wing Carnidae (Carnus)

I J

284 habitus Chloropidae (Meromyza)

DM-Cu

R2+3 289 wing Carnidae (Meoneura)

CuA1 with flexure

285 wing Chloropidae (Rhodesiella)

P-bristles absent
lower
F-bristles
incurved

interfrontal
bristles

290 habitus Ephydridae (Coenia)

286 head Carnidae

cell cup

BM-Cu absent
74
291 wing Ephydridae (Paralimna)
 - C e l l r4+5 n o t as d e s c r i b e d a b o v e , i f n a r r o w i n g , t h e n u s u a l l y b e c a u s e v e i n s
R 4 + 5 a n d M i are b o t h c u r v i n g t o w a r d t h e c o s t a ; c o s t a n e a r l y a l w a y s w i t h
a s u b c o s t a l b r e a k ( c o n t i n u o u s i n s o m e s p e c i e s o f C h l o r o p i d a e ) ; v e i n R2+3
n o t s t r i k i n g l y s h o r t ( w i n g figures 284-298). A r i s t a bare, p u b e s c e n t or w i t h
s h o r t e r t o l o n g e r r a y s , b u t w i t h o u t a characteristic z i g - z a g a r r a n g e m e n t
(fig. 282, 2 9 0 , 297). 130
130(129) Frons nearly a l w a y s w i t h a strikingly large, often l u s t r o u s ocellar triangle (fig.
282 b - d ) . P r o e p i s t e r n u m w i t h a sharp anterior carina (fig. 283). V e i n C u A i
usually w i t h a flexure h a l f w a y cell b m + d m ; often v e i n R4+5 and s o m e t i m e s
also v e i n M i c u r v i n g t o w a r d t h e costa; costa w i t h a subcostal break (fig. 2 8 4 ,
285; i n o n l y a f e w species the costa is c o n t i n u o u s ) . Chloropidae
O c e l l a r triangle n o t c o n s p i c u o u s l y large or l u s t r o u s ( e x c e p t i n C a r n i d a e ,
fig. 286). P r o e p i s t e r n u m a n t e r i o r l y w i t h o u t a s h a r p carina. V e i n C u A i
w i t h o u t flexure; v e i n s R 4 + 5 a n d M i rarely (in s o m e E p h y d r i d a e ) c u r v i n g
t o w a r d the costa; costa t w i c e interrupted, w i t h humeral and subcostal
b r e a k s (fig. 288-291, 2 9 6 , 298). 131
131(130) 4 pairs o f F-brisdes present, the l o w e r 2 pairs c u r v i n g i n w a r d , the u p p e r 2 pairs
c u r v i n g o u t w a r d or b a c k w a r d ; 1 pair o f interfrontal brisdes present, inserted
just a b o v e the antennae (fig. 286,287). Carnidae: Camus, Meoneura
> [Camus: P-bristles a b s e n t (fig. 286); c r o s s v e i n D M - C u a b s e n t (fig. 288),
w i n g m i g h t b e s h e d o f f near t h e b a s e (fig. 4 4 9 ) . Meoneura: P - b r i s t l e s par
allel (fig. 287); c r o s s v e i n D M - C u p r e s e n t (fig. 289).]
F-bristles a n d i n t e r f r o n t a l b r i s t l e s n o t as d e s c r i b e d a b o v e . 132
132(131) N o real P-bristles p r e s e n t , b u t s o m e t i m e s parallel or d i v e r g i n g p s e u d o p o s -
tocellar bristles p r e s e n t (fig. 293); arista u s u a l l y w i t h l o n g r a y s , l i m i t e d t o
t h e u p p e r side (fig. 290), arista s o m e t i m e s bare (fig. 202) or s h o r t p u b e s
c e n t ; face u s u a l l y s w o l l e n , c o n v e x (fig. 2 9 0 , 292). C e l l c u p o p e n , anal v e i n
a b s e n t (fig. 291). Ephydridae
P-bristles p r e s e n t , c o n v e r g i n g ; i f arista p l u m o s e t h e n w i t h l o n g r a y s o n
b o t h sides (e.g. fig. 297), b u t arista m a y b e bare or p u b e s c e n t o n l y . Face n o t
s w o l l e n (fig. 2 9 4 , 297). C e l l c u p o p e n or c l o s e d ; anal v e i n p r e s e n t or a b s e n t
(fig. 2 9 6 , 298). 133

292 head Ephydridae (Limnellia)

294 head
Pseudopomyzidae
(Pseudopomyza)

295 head Pseudopomyzidae

(Pseudopomyza)

75
293 head Ephydridae (Thinoscatella)
298 wing Drosophilidae (Drosophila)
303 wing Lauxaniidae (Eusapromyza)

300 habitus Conopidae (Leopoldius)

 2
133O3 ) A p a r t f r o m the c o n v e r g i n g P - b r i s t l e s , a n o t h e r pair o f c o n v e r g i n g bristles
p r e s e n t ; all 3 pairs o f F - b r i s t l e s c u r v i n g b a c k w a r d ; arista p u b e s c e n t (fig.
2 9 4 , 295). W i n g v e n a t i o n as s h o w n i n fig. 2 9 6 . Pseudopomyzidae
(In E u r o p e 1 w i d e s p r e a d s p e c i e s , Pseudopomyza atrimana ( M e i g e n ) . )
O n l y t h e c o n v e r g i n g P - b r i s t l e s p r e s e n t ; o f t h e 3 pairs o f F - b r i s t l e s , 1 pair is
c u r v i n g f o r w a r d or o b l i q u e l y i n w a r d ; arista u s u a l l y w i t h l o n g r a y s (fig.
297; see for F - b r i s t l e s a n d arista also c o u p l e t 9 9 a n d fig. 271). W i n g v e n a
t i o n as in fig. 298. Drosophilidae
134(84) A t the same time cell c u p elongate (approaching the h i n d m a r g i n o f the w i n g )
a n d v e i n C11A2 (i.e., the v e i n c l o s i n g cell cup) m o r e or less straight; cell r4+5
n a r r o w i n g t o w a r d the w i n g tip, or c l o s e d (fig. 299, 300). M o u t h p a r t s u s u a l
ly l o n g , slender, g e n i c u l a t e at the b a s e , i n s o m e cases t w i c e g e n i c u l a t e (fig.
299); i n s o m e cases m o u t h p a r t s s h o r t (fig. 300). Conopidae
- Cell cup short or absent, its apex not near the hind margin o f the w i n g ( w i n g fig
ures 302-347), or if long, then closed b y a geniculate v e i n Q 1 A 2 (fig. 301). 135
> [A g e n i c u l a t e v e i n C u A 2 , t h e v e i n c l o s i n g cell c u p , o n l y o c c u r s i n U l i d i i d a e
(fig. 301), Salticella ( S c i o m y z i d a e , fig. 307), P y r g o t i d a e (fig. 324) a n d
T e p h r i t i d a e (fig. 330).]
*35C34) D o r s a l preapical bristle p r e s e n t at l e a s t o n tibia o f m i d l e g b u t u s u a l l y o n
tibiae o f all l e g s (fig. 309). V e i n Sc c o m p l e t e , r e a c h i n g t h e c o s t a i n d e p e n d
e n t l y ( w i n g figures 301-320). 136
Tibiae w i t h o u t dorsal preapical b r i s d e . V e i n Sc c o m p l e t e or i n c o m p l e t e . 147
136(135) A n a l v e i n e n d i n g d i s t i n c t l y b e f o r e t h e w i n g m a r g i n (fig. 302, 303). 137
- A n a l v e i n l o n g , r e a c h i n g the w i n g m a r g i n ( w i n g figures 3 0 6 - 3 2 4 ) or e n d i n g
close to it, in s o m e cases v e r y t h i n or a l m o s t f o l d - l i k e near t h e e n d . 139
137(136) W i n g w i t h characteristic, b a n d e d m a r k i n g s (fig. 302).
S c i o m y z i d a e : Colobaea
- W i n g u n m a r k e d or w i t h m a r k i n g s different. 138
138(137) P - b r i s t l e s a l m o s t a l w a y s c o n v e r g i n g (fig. 3 0 4 ) , v e r y r a r e l y p a r a l l e l .
P o s t e r i o r spiracle w i t h o u t b r i s t l e s at t h e p o s t e r o v e n t r a l m a r g i n (see a l s o
c o u p l e t 91). Lauxaniidae
> [Some s p e c i m e n s o f the s c i o m y z i d g e n u s Pteromicra m a y k e y o u t here
because s o m e t i m e s their P-brisdes are s l i g h d y c o n v e r g i n g a n d the c o n t i n u a
tion o f the anal v e i n to the w i n g m a r g i n can be a v e r y t h i n fold only.
Pteromicra differs from Lauxaniidae b y the f o l l o w i n g c o m b i n a t i o n o f charac
ters: b o d y s h i n y b r o w n i s h black or b r o w n i s h black and y e l l o w , fore legs large
ly b r o w n i s h black, other legs m a i n l y y e l l o w , a n e p i s t e r n u m w i t h o u t bristles.]
- P-bristles d i v e r g i n g (fig. 305) or a b s e n t . P o s t e r i o r spiracle w i t h 1 or m o r e
bristles at t h e p o s t e r o v e n t r a l m a r g i n (fig. 1 9 6 , 1 9 7 ) (see also c o u p l e t 91).
Sepsidae
307 wing Sciomyzidae (Salticella)

311 head Pallopteridae (Eurygnathomyia)

, Sc R1

308 head Sciomyzidae cell r4+5

(Coremacera)

312 wing Pallopteridae (Eurygnathomyia)

preapical-^'l^ katepisternum
/yfSi \ katepisternum
bristle % fl^apical bristle

309 habitus Sciomyzidae (Psacadina) V

313 katepisternum Pallopteridae
(Eurygnathomyia)

310 habitus Phaeomyiidae (Pelidnoptera) 314 head Dryomyzidae (Neuroctena)

78
139(136) V e i n R i w i t h s e t u l a e o n its dorsal side (as i n fig. 306) a n d at t h e s a m e t i m e
w i n g m e m b r a n e w i t h a p a t t e r n e d ( n o t n e c e s s a r i l y as i n fig. 306).
Ulidiidae
- C o m b i n a t i o n o f c h a r a c t e r s n o t as g i v e n a b o v e ; i f v e i n R i w i t h s e t u l a e o n its
dorsal side, t h e n w i n g n o t p a t t e r n e d b u t clear, s h a d e d a r o u n d t h e v e i n s
( D r y o m y z i d a e ) o r e n t i r e l y i n f u s c a t e d (fig. 310: P h a e o m y i i d a e ) . 140
140(139) V e i n C u A 2 (the v e i n c l o s i n g cell c u p ) g e n i c u l a t e , c a u s i n g cell c u p t o t a p e r
t o a n a c u t e apical a n g l e ; cell r4+5 n a r r o w i n g t o w a r d t h e w i n g tip (fig. 301,
307)- H i
V e i n C u A 2 straight or c o n v e x ; cell R 4 + 5 n e a r l y a l w a y s b r o a d t o w a r d t h e
w i n g tip (fig. 3 0 9 , 310, 3 1 2 , 3 2 0 ) . 142
141(140) Thorax lustrous, greenish blue; a b d o m e n lustrous, blackish b r o w n . W i n g
w i t h o u t a b r o w n i s h t i n g e ; cell c u p l o n g , n a r r o w i n g t o a n a c u t e apical e n d
(fig. 301). F e m a l e w i t h a l a n c e t - s h a p e d , i n c o m p l e t e l y retractable o v i p o s i
tor (as i n fig. 350, 351). Ulidiidae
- B o d y n o t c o n s p i c u o u s l y l u s t r o u s ; t h o r a x b r o w n w i t h darker l o n g i t u d i n a l
stripes o r s p o t s a b o v e a n d at s i d e s ; a b d o m e n l a r g e l y grey. W i n g w i t h
b r o w n i s h t i n g e , c r o s s v e i n R - M w i t h dark b o r d e r s ; cell c u p less e l o n g a t e
a n d less a c u t e (fig. 307). F e m a l e w i t h o u t a l a n c e t - s h a p e d o v i p o s i t o r .
S c i o m y z i d a e : Salticella
142(140) C l y p e u s s m a l l a n d w i t h d r a w n , i n lateral v i e w m o r e o r less h i d d e n b e h i n d
t h e oral m a r g i n ; s e c o n d a n t e n n a l s e g m e n t o f t e n e l o n g a t e (fig. 3 0 8 ) .
K a t e p i s t e r n u m bare o r s l i g h t l y p u b e s c e n t , rarely w i t h 1 o r 2 b r i s t l e s near its
u p p e r m a r g i n (fig. 3 0 9 , 310). 143
C l y p e u s w e l l v i s i b l e , p r o t r u d i n g b e l o w t h e oral m a r g i n (fig. 311, 314) o r
p r o t r u d i n g f r o m a n e m a r g i n a t i o n o f t h i s m a r g i n (fig. 3 1 9 , 321); s e c o n d
antennal segment always short. Katepisternum always w i t h m o r e than 1
bristle near its u p p e r m a r g i n ( u s u a l l y 3 o r m o r e ) (fig. 313, 315) or, i n case o f
Helcomyza, k a t e p i s t e r n u m g r e y a n d w i t h a d e n s e c o v e r o f dark h a i r - l i k e
bristles. 144
143(142) T i b i a e w i t h apical a n d dorsal p r e a p i c a l b r i s t l e s o n l y , n o b r i s t l e s h a l f w a y u p
(fig. 309). W i n g o f t e n w i t h a p a t t e r n , i n s o m e s p e c i e s o n l y i n f u s c a t e d o r
all clear; v e i n R i w i t h o u t s e t u l a e o n its dorsal side. A r i s t a p u b e s c e n t o r
w i t h l o n g e r rays; s e c o n d a n t e n n a l s e g m e n t o f t e n e l o n g a t e (fig. 308).
Sciomyzidae
T i b i a o f h i n d l e g , a n d s o m e t i m e s tibia o f m i d l e g as w e l l , w i t h apical a n d
dorsal preapical bristles b u t also w i t h b r i s t l e s h a l f w a y u p their l e n g t h (fig.
310). W i n g i n f u s c a t e d , w i t h o u t p a t t e r n ; v e i n R i w i t h s e t u l a e o n its u p p e r
side. A r i s t a p u b e s c e n t ; s e c o n d a n t e n n a l s e g m e n t a l w a y s s h o r t .
Phaeomyiidae
(In E u r o p e 1 g e n u s , Pelidnoptera, w i t h 3 species.)
144(142) H e a d as i n fig. 311: 2 pairs o f F - b r i s t l e s c u r v i n g b a c k w a r d , t h e l o w e r o n e t h e
s m a l l e s t . W i n g as i n fig. 312: c o s t a w e a k e n e d j u s t b e y o n d c r o s s v e i n H a n d
w i t h a subcostal break; v e i n C u A 2 c o n v e x .
Pallopteridae: Eurygnathomyia

7 9
 Oc-bristles absent

clypeus '

319 head Helcomyzidae (Helcomyza)

315 habitus Dryomyzidae (Dryomyza)

lateral plate

prosternum
320 wing Helcomyzidae (Helcomyza)

316 ventral view thorax Dryomyzidae (Dryomyza)

clypeus
321 head
Heterocheilidae
(Heterocheila)

317 ventral view thorax Helcomyzidae (Helcomyza)

^ B M - C u absent

318 ventral view thorax Helcomyzidae 322 habitus Cryptochetidae

(Helcomyza) (Cryptochetum)

8 o
 H e a d a n d w i n g different; c o s t a c o n t i n u o u s ; v e i n C u A 2 straight (fig. 315,
320). 145
145(144) A n t e n n a e i n s e r t e d c l o s e t o g e t h e r , n o t further apart t h a n t h e l e n g t h o f
their s e c o n d s e g m e n t (fig. 314). C o s t a w i t h c l o s e l y s p a c e d p u b e s c e n c e ,
w i t h o u t s t r o n g s h o r t b r i s t l e s (fig. 315). U n d e r s i d e o f t h o r a x w i t h o u t p r e -
c o x a l b r i d g e ( p r o s t e r n u m linear t o o v o i d , separate f r o m t h e lateral p l a t e s
b y a m e m b r a n e , fig. 316). Y e l l o w t o r e d d i s h c o l o u r e d flies. D r y o m y z i d a e
A n t e n n a l i n s e r t i o n s further apart, a b o u t t w i c e t h e l e n g t h o f t h e i r s e c o n d
s e g m e n t (fig. 3 1 9 , 321). C o s t a i n s o m e c a s e s w i t h s t r o n g s h o r t b r i s t l e s (fig.
320). U n d e r s i d e o f t h o r a x w i t h a p r e c o x a l b r i d g e ( p r o s t e r n u m f u s e d w i t h
t h e lateral p l a t e s , fig. 317, 318). Flies o f c o a s t a l habitats, b r o w n i s h or g r e y
coloured. 146

146(145) B o d y c o l o u r m a i n l y grey. O c - b r i s t l e s a b s e n t o r w e a k l y d e v e l o p e d ; l-z pairs

o f less s t r o n g F-bristles p r e s e n t (fig. 319). W i n g w i t h s t r o n g b r i s t l e s a l o n g
c o s t a (fig. 320). P r o s t e r n u m w i t h l o n g b r i s t l e s (fig. 318). [Flies o f c o a s t a l
habitats.] Helcomyzidae
(In E u r o p e 1 g e n u s , Helcomyza, w i t h 2 s p e c i e s . )
B o d y c o l o u r m a i n l y b r o w n . O c - and F-bristles w e l l d e v e l o p e d ; 3 pairs o f
F-bristles p r e s e n t (fig. 321). W i n g w i t h o u t s t r o n g b r i s t l e s a l o n g c o s t a .
P r o s t e r n u m bare. [Flies o f c o a s t a l habitats.] Heterocheilidae
(In E u r o p e , a l o n g N o r t h Sea a n d Baltic Sea, 1 s p e c i e s , Heterocheila bucca-
ta (Fallen).)

147035) A r i s t a a b s e n t ; t h i r d a n t e n n a l s e g m e n t c o n s p i c u o u s l y large; ocellar triangle

large. V e i n Sc c o m p l e t e u p t o t h e c o s t a , b u t w e a k , h a r d l y v i s i b l e ; c r o s s v e i n
B M - C u a b s e n t (cells b m a n d d m f u s e d ) . S m a l l , s t o u t flies w i t h a b l a c k or
m e t a l l i c b l u e lustre (fig. 322). Cryptochetidae
(In E u r o p e 1 g e n u s , Cryptochetum, w i t h 3 species.)
A r i s t a p r e s e n t and c o m b i n a t i o n o f o t h e r characters n o t as g i v e n a b o v e .

148(147) O c e l l i absent; s e c o n d a n t e n n a l s e g m e n t v e r y e l o n g a t e (fig. 323). V e i n Sc

e n d i n g j u s t b e f o r e t h e c o s t a ; v e i n R2+3 n e a r l y a l w a y s w i t h an a p p e n d i x
(fig. 324). Pyrgotidae
(In C e n t r a l and S o u t h e r n E u r o p e 1 s p e c i e s , Adapsilia coarctata W a g a . )

appendix

324 wing Pyrgotidae (Adapsilia)

8 1
- O c e l l i p r e s e n t ; s e c o n d a n t e n n a l s e g m e n t u s u a l l y n o t e l o n g a t e (fig. 325,
327). W i n g v e n a t i o n different. 149
0
149(148) V e i n Sc a b r u p t l y b e n t f o r w a r d t o w a r d t h e c o s t a at n e a r l y 9 0 , g o i n g
t o w a r d t h e c o s t a as a t r a n s p a r e n t line or trace (fig. 3 2 6 , 3 2 9 , 330). 150
V e i n Sc n o t a b r u p t l y b e n t f o r w a r d . 151
150(149) F-bristles, if present, restricted to the upper part o f the head, not c u r v i n g
i n w a r d ; face o f t e n r e c e d i n g , t h i r d a n t e n n a l s e g m e n t i n s o m e c a s e s e l o n
gate (fig. 325). W i n g u n m a r k e d ; cell c u p c l o s e d b y a s t r a i g h t v e i n C u A 2 ,
n o t n a r r o w i n g t o an a c u t e e n d ; basal p a r t o f w i n g w i t h a t r a n s p a r e n t line
c o n n e c t i n g t h e (virtual) e n d i n g o f Sc a n d t h e e n d o f cell b m (fig. 326, 327).
Psilidae
- A l m o s t a l w a y s s e v e r a l l o w e r pairs o f F - b r i s t l e s c u r v i n g i n w a r d (fig. 328).
W i n g o f t e n m a r k e d ; cell c u p n e a r l y a l w a y s c l o s e d b y a g e n i c u l a t e v e i n
Q 1 A 2 a n d w i t h an a c u t e apical e n d , v e i n C u A 2 rarely s t r a i g h t or c o n v e x ; n o
t r a n s p a r e n t line i n basal p a r t o f t h e w i n g (fig. 330, 331). Tephritidae
151(149) V e i n Sc c o m p l e t e , its apical p a r t n o t p a l l i d , r e a c h i n g t h e c o s t a separately
f r o m v e i n R i ( w i n g figures 333-357). N o t e t h a t t h e e n d s o f v e i n s Sc a n d R i
m a y b e c l o s e t o g e t h e r , as i n fig. 333, 339). 152
- V e i n Sc a b s e n t , s h o r t e n e d , m e r g i n g w i t h R i b e f o r e t h e c o s t a , or its apical
p a r t pallid a n d v a g u e (fig. 332, w i n g figures 358-369). 164
152(151) M o u t h p a r t s e l o n g a t e , slender, t w i c e g e n i c u l a t e . V e i n s Sc a n d R i e n d i n t h e
costa c l o s e t o g e t h e r , h a l f w a y u p t h e w i n g ; c o s t a c o n t i n u o u s , cell r4+5 nar
r o w i n g t o w a r d w i n g tip, cell c u p c l o s e d b y s t r a i g h t v e i n C11A2; anal v e i n
l o n g (fig. 333). C o n o p i d a e : Dalmannia
- M o u t h p a r t s a n d w i n g v e n a t i o n n o t as d e s c r i b e d a b o v e . 153

332 wing base Strongylophthalmyiidae 333 habitus Conopidae (Dalmannia)

(Strongylopthalmyia)

83
84
1 I 2
53( 5 ) F e m u r o f h i n d l e g s w o l l e n , its l o w e r side w i t h 2 r o w s o f s p i n e s (fig. 3 3 4 ,
335a). O c - b r i s t l e s a b s e n t . H a b i t u s , e s p e c i a l l y h e a d a n d w i n g as i n figure
334 o r 336. 154
F e m u r o f h i n d l e g n o t s w o l l e n , w i t h o u t s p i n e s o n l o w e r side. O c - b r i s t l e s
absent or present. Habitus o t h e r w i s e . 155
154(153) S e c o n d a n t e n n a l s e g m e n t l e n g t h e n e d . H a b i t u s as i n fig. 3 3 4 .
S c i o m y z i d a e : Sepedon
- S e c o n d a n t e n n a l s e g m e n t n o t l e n g t h e n e d (fig. 335b). H a b i t u s as i nfig.336.
Megamerinidae
(In E u r o p e 1 w i d e s p r e a d s p e c i e s , Megamerina dolium (Fabricius).
155(153) H e a d u n u s u a l l y w i d e , e y e s o n s h o r t s t a l k s . C r o s s v e i n B M - C u a b s e n t (cells
b m a n d d m fused) (fig. 337). Diopsidae
(In C e n t r a l E u r o p e 1 s p e c i e s , Sphyracephala europaea P a p p &. Foldvari.)
- H e a d n o t as w i d e , e y e s n o t o n stalks. C r o s s v e i n B M - C u p r e s e n t (cells b m
a n d d m separate) ( w i n g figures 338-357), e x c e p t i n M i c r o p e z i d a e , s u b f a m
i l y M i c r o p e z i n a e (fig. 339, cell r4+5 n a r r o w i n g ) . 156
156(155) Largely b l a c k l u s t r o u s , s m a l l , a n t - l i k e flies w i t h r e l a t i v e l y f e w b r i s t l e s o r
p u b e s c e n c e . H e a d r o u n d , a b d o m e n b a s a l l y c o n s t r i c t e d . In m a l e , f e m u r a n d
tibia o f fore l e g o f t e n w i t h p r o t r u s i o n s , s p i n e s , serrate s t r u c t u r e s , etc. (fig.
338). A r i s t a bare. W i n g o f t e n w i t h a dark s p o t at t h e w i n g t i p (Sepsis, fig.
198). P o s t e r i o r spiracle w i t h 1 o r m o r e b r i s t l e s at t h e p o s t e r o v e n t r a l m a r g i n
(fig. 1 9 6 , 1 9 7 ) . Sepsidae
C o m b i n a t i o n o f c h a r a c t e r s n o t as g i v e n a b o v e . P o s t e r i o r spiracle w i t h o u t
bristles along the posteroventral margin. 157
> [Black l u s t r o u s U l i d i i d a e , s u c h as s o m e s p e c i e s o f Herina a n d Seioptera,
fig. 350, m a y k e y o u t h e r e . In t h e U l i d i i d a e t h e p o s t e r i o r spiracle h a s n o
bristles, t h e f e m a l e h a s a n i n c o m p l e t e l y retractable o v i p o s i t o r , t h e apical
e n d o f cell c u p is a c u t e (fig. 3 4 9 - 3 5 1 ) a n d / o r v e i n R i h a s s e t u l a e o n its
u p p e r side (fig. 354).]

338 habitus Sepsidae

(Themira)

85
 cell sc RI
cell cup
CuA2 straight
BM-Cu absent

339 wing Micropezidae (Micropeza)

344 wing Chamaemyiidae (Lipoleucopis)

340 head Micropezidae (Micropeza)

P- bristles absent
Oc-bristles absent

anal vein \

345 wing base Chamaemyiidae (Parochthiphila)

341 head Micropezidae (Calobata)

cell cup

CuA2 convex

342 wing Tanypezidae (Tanypeza)

346 head Chamaemyiidae (Parochthiphila)

343 habitus Tanypezidae (Tanypeza)

347 wing Cremifaniidae

(Cremifania)
86
157(156) C e l l r4+5 n a r r o w i n g t o w a r d w i n g tip, apical e n d o f cell c u p n o t a c u t e (fig.
339, 342); w i n g n o t m a r k e d . L e g s c o n s p i c u o u s l y l o n g and slender. 158
C e l l r4+5 parallel or w i d e n i n g t o w a r d t h e w i n g tip (fig. 3 4 4 ) ; i f n a r r o w i n g
t o w a r d w i n g tip, t h e n apical e n d o f cell c u p a c u t e a n d v e i n C u A 2 g e n i c u
late (fig. 349) a n d / o r w i n g w i t h m a r k i n g s . L e g s n o t c o n s p i c u o u s l y l o n g
and slender. 159
158(157) O c - b r i s t l e s absent; P-bristles d i v e r g i n g (fig. 3 4 0 ) or a b s e n t (fig. 3 4 1 ) . V e i n
Sc separate f r o m R i ; cell c u p c l o s e d b y a straight v e i n C u A 2 ; c r o s s v e i n B M -
C u s o m e t i m e s a b s e n t (fig. 339: s u b f a m i l y M i c r o p e z i n a e ) . M i c r o p e z i d a e
O c - b r i s t l e s p r e s e n t ; P - b r i s t l e s d i v e r g i n g . V e i n s Sc a n d R i t o u c h or
a p p r o a c h e a c h o t h e r o v e r s o m e d i s t a n c e b e f o r e t h e y separately e n d i n t h e
c o s t a ; cell c u p c l o s e d b y a c o n v e x v e i n C u A 2 ; c r o s s v e i n B M - C u p r e s e n t
(fig. 342). H a b i t u s as in fig. 343. Tanypezidae
(In E u r o p e 1 w i d e s p r e a d s p e c i e s , Tanypeza longimana Fallen.)
159(157) S i l v e r y g r e y flies. F e m a l e w i t h o u t a l a n c e t - s h a p e d o v i p o s i t o r . W i n g o f t e n
w i t h d a r k e n e d cell(s) c a n d / o r sc; anal v e i n s h o r t (fig. 3 4 4 , 3 4 5 , 3 4 7 ) . P -
bristles c o n v e r g i n g (fig. 3 4 6 ) or a b s e n t (fig. 348). 160
- Flies o f d i f f e r e n t c o l o u r . F e m a l e w i t h a l a n c e t - s h a p e d , i n c o m p l e t e l y
retractable o v i p o s i t o r (fig. 350, 351, 353, 357). W i n g o f t e n w i t h e x t e n s i v e
m a r k i n g s ; anal v e i n l o n g , o f t e n r e a c h i n g t h e h i n d m a r g i n o f t h e w i n g . P-
b r i s t l e s d i v e r g i n g or a b s e n t (fig. 355, 356). 161
160(159) V e i n Sc r e a c h i n g the c o s t a near t h e e n d o f v e i n R i (fig. 3 4 4 ; i n m o s t s p e c i e s
v e i n s Sc and R i t o u c h or a p p r o a c h e a c h o t h e r b e f o r e t h e y s e p a r a t e l y e n d i n
t h e c o s t a , fig. 345). P-bristles c o n v e r g i n g (fig. 346) or absent. E s p e c i a l l y i n
t h e large g e n u s Leucopis dorsal b r i s t l e s o f t h e h e a d largely a b s e n t (fig. 257b)
Chamaemyiidae
V e i n s Sc a n d R i r e a c h i n g the c o s t a at s o m e d i s t a n c e o f e a c h o t h e r (fig. 3 4 7 ) .
P-bristles a b s e n t (fig. 348). Cremifaniidae
(In E u r o p e 1 g e n u s , Cremifania, w i t h 2 species.)

348 head Cremifaniidae 349 wing Ulidiidae

(Cremifania) (Ulidia)

87
161(159) A p i c a l e n d o f cell c u p a c u t e ; v e i n C u A 2 g e n i c u l a t e or w i t h an S - c u r v e (fig.
349-351). Ulidiidae
C e l l c u p different, v e i n C u A 2 c o n v e x (fig. 352) or s t r a i g h t (fig. 357). 162
162(161) V e i n R i o n u p p e r side w i t h o u t s e t u l a e (fig. 352, 353). Pallopteridae
V e i n R i o n u p p e r side w i t h s e t u l a e (fig. 3 5 4 ) . 163
163(162) C o s t a w i t h o n l y a h u m e r a l b r e a k (just b e y o n d c r o s s v e i n H) (fig. 3 5 4 ) . P -
bristles r e d u c e d or a b s e n t (fig. 355). Platystomatidae
C o s t a c o n t i n u o u s , or w i t h o n l y a s u b c o s t a l b r e a k , or w i t h b o t h h u m e r a l
a n d s u b c o s t a l b r e a k s . P - b r i s t l e s u s u a l l y p r e s e n t , d i v e r g i n g (fig. 356).
Ulidiidae
164(151) V e i n R2+3 l o n g , n e a r l y r e a c h i n g R 4 + 5 a n d w i n g tip; w i n g b a s e u s u a l l y nar
r o w ; anal area r e d u c e d ; o f t e n a s m a l l cell p r e s e n t w h e r e R i e n d s in t h e
c o s t a ; w i n g o f t e n w i t h m a r k i n g s ( w i n g tip w i t h dark s p o t , c r o s s v e i n s
b a n d e d ) ; cell c u p a n d c r o s s v e i n B M - C u p r e s e n t (fig. 358, 359). 1 pair o f F -
bristles c u r v i n g b a c k w a r d (fig. 360). Opomyzidae
> [In case o f h a v i n g o v e r l o o k e d t h e pale y e l l o w v i b r i s s a e i n C h y r o m y i d a e ,
this f a m i l y w i l l k e y o u t h e r e . C h y r o m y i d a e are m o s t l y y e l l o w flies, w i t h a
n o r m a l w i n g b a s e w i d t h (fig. 256), n o m a r k i n g s o n t h e w i n g , c o n v e r g i n g
P-bristles a n d u s u a l l y m o r e t h a n 1 pair o f F - b r i s t l e s (fig. 270).]

359 wing Opomyzidae (Opomyza) 360 head Opomyzidae (Anomalochaeta)

89
 361 head Chloropidae
(Calamoncosis)

R4+5

alula 366 habitus Ephydridae

cell cup (Ochthera)
BM-Cu absent

CuA1 with flexure

362 wing Chloropidae

(Rhodesiella)

DM-Cu

367 habitus
Tethinidae
(Pelomyiinae)
(Pelomyia)

363 habitus Nannodastiidae (Azorastia)

R2+3

I i

alula absent
cell cup absent 7i ^
;

1
BM-Cu absent '''''''niMMiiiniTO^ow*" *
368 wing base
364 wing Nannodastiidae (Azorastia) Strongylophthalmyiida
(Strongylophthalmyia)

9 o
 V e i n R2+3 n o t (nearly) r e a c h i n g R 4 + 5 a n d w i n g tip; w i n g b a s e n o t n a r r o w ;
anal area w e l l d e v e l o p e d (fig. 362); w i n g rarely w i t h m a r k i n g s ; i f w i n g b a s e
s o m e w h a t n a r r o w , t h e n v e i n R2+3 s h o r t , e n d i n g a b o u t h a l f w a y u p t h e
costa ( N a n n o d a s t i i d a e : fig. 363, 3 6 4 ) ; cell c u p and c r o s s v e i n B M - C u p r e s
e n t or absent. F-bristles variable. 165
165(164) C e l l c u p o p e n (fig. 362, 365) or a b s e n t (fig. 3 6 4 ) ; c r o s s v e i n B M - C u a b s e n t
(cells b m a n d d m f u s e d ) (fig. 362-365). 166
C e l l c u p p r e s e n t , c l o s e d ; c r o s s v e i n B M - C u p r e s e n t or a b s e n t (fig. 3 6 7 - 3 6 9 ) .
168
166(165) O c e l l a r triangle c o n s p i c u o u s l y large (fig. 361,282). V e i n C u A i u s u a l l y w i t h
a flexure h a l f w a y d o w n cell b m + d m (fig. 362); o f t e n v e i n R 4 + 5 a n d s o m e
t i m e s also v e i n M i c u r v i n g t o w a r d t h e c o s t a ; c o s t a w i t h a s u b c o s t a l b r e a k
(fig. 362; i n o n l y a f e w s p e c i e s t h e c o s t a is c o n t i n u o u s ) . P r o e p i s t e r n u m
w i t h a s h a r p anterior carina (fig. 283). Chloropidae
- O c e l l a r triangle n o t c o n s p i c u o u s l y large. V e i n C u A i w i t h o u t f l e x u r e ; c o s t a
w i t h b o t h h u m e r a l a n d s u b c o s t a l b r e a k s (fig. 3 6 4 - 3 6 6 ) . P r o e p i s t e r n u m
w i t h o u t carina. 167
167(166) V e i n R2+3 s h o r t , e n d i n g a b o u t h a l f w a y u p t h e c o s t a ; w i n g b a s e n a r r o w ,
cell c u p a n d alula a b s e n t (fig. 363, 3 6 4 ) . S e c o n d and t h i r d a n t e n n a l s e g
m e n t s w i t h l o n g b r i s t l e s (fig. 363). [Flies o f coastal habitats.]
Nannodastiidae
(In E u r o p e 1 g e n u s , Azorastia, w i t h 3 s p e c i e s . )
- C o m b i n a t i o n o f characters n o t as g i v e n a b o v e . W i n g b a s e rarely n a r r o w ,
cell c u p o p e n or a b s e n t , alula u s u a l l y p r e s e n t ; v e i n r2+3 n e a r l y a l w a y s l o n g
(fig. 365), rarely r e a c h i n g o n l y h a l f w a y u p t h e w i n g . A r i s t a u s u a l l y w i t h
l o n g rays o n t h e u p p e r side o n l y (fig. 366), s o m e t i m e s bare or p u b e s c e n t ;
face u s u a l l y s w o l l e n , c o n v e x . In Ochthera fore f e m u r raptorial (fig. 366).
Ephydridae
168(165) C r o s s v e i n B M - C u a b s e n t (cells b m and d m f u s e d ) . P-bristles c o n v e r g i n g .
S m a l l , s t o u t flies w i t h g r e y i s h , y e l l o w i s h or b r o w n i s h p r u i n o s i t y (fig. 367).
Tethinidae: Pelomyiinae
- C r o s s v e i n B M - C u p r e s e n t (cells b m a n d d m separate) (fig. 368). P-bristles
d i v e r g i n g . L o n g , slender, b l a c k flies (fig. 369). S t r o n g y l o p h t h a l m y i i d a e
(In E u r o p e 1 g e n u s , Strongylophthalmyia, w i t h 2 species).
 373 head and fore
legs Streblidae
(Brachytarsina)

372 wing Streblidae

(Brachytarsina)

9 2
169(81) H e a d a d p r e s s e d t o t h e t h o r a x a n d e y e s w e l l - d e v e l o p e d (fig. 371). Tarsal
c l a w s l o n g , d e n t a t e at t h e b a s e (fig. 370). V e i n s in front p a r t o f w i n g s t r o n g ,
r e m a i n i n g v e i n s w e a k (fig. 371; w i n g m a y b e p r e s e n t or a b s e n t , see c o u p l e t
209). Flies e c t o p a r a s i t i c o n b i r d s a n d m a m m a l s ( n o t o n bats).
Hippoboscidae
H e a d n o t a d p r e s s e d to the t h o r a x . E y e s a n d o c e l l i r u d i m e n t a r y or a b s e n t .
Last tarsal s e g m e n t b r o a d e n e d (fig. 373). W i n g v e n a t i o n as i n fig. 372. Flies
ectoparasitic o n b a t s . Streblidae
(In S o u t h e r n E u r o p e 1 s p e c i e s , Brachytarsinaflavipennis Macquart.)
170(82) M o u t h o p e n i n g s m a l l , its d i a m e t e r u p t o 1 / 4 to 1 / 8 o f t h e w i d t h o f t h e
head; mouthparts minute, rudimentary, in s o m e cases o n l y the s w o l l e n
tip o f t h e p a l p v i s i b l e (fig. 375, 378, 3 7 9 , 382). 171
- M o u t h o p e n i n g large, its d i a m e t e r at l e a s t 1 / 4 o f t h e w i d t h o f the h e a d ;
m o u t h p a r t s w e l l d e v e l o p e d , w e l l v i s i b l e (fig. 4 , 7 , 8, 393, 3 9 4 ) . 175
171(170) Ocelli absent. T a c h i n i d a e : Therobia
Ocelli present 172
172(171) H e a d (fig. 375), t h o r a x , l e g s a n d a b d o m e n w i t h s t o u t b l a c k b r i s t l e s . M e r o n
w i t h a single or d o u b l e r o w o f b r i s t l e s a l o n g its h i n d m a r g i n n e a r t h e p o s
terior spiracle; s u b s c u t e l l u m c o n v e x ( s w o l l e n , b u l g i n g ) (fig. 374).
Tachinidae: Trixa, Trixiceps
H e a d a n d u s u a l l y also l e g s a n d r e m a i n d e r o f b o d y c o m p l e t e l y or a l m o s t
completely devoid o f stout black bristles, in m o s t species covered w i t h
soft h a i r - l i k e b r i s t l e s , g i v i n g t h e flies t h e a s p e c t o f b e e s or b u m b l e b e e s (fig.
376-383). M e r o n largely w h i t e or y e l l o w p u b e s c e n t . S u b s c u t e l l u m v a r i a b l e
(from flat t o c o n v e x ) . 173
173(172) V e i n M i m o r e or l e s s straight a n d e n d i n g b e y o n d t h e w i n g tip (fig. 3 7 6 ) .
Gasterophilidae
(In E u r o p e 1 g e n u s , Gasterophilus, w i t h 6 s p e c i e s . )

376 habitus Gasterophilidae

(Gasterophilus)

9 3
 V e i n M i c u r v i n g a n d e n d i n g near (fig. 377, 380) or in v e i n R 4 + 5 (fig. 381,
383)- 174
174(173) V e i n M i b e y o n d c r o s s v e i n D M - C u g r a d u a l l y c u r v i n g t o w a r d t h e c o s t a , or
p r o c e e d i n g m o r e or less straight; cell r4+5 o p e n or c l o s e d at t h e w i n g m a r
g i n , w i t h o u t a p e t i o l e (fig. 377). E n d s o f t h e p t i l i n a l s u t u r e b e l o w t h e
antennae n o t s t r o n g l y c u r v e d t o w a r d e a c h o t h e r (fig. 378). F e m a l e
o v i p a r o u s a n d p o s s e s s i n g a n o v i p o s i t o r (fig. 377). Hypodermatidae
- V e i n M i s t r o n g l y c u r v i n g or w i t h a n angle t o w a r d t h e c o s t a (fig. 380) or
e n d i n g in v e i n R 4 + 5 m a k i n g cell r4+5 p e t i o l a t e (fig. 381, 383). E n d s o f t h e
p t i l i n a l s u t u r e b e l o w t h e a n t e n n a e s t r o n g l y c u r v e d t o w a r d e a c h o t h e r (fig.
379), e x c e p t in Cephalopina (fig. 382) w h i c h s h o w s a w i n g v e n a t i o n as i n
fig. 383. F e m a l e l a r v i p a r o u s a n d w i t h o u t o v i p o s i t o r . Oestridae

382 head Oestridae (Cephalopina) 383 habitus Oestridae (Cephalopina)

9 5
large bristles i \

384 thorax Anthomyiidae (Pegomyia)

(Winthemia)
392 head Anthomyiidae
fi (Craspedochoeta) (fem<
 175(17) M e r o n w i t h o u t o n e or m o r e r o w s o f b r i s t l e s near its h i n d m a r g i n (fig. 384).
V e i n M i u s u a l l y s t r a i g h t (fig. 391, 3 9 4 , 395), in M u s c i d a e s o m e t i m e s c u r v
i n g g e n t l y or s h a r p l y t o w a r d t h e c o s t a (fig. 396). 176
- M e r o n w i t h b r i s t l e s near t h e h i n d m a r g i n , a r r a n g e d i n o n e (fig. 386) or
several (fig. 387) r o w s , o r i n a s m a l l c l u s t e r (fig. 4 0 8 ) . V e i n M i u s u a l l y c u r v
i n g s h a r p l y f o r w a r d (fig. 3 8 8 , 4 0 9 ) , s o m e t i m e s e n d i n g in v e i n R 4 + 5 m a k
i n g cell r4+5 p e t i o l a t e (fig. 389), M i rarely c u r v i n g g e n t l y (fig. 385) or
a l m o s t straight (fig. 4 0 7 ) . 181
> [A k e y w i t h g o o d figures for t h e f a m i l i e s a n d g e n e r a w h i c h k e y o u t i n c o u
p l e t s 176-183 is g i v e n b y G r e g o r et al. 2002.]
176(175) V e n t r a l surface o f s c u t e l l u m i n t h e m i d d l e w i t h a p a t c h o f fine, p a l e hairs
(fig. 3 8 4 , 390). A n a l v e i n l o n g (fig. 391). F e m a l e o f t e n w i t h i n t e r f r o n t a l
bristles (fig. 392). Anthomyiidae
- V e n t r a l surface o f s c u t e l l u m i n t h e m i d d l e bare, a l t h o u g h i n s o m e c a s e s
bristles m a y b e p r e s e n t laterally. A n a l v e i n v a r i a b l e . Interfrontal b r i s t l e s
p r e s e n t or a b s e n t . 177
177(176) T h e p a r t o f t h e h e a d b e h i n d a n d o b l i q u e l y b e l o w t h e e y e s w i t h a f e w to
m a n y l o n g , p a l e , t h i n hairs (fig. 3 9 3 , 3 9 4 ) ; i n t e r f r o n t a l b r i s t l e s a b s e n t . A n a l
v e i n l o n g , u s u a l l y r e a c h i n g t h e w i n g m a r g i n (fig. 3 9 4 ) . S c a t h o p h a g i d a e
If p r e s e n t , bristles b e h i n d a n d o b l i q u e l y b e l o w t h e e y e s n o t l o n g , p a l e , t h i n
or hair-like. Interfrontal b r i s d e s p r e s e n t o r a b s e n t . A n a l v e i n v a r i a b l e . 178
178(177) A n a l v e i n r e a c h i n g t h e w i n g m a r g i n (fig. 3 9 4 ) , i n s o m e cases v e r y t h i n or
a l m o s t f o l d - l i k e near t h e e n d . 179

393 head Scathophagidae (Scathophaga) 394 habitus Scathophagidae (Spaziphora)

97
398 head Muscidae (Azelia) (female)

98
 A n a l v e i n n o t r e a c h i n g t h e w i n g m a r g i n (fig. 395-397). 180
179(178) K a t e p i s t e r n u m w i t h at least 2 large b r i s t l e s , u s u a l l y 3 (fig. 384). W i n g clear.
F e m a l e o f t e n w i t h i n t e r f r o n t a l b r i s t l e s (fig. 392). Anthomyiidae
K a t e p i s t e r n u m w i t h 1 large bristle (fig. 3 9 4 ) or w i n g d a r k e n e d at t h e b a s e ,
a l o n g t h e c o s t a a n d at t h e tip. Interfrontal b r i s t l e s a b s e n t .
Scathophagidae
180(178) V e i n A 2 c u r v i n g t o w a r d the e n d o f t h e anal v e i n w h i c h it w o u l d m e e t i f it
w e r e e x t e n d e d ; apical p o r t i o n o f v e i n Sc a l m o s t straight; v e i n M i a l m o s t
straight (fig. 395). Interfrontal b r i s t l e s a b s e n t . Fanniidae
- V e i n A 2 a n d the anal v e i n w o u l d n o t m e e t i f b o t h w e r e e x t e n d e d ; apical
p o r t i o n o f v e i n Sc d i s t i n c t l y c u r v i n g (fig. 397); v e i n M i d i s t i n c t l y c u r v i n g
f o r w a r d (fig. 396) or n o t (fig. 397). Interfrontal b r i s t l e s u s u a l l y a b s e n t .
Muscidae
> [ T h e g e n u s Azelia ( M u s c i d a e ) s h o w s a v e n a t i o n s i m i l a r t o t h a t o f
F a n n i i d a e w i t h r e s p e c t to A 2 a n d t h e anal v e i n ; t h e f e m a l e o f Azelia p o s
s e s s e s i n t e r f r o n t a l bristles (fig. 398), w h i c h in t h e F a n n i i d a e are a l w a y s
absent.]
181(175) S u b s c u t e l l u m strongly d e v e l o p e d , inflated (convex), closely adpressed to
the scutellum (membrane b e t w e e n scutellum and subscutellum distinct
ly n a r r o w e r t h a n t h e s u b s c u t e l l u m itself; fig. 3 9 9 , 374). L o w e r c a l y p t e r
u s u a l l y r o u n d e d - t r i a n g u l a r , r a r e l y e l l i p t i c a l (Cinochira, Catharosia).
P o s t e r i o r spiracle w i t h o u t l o n g hairs o n anterior l a p p e t (fig. 4 0 2 ) .
Tachinidae
> [Vein M i u s u a l l y c u r v i n g or g e n i c u l a t e t o w a r d t h e c o s t a (fig. 4 0 5 ) or e n d
i n g i n v e i n R 4 + 5 (cell r4+5 p e t i o l a t e , fig. 4 0 6 ) ; rarely faintly c u r v i n g ,
straight or n o t r e a c h i n g t h e w i n g m a r g i n . ]
S u b s c u t e l l u m a b s e n t or w e a k l y d e v e l o p e d (fig. 4 0 1 ) ; if s o m e w h a t i n f l a t e d ,
t h e n s u b s c u t e l l u m a b o u t as h i g h as t h e m e m b r a n e b e t w e e n s c u t e l l u m a n d
s u b s c u t e l l u m (fig. 4 0 0 ) a n d p o s t e r i o r spiracle w i t h l o n g h a i r - l i k e b r i s t l e s
a l o n g t h e a n t e r i o r m a r g i n (fig. 4 0 3 , 4 0 4 ) . 182

9 9
405 habitus Tachinidae (Ectophasia) 406 habitus Tachinidae (Cylindromyia)

407 habitus Muscidae (Eginia)

411 habitus
Calliphoridae
(Melanomya)

posterior spiracie
meral bristles
WM
408 meron bristles Muscidae (Eginia)

V Jak. ?
ff$$8 412 posterior spiracle
Rhinophoridae
(Melanophora)

409 wing Calliphoridae (Lucilia)

anterior lappet

posterior lappet

410 wing Rhinophoridae (Stevenia)

413 posterior spiracle Calliphoridae (Phumosia)

182(181) V e i n M i s l i g h t l y c u r v i n g b a c k w a r d (fig. 4 0 7 ) . M e r a l b r i s t l e s a r r a n g e d i n a
s m a l l cluster (fig. 4 0 8 ) . M u s c i d a e : Eginia
> [ A b d o m e n y e l l o w i s h to o r a n g e . L o w e r c a l y p t e r s m a l l , elliptical; anal v e i n
c o n t i n u e s u p t o j u s t b e f o r e t h e w i n g m a r g i n . S u b s c u t e l l u m absent; p o s t e
rior spiracle w i t h o u t l a p p e t s , w i t h b r i s t l e s a l o n g t h e anterior margin.]
- V e i n M i c u r v i n g t o w a r d t h e c o s t a g e n t l y (fig. 385, 411) or g e n i c u l a t e (fig.
4 0 9 ) or e n d i n g i n v e i n R 4 + 5 (cell r4+5 p e t i o l a t e , fig. 4 1 0 ) . 183
183(182) M e r o n w i t h a r o w o f delicate bristles a l o n g its h i n d m a r g i n , in s i z e a n d
t h i c k n e s s similar t o t h e b r i s t l e s a l o n g t h e u p p e r m a r g i n o f t h e m e r o n . V e i n
M i c u r v i n g g e n t l y t o w a r d t h e costa. H a b i t u s as i n fig. 385.
M u s c i d a e : Graphomya
> [ B o d y pale w i t h g r e y p r u i n o s i t y , t h o r a x w i t h dark l o n g i t u d i n a l s t r i p e s ,
a b d o m e n c h e c k e r e d w i t h r o u n d to o v a l s p o t s in t h e m i d d l e (fig. 385).]
M e r o n bristles s t r o n g , a r r a n g e d i n o n e (fig. 386, 399) or s e v e r a l (fig. 387)
rows. Vein M i usually c u r v i n g s h a r p l y t o w a r d t h e c o s t a or g e n i c u l a t e (fig.
4 0 9 ) or e n d i n g i n v e i n R 4 + 5 (cell H + 5 p e t i o l a t e , fig. 4 1 0 ) , o n l y i n f e w c a s e s
gently curving toward t h e costa (fig. 411). 184
> [A k e y w i t h g o o d figures for t h e T a c h i n i d a e a n d t h e f a m i l i e s a n d g e n e r a
w h i c h k e y o u t i n c o u p l e t s 1 8 4 - 1 9 0 is g i v e n b y T s c h o r s n i g & H e r r i n g 1 9 9 4
(in G e r m a n ) . ]
184(183) P o s t e r i o r spiracle w i t h a l i n i n g o f l o n g hair-like b r i s t l e s , w i t h o u t l a p p e t s
(fig. 4 0 3 , 4 1 2 ) . L o w e r c a l y p t e r o v o i d , s p o o n - s h a p e d , e r e c t , l a t e r a l l y
e x t e n d e d (fig. 4 1 4 ) . S u b s c u t e l l u m m a y b e s w o l l e n or n o t . 185
P o s t e r i o r spiracle at least partially c o v e r e d b y an anterior a n d / o r p o s t e r i o r
l a p p e t , the latter in the f o r m o f a l o c k i n g plate ( o p e r c u l u m ) (fig. 4 1 3 ) .
L o w e r c a l y p t e r variable: u s u a l l y large, triangular, in s o m e cases s m a l l a n d
s p o o n - s h a p e d . S u b s c u t e l l u m n a r r o w , flat, n o t s w o l l e n (fig. 4 1 7 ) . 186
185(184) S u b s c u t e l l u m d i s t i n c t l y s w o l l e n ( c o n v e x ) b u t n o t s t r o n g l y d e v e l o p e d , at
m o s t h a l f as d e e p as h i g h (fig. 4 1 5 ) . Pdiinophoridae
> [Cell r4+5 o f t e n p e t i o l a t e (fig. 4 1 0 ) . In s o m e c a s e s , b r i s t l e s p r e s e n t o n t h e
g e n a , a l o n g s i d e t h e l o w e r e y e m a r g i n (fig. 416).]

101
 420a habitus Sarcophagidae (Sarcophaga)

420b wing Sarcophagidae (Sphenometopa)

421 thorax Calliphoridae (spec.)

1O 2
 S u b s c u t e l l u m d i s t i n c t l y flat, n o t s w o l l e n (fig. 4 1 7 ) .
Calliphoridae: Morinia, Melanomya, Angioneura
> [Cell r4+5 o p e n or c l o s e d at t h e w i n g m a r g i n , rarely w i t h a s h o r t p e t i o l e
(fig. 4 1 8 ) . N o bristles o n t h e g e n a near t h e l o w e r e y e m a r g i n . B o d y b l a c k or
b l a c k w i t h g r e y p r u i n o s i t y , n o stripes o n u p p e r side o f t h o r a x a n d n o
checkered pattern o n the abdomen.]
186(184) Flies largely g r e e n or b l u e m e t a l l i c l u s t r o u s . Calliphoridae
- C o l o u r v a r y i n g f r o m y e l l o w t o b l a c k , b u t w i t h o u t m e t a l l i c g r e e n or b l u e
lustre. 187
187(186) P r e s u t u r a l bristle (PS) i n s e r t e d m o r e o u t w a r d t h a n p o s t h u m e r a l bristle
(PH) (fig. 4 1 9 ) or p o s t h u m e r a l bristle a b s e n t . N o t o p l e u r o n w i t h 2 b r i s t l e s ,
u s u a l l y i n t e r s p e r s e d w i t h 1 or 2 s m a l l e r bristle(s) (fig. 4 1 9 ) . P r o e p i s t e r n u m
u s u a l l y bare. 188
Presutural bristle (PS) i n s e r t e d m o r e i n w a r d t h a n p o s t h u m e r a l bristle
(PH) (fig. 4 2 1 , 4 2 2 ) . In s o m e g e n e r a , t h e p r e s u t u r a l b r i s t l e (PS) a n d
p o s t h u m e r a l bristle (PH) i n s e r t e d at t h e s a m e l e v e l (fig. 423) i n w h i c h case
t h e p r o e p i s t e r n u m is s e t w i t h hairs. N o t o p l e u r o n w i t h 2 bristles (fig. 4 2 1 -
423). Calliphoridae
188 (187) Flies n o t c o m p l e t e l y b l a c k a n d l u s t r o u s . Sarcophagidae
> [Often gena w i t h bristles b e l o w the e y e . M o s t species grey w i t h black
markings consisting o f stripes o n the thorax and a checkered pattern o n
the a b d o m e n (fig. 420a) a n d w i t h a f o l d or a p p e n d i x at t h e c o r n e r o f v e i n
M i (fig. 420b).]
- L u s t r o u s b l a c k flies. 189
189(188) Flies o v e r 7 m m b o d y l e n g t h . V e i n M i g e n i c u l a t e t o w a r d R 4 + 5 , c a r r y i n g a
fold or a p p e n d i x at t h e c o r n e r ; v e i n M i e n d i n g i n t h e c o s t a near v e i n R 4 + 5
(fig. 4 2 0 b ) or i n v e i n R 4 + 5 (cell 14+5 w i t h a s h o r t p e t i o l e ) .
Sarcophagidae
S m a l l flies (2-3 m m ) . V e i n M i r e g u l a r l y c u r v i n g t o w a r d v e i n R 4 + 5 , w i t h
o u t fold or a p p e n d i x ; cell r4+5 o p e n or w i t h a l o n g p e t i o l e (as i n fig. 4 1 0 ) .
190
190(189) C e l l r4+5 w i t h a l o n g p e t i o l e (as i nfig.4 1 0 ) .
Tachinidae: Litophasia hyalipennis (Fallen)
C e l l r4+5 o p e n . Calliphoridae: Angioneura acerba ( M e i g e n )

PH PS notopleuron

422 detail thorax 423 detail thorax 424 leg Cecidomyiidae (Frirenia)
Calliphoridae Calliphoridae
(Protophormia (Protophormia
terraenovae) atriceps)

1o3
428 head Pediciidae (Tricyphona) 432 thorax Pediciidae (Tricyphona)
191(1) A n t e n n a u s u a l l y l o n g , f i l i f o r m or r e s e m b l i n g a s t r i n g o f b e a d s , w i t h 6 or
( u s u a l l y ) m o r e s e g m e n t s ; u s u a l l y all s e g m e n t s or t h o s e b e y o n d t h e s e c o n d
m o r e or less s i m i l a r (fig. 4 2 6 , 4 3 7 ) ; i f less t h a n 6 s e g m e n t s (4 s e g m e n t s i n
fig. 428), t h e n o c e l l i a b s e n t a n d l e g s a n d a b d o m e n l e n g t h e n e d (fig. 4 2 9 ) .
Palp u s u a l l y w i t h 3-5 s e g m e n t s . Nematocera - 1 9 2
A n t e n n a n o t f i l i f o r m or r e s e m b l i n g a s t r i n g o f b e a d s , w i t h less t h a n 6 s e g
m e n t s ; u s u a l l y t h e first 2 s e g m e n t s s h o r t , t h e third s e g m e n t large a n d t h e
s u b s e q u e n t s e g m e n t s m a k i n g u p an arista or s t y l e . P a l p u s u a l l y w i t h 1-2
segments. Brachycera - 206
192(191) O c e l l i a b s e n t (fig. 4 2 6 , 4 2 8 ) . 193
- Ocelli present. 199
193(192) Tarsi w i t h less t h a n 5 s e g m e n t s , o r first tarsal s e g m e n t far s h o r t e r t h a n s e c
o n d s e g m e n t (fig. 4 2 4 ) . S m a l l , delicate i n s e c t s , g e n e r a l l y w i t h c o n s p i c u
ously long antennae. Cecidomyiidae
- First tarsal s e g m e n t n o t d i s t i n c t l y s h o r t e r t h a n t h e s e c o n d ; t a r s u s w i t h 5
segments. 194
194(193) U p p e r side o f t h o r a x w i t h a V - s h a p e d t r a n s v e r s e s u t u r e (fig. 425). 195
U p p e r side o f t h o r a x w i t h o u t V - s h a p e d t r a n s v e r s e s u t u r e . 197
195(194) T e r m i n a l (5th) s e g m e n t o f p a l p e l o n g a t e , d i s t i n c t l y l o n g e r t h a n t h e p r e
c e d i n g s e g m e n t s ; r o s t r u m w e l l d e v e l o p e d , w i t h o u t (fig. 4 2 6 ) or w i t h
n a s u s (fig. 22). B o d y l e n g t h g e n e r a l l y o v e r 8 m m (fig. 4 2 7 ) . Tipulidae
T e r m i n a l (5th) p a l p a l s e g m e n t s h o r t ; r o s t r u m s h o r t , w i t h o u t n a s u s (fig.
4 2 8 ) ; b o d y l e n g t h u s u a l l y less t h a n 8 m m (fig. 4 2 9 - 4 3 1 ) . 196
196(195) E y e p u b e s c e n t , w i t h o m m a t r i c h i a i n b e t w e e n t h e facets (fig. 428).
Pediciidae
E y e bare. Limoniidae
197(194) U p p e r side o f t h o r a x w i t h r e m n a n t s o f a V - s h a p e d t r a n s v e r s e s u t u r e (fig.
432). 198
- U p p e r side o f t h o r a x w i t h o u t r e m n a n t s o f t h i s s u t u r e ( e x a m p l e s o f h a b i t u s
a n d w i n g v e n a t i o n cf. fig. 4 3 3 , 4 3 4 ) . Chironomidae
198(197) E y e p u b e s c e n t , w i t h o m m a t r i c h i a i n b e t w e e n t h e facets (fig. 428).
Pediciidae
E y e bare. Limoniidae
199(192) E y e s c o n n e c t e d b y a n e y e b r i d g e a b o v e t h e a n t e n n a e , or t o u c h i n g e a c h
o t h e r (fig. 58,73). 200
- E y e s separate. 202

433 copula Chironomidae (Clunio) 434 wing Chironomidae (Telmatogeton)

105
435 habitus Sciaridae (Epidapus)

436 antenna Scatopsidae

(Swammerdamella)

440 habitus Braulidae

(Braula)

437 habitus Cecidomyiidae (Aprionus)

441 habitus
438 ventral view head and thorax
Nycteribiidae
Hippoboscidae (Melophagus)
(Penicillidia)
dorsal view

i o6
200(199) T i b i a e o f m i d a n d h i n d l e g s w i t h apical bristles (fig. 4 3 5 ) . Sciaridae
T i b i a e o f m i d a n d h i n d l e g s w i t h o u t apical b r i s d e s . 201
201(200) A n t e n n a l segments m o r e or less c o m p a c t , w i t h a strong terminal s e g m e n t
(fig. 4 3 6 ) . A b d o m e n o f e v e n w i d t h t h r o u g h o u t o r e v e n s l i g h t l y w i d e r t o
t h e rear. Scatopsidae
> [The o n l y s c a t o p s i d i n E u r o p e w i t h r u d i m e n t a r y w i n g s is Thripomorpha
paludicola Enderlein.]
- A n t e n n a l s e g m e n t s fragile, c a r r y i n g a r i n g o f l o n g , t h i n v e r t i c i l s basally.
A b d o m e n t a p e r i n g t o a p o i n t (fig. 4 3 7 ) . Cecidomyiidae
202(199) T i b i a e o f m i d a n d h i n d l e g s w i t h o u t apical b r i s t l e s . Cecidomyiidae
T i b i a e o f m i d a n d h i n d l e g s w i t h apical b r i s t l e s (fig. 4 3 5 ) . 203
203(202) A n t e n n a m o r e o r less as l o n g as h e a d a n d t h o r a x t o g e t h e r ; h e a d a n d t h o r a x
black, a b d o m e n b l a c k i s h b r o w n , s i z e 4 - 5 m m , w i n g d i s t i n c t l y s h o r t e n e d
(female o f Hesperinus imbecillus M e i g e n ) . Hesperinidae
C o m b i n a t i o n o f characters n o t as g i v e n a b o v e . 204
204(203) A n t e n n a inserted halfway u p the eye, or higher.
Sciaridae ( & M y c e t o p h i l i d a e ? )
> [ T h e u n k n o w n f e m a l e o f t h e m y c e t o p h i l i d Baeopterogyna mihalyii
M a t i l e , w h i c h is b e l i e v e d t o b e s h o r t - w i n g e d , m i g h t k e y o u t here.]
- A n t e n n a i n s e r t e d at l o w e r p a r t o f t h e h e a d , u s u a l l y j u s t a b o v e t h e l o w e r
m a r g i n o f the face. 205
205(204) V e i n R forked (fig. 67). Pleciidae
> [ S o m e w h a t s h o r t - w i n g e d m a l e o f Penthetriafunebris M e i g e n , fig. 68.]
Vein R n o t forked or w i n g absent. Sciaridae
206(191) C o x a e o f m i d l e g s , a n d u s u a l l y o f t h e o t h e r l e g s as w e l l , far apart (fig. 4 3 8 ,
441). 207
C o x a e o f m i d l e g s n o t c o n s p i c u o u s l y w i d e apart, s o m e t i m e s e v e n t o u c h
i n g e a c h other. 210
207(206) Tarsal c l a w s t r a n s f o r m e d i n t o a c o m b - l i k e s t r u c t u r e , p o i n t i n g i n w a r d (fig.
4 3 9 , 4 4 0 ) . E y e r e d u c e d t o a s p o t b e h i n d t h e s u n k e n a n t e n n a ; ocelli, h a l -
teres a n d s c u t e l l u m a b s e n t . T h o r a x s m a l l , m o r e o r less s i m i l a r t o t h e first
a b d o m i n a l s e g m e n t s (fig. 4 4 0 ) . Flies l i v i n g e c t o p a r a s i t i c o n b e e s .
Braulidae
(In E u r o p e 1 g e n u s , Braula, w i t h 3 s p e c i e s . )
C o m b i n a t i o n o f characters n o t as g i v e n a b o v e . If t h o r a x s i m i l a r t o t h e first
a b d o m i n a l s e g m e n t s , t h e n t h e e y e s w e l l d e v e l o p e d (e.g. fig. 4 4 5 ) . 208
208(207) Head small, in resting position bent backward o n to thorax. Eyes and ocel
li s m a l l o r a b s e n t . L e g s l o n g , w i t h s w o l l e n f e m o r a a n d tibiae; first tarsal
s e g m e n t at least as l o n g as all o t h e r tarsal s e g m e n t s t o g e t h e r (fig. 4 4 1 ) . Flies
l i v i n g e c t o p a r a s i t i c o n bats. Nycteribiidae

107
 1st tarsal segment
of hind-leg

447 habitus Sphaeroceridae

(Aptilotus)

442 habitus Hippoboscidae

(Crataerina)

446 habitus Hybotidae

(Ariasella)

wing cast off at some

distance from base 448 habitus Chloropidae
(Tricimba)

443 habitus Hippoboscidae

(Melophagus)
m
449 habitus Carnidae
(Carnus) (female)
450 head Ephydridae
(Rhynchopsilopa)

eyes rudimentary 444 head and fore

or absent legs Streblidae
(Brachytarsina)

445 habitus Phoridae (Aenigmatias) 451 habitus Anthomyzidae (Stiphrosoma)

l o 8
 Head not bent b a c k w a r d o n the thorax. C o m b i n a t i o n o f other characters
n o t as g i v e n a b o v e . Flies l i v i n g e c t o p a r a s i t i c o n b i r d s a n d m a m m a l s
( H i p p o b o s c i d a e n o t o n bats; S t r e b l i d a e o n bats o n l y ) . 209
209(208) H e a d a d p r e s s e d t o t h e t h o r a x a n d e y e s w e l l - d e v e l o p e d . Tarsal c l a w s l o n g ,
d e n t a t e at t h e b a s e (fig. 4 4 2 , 4 4 3 ) . Flies l i v i n g as e c t o p a r a s i t e s o n b i r d s a n d
m a m m a l s ( e x c e p t bats). Hippoboscidae
H e a d n o t a d p r e s s e d to the t h o r a x . E y e s a n d o c e l l i r u d i m e n t a r y or a b s e n t .
Last tarsal s e g m e n t b r o a d e n e d (fig. 4 4 4 ) . Flies e c t o p a r a s i t i c o n b a t s .
Streblidae
210(206) A n t e n n a s e e m s t o c o n s i s t o f a s i n g l e s p h e r i c a l s e g m e n t , w i t h a l o n g arista.
W i n g s r e d u c e d or w i n g s a n d h a l t e r e s a b s e n t (fig. 4 4 5 ) . Phoridae
Antenna distinctly w i t h 3 segments. 211
211(210) T h i r d a n t e n n a l s e g m e n t e n d i n g m o r e or less a c u t e ; p t i l i n a l s u t u r e a b s e n t .
212
T h i r d a n t e n n a l s e g m e n t o v o i d or r o u n d e d ; p t i l i n a l s u t u r e p r e s e n t . 213
212(211) M o u t h p a r t s l o n g . D o r s a l side o f h e a d n o t c o n c a v e b e t w e e n t h e e y e s
( e x a m p l e o f h a b i t u s : fig. 4 4 6 ) . Hybotidae
M o u t h p a r t s s h o r t ; h e a d w i t h dorsal part, b e t w e e n e y e s , c o n c a v e .
Dolichopodidae
213(211) First tarsal s e g m e n t o f h i n d l e g s w o l l e n , u s u a l l y s h o r t e r t h a n t h e s e c o n d
tarsal s e g m e n t (fig. 4 4 7 ) . Sphaeroceridae
- First tarsal s e g m e n t o f h i n d l e g n o t s h o r t e r t h a n t h e s e c o n d tarsal s e g m e n t .
214
214(213) Bristles o n t h e h e a d w e a k l y d e v e l o p e d or a b s e n t . Chloropidae
- Bristles o n t h e h e a d n o r m a l l y d e v e l o p e d . 215
215(214) O c e l l a r triangle s t r i k i n g l y large; 1 pair o f interfrontal b r i s t l e s i m m e d i a t e l y
a b o v e the a n t e n n a e . W i n g cast off at s o m e d i s t a n c e f r o m t h e b a s e . H a b i t u s
fig. 4 4 9 . Carnidae
O c e l l a r triangle n o t c o n s p i c u o u s l y large. 216
216(215) T i b i a e w i t h o u t dorsal preapical bristle. 217
T i b i a e w i t h dorsal preapical b r i s t l e . 219
217(216) Face d i s t i n c t l y s w o l l e n , c o n v e x ; i f arista w i t h l o n g r a y s , t h e n o n l y o n t h e
u p p e r side (fig. 4 5 0 ) . Ephydridae
Face n o t d i s t i n c t l y c o n v e x ; i f arista w i t h l o n g r a y s , t h e n o n b o t h s i d e s
(plumose). 218
218(217) A t least 2 pairs o f F - b r i s t l e s p r e s e n t (fig. 4 5 1 ) . Anthomyzidae
- O n l y 1 pair o f F - b r i s t l e s p r e s e n t (fig. 360). Opomyzidae
219(216) S u b s c u t e l l u m w e l l d e v e l o p e d , i n f l a t e d (fig. 399). Tachinidae
S u b s c u t e l l u m n o t inflated. 220
220(219) A r i s t a p l u m o s e (fig. 218). Drosophilidae
A r i s t a bare or p u b e s c e n t o n l y (fig. 1 0 ) . Heleomyzidae

109
FAMILY DESCRIPTIONS

C A T = A l t h o u g h they d o n o t contain k e y s , t h e identification references include recent cata

logues as valuable source o n genera, species, distribution and references.
C M P D = C o n t r i b u t i o n s to a Manual o f Palaearctic Diptera.
Lindner = Chapter in Lindner, E., D i e Fliegen der Palaarktischen R e g i o n .
( ) Family n a m e s b e t w e e n brackets refer to n a m e s as found in t h e literature, n o t recognised
here as a separate family b u t , as indicated, considered part o f another family,
et al. References w i t h m o r e than t w o authors are g i v e n as First author et al.

A s far as n o t y e t outdated, the n u m b e r of genera and species in Europe is largely based o n the
Catalogue o f Palaearctic Diptera, the C M P D and Fauna Europaea, t h e latter available online
at: w w w . f a u n a e u r . o r g (consulted w a s version 1.2, updated 7 March 2005).

A s to size, the f o l l o w i n g categories are distinguished: m i n u t e : smaller than 2 m m ; small: 2-

5 m m ; m e d i u m sized: 5-10 m m ; large: 10-20 m m ; v e r y large: o v e r 20 m m .

Acartophthalmidae (key couplet 113;fig.243)

Systematics: Acalyptrate Brachycera; superfamily
O p o m y z o i d e a ; in Europe 1 g e n u s , Acartophthalmus, w i t h 3
species.
Characters: M i n u t e to small (1-2.5 m m ) , b r o w n i s h grey flies.
Arista pubescent, ocelli present; Oc-bristles present; P-bris-
tles strong, far apart, diverging; 3 pairs o f F-bristles, c u r v i n g
obliquely o u t - b a c k w a r d , increasing in size, the u p p e r pair the
largest; scattered interfrontal setulae present; vibrissae absent
b u t w i t h a series o f strong bristles near the vibrissal angle.
W i n g u n m a r k e d or tinged along costa; costa w i t h a humeral
break o n l y ; v e i n Sc complete; crossvein B M - C u present; cell
c u p closed. Tibiae w i t h o u t dorsal preapical bristle.
Biology: T h e larvae o f this small family are p r e s u m e d to feed o n
rotting organic matter. T h e adults appear to inhabit w o o d l a n d
and are found o n rotting bracket fungi and m u s h r o o m s , decay
ing w o o d , d r o p p i n g s and carrion; d e p o s i t i o n o f eggs o n a dead 452 Acartophthalmus nigrinus
snail and carrion has b e e n o b s e r v e d . (Zetterstedt), female; Papp 1978
Identification references: O z e r o v 1986 ( r e v i e w ) ; Stackelberg
1989a! (former U S S R ) ; C M P D : Papp 8t O z e r o v 1998; C A T : Papp 19846.

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Acroceridae (key couplet 45; fig. 98-99)
Systematics: L o w e r m u s c o m o r p h B r a c h y c e r a ; s u p e r f a m i l y
Nemestrinoidea; i n Europe 8 genera and about 35 species.
Characters: D e s p i t e variability in size (2.5-20 m m ) , colour and
w i n g venation, these flies are easily recognised b y their small head
situated almost b e l o w the h u n c h b a c k e d thorax, the large ear-
shaped l o w e r calypter and large a b d o m e n . B o d y colour dull or l u s
trous, usually black w i t h w h i t e , y e l l o w , or orange markings o n
thorax and a b d o m e n . T h e small head largely o c c u p i e d b y the rela
tively large eyes, holoptic in b o t h sexes. W i n g clear or tinged;
venation v a r y i n g from complete to strongly reduced. Legs simple,
femora s o m e t i m e s s w o l l e n ; e m p o d i u m p u l v i l l i f o r m .
Biology: T h e larvae are endoparasitoids o f spiders, usually o n e larva
per spider, rarely t w o or m o r e . T h e spider continues functioning 4 5 ii paniae Lamarck,
3 A s t o m e a niS

normally until just prior to c o m p l e t i o n o f the fourth instar. A t the male; Sack 1936
end o f this stage the spider is largely eaten and the larva pupates
outside the spider in the w e b the spider has w o v e n at the v e r y last. T h e r e are apparently
no fixed host-parasitoid relationships at the species level. A d u l t s m a y m o r e or less a b o u n d
locally b u t apart from species visiting flowers and feeding o n nectar, they are rarely
observed. T h e y are m a i n l y active o n w a r m s u n n y days. Males m a y s o m e t i m e s gather in
large n u m b e r s at higher points in the landscape (hilltopping). M a t i n g takes place in flight.
Identification references: C h v a l a 1980 (Central Europe); V a n der G o o t 1963, D e Jong et al.
2000 (Netherlands), N a r t s h u k l g S g f (former U S S R ) ; O l d r o y d 1969; S t u b b s 8s. Drake 2001
(British Isles); W e i n b e r g & Bachli 1997 (Switzerland); C M P D : Nartshuk 1997; C A T :
Nartshuk 1988.

(Aenigmatiidae): part o f the Phoridae.

Agromyzidae (key couplet n o , 123; fig. 241-242,263-266)

Systematics: Acalyptrate Brachycera; superfamily O p o m y z o i d e a ;
in Europe s o m e 23 genera and about 910 species.
Characters: M i n u t e to m e d i u m sized (1-6 m m ) flies, v a r y i n g in
colour from all y e l l o w to black to metallic green. A r i s t a bare to
pubescent; ocelli present; Oc-bristles present; P-bristles diverg
ing; 2-8 pairs o f F-bristles, the l o w e r 1-3 pairs c u r v i n g i n w a r d , the
other pairs b a c k w a r d ; interfrontal bristles absent; interfrontal
setulae s o m e t i m e s present; vibrissae present b u t i n s o m e cases
w e a k . W i n g u n m a r k e d ; costa w i t h subcostal break; v e i n Sc c o m
plete or incomplete, apically e n d i n g in v e i n R i ( A g r o m y z i n a e ) or
separate from v e i n R i b u t reduced to a fold that m a y or m a y n o t
reach the costa ( P h y t o m y z i n a e ) ; crossvein B M - C u present; cell
cup closed. Tibiae w i t h o u t dorsal preapical bristle. Female w i t h
oviscape, n o n retractable basal s e g m e n t o f the ovipositor.
Biology: T h e larvae are m i n i n g , generally i n the stalks or leaves o f 454 Mines of the larva of Aulagromyza
uli
herbs, s o m e t i m e s in roots, seeds or under bark. In the f e w species PP (Kaltenbach) on Populus; Hering
1 9 5 7
living o n w o o d y plants this m a y lead to gall formation. M o s t
species are m o r e or less h o s t specific and several species cause
damage to e c o n o m i c a l l y important plants. T h e shape of the m i n e is often characteristic o f
the species and therefore useful for identification. A d u l t s occur in a v a r i e t y o f habitats,
d e p e n d i n g o n the larval h o s t plants.
Identification references: D e m p e w o l f 2 0 0 4 ( W o r l d ) ; H e r i n g 1 9 5 7 ( m i n e s ) ; E . B .
R o h d e n d o r f 1989 (former U S S R ) ; Spencer 1972 (British Isles), 1976 ( N o r t h w e s t e r n
Europe), 1987 (North A m e r i c a ) ; C M P D : D a r v a s 8t Papp 2000 (genera); C A T : Papp i g 8 4 d
(see also Spencer & Martinez 1987).

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Anisopodidae (key couplet 38; fig. 38, 84)
Systematics: N e m a t o c e r a ; superfamily A n i s o p o d o i d e a ; tradi
tionally this family includes the Mycetobiidae; in Europe 1
g e n u s , Sylvicola, w i t h about 10 species.
Characters: Small to large (4-12 m m ) , y e l l o w i s h to b r o w n i s h
N e m a t o c e r a w i t h slender antennae and a b d o m e n . Head small
and rounded; m o u t h p a r t s small; ocelli present; e y e s dichoptic
or holoptic; antenna w i t h 16 s e g m e n t s v a r y i n g from relatively
short to longer than head and thorax together. W i n g w i d e ,
w i t h clear anal lobe and a pattern o f smaller markings. Legs
slender, relatively long; tibiae w i t h apical spurs.
Biology: T h e larvae live in, and feed o n , all sorts o f rotting and
d e c o m p o s i n g material like d r o p p i n g s , d u n g , vegetables, d y i n g
w o o d , cracks and fissures in trees filled w i t h organic matter, 455 Sylvicola fenestralis (Scopoli),
bark, bracket fungi and m u s h r o o m s , leaves, roots, sap e x u d i n g female; Seguy 1940
from tree w o u n d s and other liquids rich in organic matter. T h e
adults are encountered m a i n l y in s u m m e r in w o o d l a n d , m e a d o w s , gardens, etc. T h e y are
found near the larval habitats, visiting flowers, near sap e x u d i n g from tree w o u n d s , and
are regularly seen o n w i n d o w s , b o t h indoors and outside, or other vertical surfaces such
as tree trunks, w a l l s , etc. Males form s w a r m s to attract females for mating.
Identification references: H a e n n i 1997c (Switzerland); K r i v o s h e i n a 8t M e n z e l 1998
(review, k e y to species); Michelsen 1999 (systematics); S0H
1992 ( N o r w a y ) ; Stackelberg i 9 8 g n (former U S S R ) ; C M P D :
K r i v o s h e i n a 1997c; C A T : K r i v o s h e i n a 1986c.

Anthomyiidae (key couplet 1 7 6 , 1 7 9 ; fig. 384,390-392)

Systematics: Calyptrate Brachycera; superfamily Muscoidea;
s o m e 35 genera and about 4 8 0 species in Europe.
Characters: Small to large (4-12 m m ) , slender flies, usually
coloured b r o w n i s h , greyish or blackish, s o m e t i m e s y e l l o w i s h
or w i t h a distinct pattern o n thorax and a b d o m e n , n e v e r
metallic lustrous. Arista bare to p l u m o s e ; female often w i t h
interfrontal bristles. W i n g generally clear, s o m e t i m e s tinged,
rarely marked; v e n a t i o n virtually a l w a y s w i t h straight v e i n M i 456 Anthomyia pluvialis (Linnaeus),
and l o n g anal v e i n . S c u t e l l u m u n d e r n e a t h in the middle usual male; Seguy 1951
ly w i t h a patch o f fine, pale hairs. M e r o n d e v o i d of bristles near
h i n d margin.
Biology: T h e larvae o f m a n y genera are p h y t o p h a g o u s , s o m e o f w h i c h are leaf miners. A t
the same time m a n y species are saprophagous in diverse habitats (dung, nests o f birds,
etc.). P h y t o p h a g o u s species can be o f e c o n o m i c importance, like various species of Delia
causing damage to o n i o n s , beans and other vegetables. S o m e species are saprophagous, or
w i t h larvae l i v i n g in m u s h r o o m s or as parasitoids o f insects.
T h e larvae o f Fucellia live at the coast in rotting organic matter
w a s h e d u p b y the sea. T h e adults visit flowers b u t can b e found
o n e x u d i n g tree sap and o n d u n g , carrion, rotting plants, etc.
Identification references: E l b e r g i g S g (former U S S R ) ; H e n n i g
1966-1976 (Lindner); C M P D : S u w a 8t D a r v a s 1998; C A T :
D e l y - D r a s k o v i t s 1993.

Anthomyzidae (key couplet 1 0 5 , 1 2 5 , 218; fig. 232-235, 267-

269,451)
Systematics: Acalyptrate Brachycera; superfamily
O p o m y z o i d e a ; 9 genera and about 30 species in Europe.
Characters: M i n u t e to small (1.3-4.5 m m ) , slender flies, y e l l o w
to black, s o m e t i m e s w i t h dark spots or stripes; w i n g usually
l o n g and narrow. Arista pubescent or w i t h longer rays; ocelli 457 Typhamyza bifasciata (Wood),
present; O c - b r i s t l e s present; P-bristles small, c o n v e r g i n g , male; Rohacek 1992

112
 rarely absent; 2-3 pairs o f F-bristles, c u r v i n g backward, usually preceded b y 1 or m o r e
weaker bristles; interfrontal bristles absent or present; vibrissae present. W i n g usually
clear, s o m e t i m e s marked; v e i n C w i t h subcostal break; v e i n Sc incomplete; crossvein B M -
C u present b u t s o m e t i m e s n o t complete; cell c u p closed, rarely o p e n . S o m e genera, includ
ing the more c o m m o n Anthomyza and Paranthomyza, w i t h femur o f fore leg w i t h a short
spine o n the ventral side, b e y o n d the middle; tibiae w i t h o u t dorsal preapical bristle.
Biology: T h e larvae o f m o s t species d e v e l o p in the stems and leaf sheaths o f grasses and
plants in marsh habitats such as species ofTyphaJuncus, Carex and Scirpus w i t h o u t caus
ing m u c h damage; several species are k n o w n to m i n e in d i c o t y l e d o n s w h e r e a s others live
in dead plants or as inquilines in existing galls. T w o species o f Fungomyza d e v e l o p i n the
sporocarps o f fungi. A d u l t s o f m o s t species are usually found in m o i s t habitats, b o t h
shaded and in t h e o p e n such as d a m p m e a d o w s , marshes, b o g s and d a m p d e c i d u o u s or
m i x e d forests w i t h rich u n d e r g r o w t h ; a preference for m o r e dryer habitats such as d r y
grasslands is s h o w n b y s o m e species o f Anthomyza and the b r a c h y p t e r o u s Stiphrosoma
sabulosum (Haliday).
Identification references: A n d e r s s o n 1 9 7 6 , 1984a ( N o r t h w e s t e r n Europe); S o o s 1981
(Central Europe); Rohacek 1 9 9 9 , 2006 (revision); Stackelberg 198930 (former U S S R ) ;
C M P D : Rohacek 1998b; C A T : A n d e r s s o n 1984b.

Asilidae (key couplet 58;fig.93,123-124)

Systematics: L o w e r m u s c o m o r p n B r a c h y c e r a ; s u p e r f a m i l y
Asiloidea; s o m e 77 genera and about 5 4 0 species in Europe.
Characters: Small (3 m m ) to v e r y large (50 m m ) , b u t generally
m e d i u m sized to large (8-20 m m ) flies, v a r y i n g in shape from
long and slender to short and stout. T h e colour of the b o d y and
pubescence are variable from dark metallic lustrous to paler,
w i t h orange or y e l l o w markings o n b o d y and w i n g s , or r e s e m
bling bees and w a s p s . Head dorsally concave i n b e t w e e n the
large, dichoptic eyes; ocelli implanted o n an elevation b e t w e e n
the eyes; antenna variable, third s e g m e n t elongate, usually fol
l o w e d b y 1 or 2 segment(s) w h i c h may, or m a y n o t , constitute
an arista or style; face relatively l o n g and nearly a l w a y s w i t h a
458 Philonicus albiceps (Meigen),
distinct facial k n o b bearing long bristles, n a m e d t h e m y s t a x or
male; Verrall 1909
beard; m o u t h p a r t s p o w e r f u l and piercing. W i n g usually clear,
in s o m e cases c o m p l e t e l y or partially tinged or darkened; cell
m3 and cell c u p o p e n or closed. Legs l o n g and p o w e r f u l , often armed w i t h strong bristles;
tibiae w i t h apical bristles or spurs; e m p o d i u m bristle-like or absent.
Biology: T h e larvae live in the soil and in d e c a y i n g w o o d ; as far as k n o w n they largely feed
on the larvae o f other insects. T h e adults are predators as w e l l . M o s t species prefer o p e n ,
s u n n y habitats and are m a i n l y active during t h e w a r m e s t part o f the day. T h e y are able
fliers that catch their prey - m a i n l y other insects - o n t h e w i n g . T h e y are often seen o n
l o o k o u t posts like w a l l s , branches and leaves, or o n the ground, s u r v e y i n g their s u r r o u n d
ings for possible prey flying by.
Identification references: V o n d e r D u n k 1 9 9 6 ( C e n t r a l
Europe); G e l l e r - G r i m m 2003 ( G e r m a n y ) ; H u l l 1962 (genera);
Van der G o o t 1985 ( N o r t h w e s t e r n Europe); V a n V e e n 1996
(Netherlands); Larsen & Meier 2004 ( D e n m a r k ) ; Lehr 1996
(Asilinae); R i c h t e r 1989b (former U S S R ) ; O l d r o y d 1 9 6 9 ;
Stubbs & Drake 2001 (British Isles); W e i n b e r g 8t Bachli 1995
(Switzerland); C M P D : Majer 1997c; C A T : Lehr 1988.

Asteiidae (key couplet 1 0 4 , 1 2 9 ; fig. 228-229,258,279-281)

Systematics: Acalyptrate Brachycera; superfamily
O p o m y z o i d e a ; 4 genera and about 18 species in Europe.
Characters: M i n u t e to small (1-3 m m ) , delicate, often w e a k l y
sclerotized flies w i t h y e l l o w - b l a c k or dark c o l o u r s , m a l e
almost a l w a y s w i t h distinct interfrontal stripes. Arista s o m e - 4 Asteia amoena Meigen,
5 9

times bare or pubescent b u t usually w i t h a characteristic zigzag female; Papp 1973

113
 o f longer or shorter alternating rays, arista absent in t h e Mediterranean species Asteia
mahunkai Papp; ocelli present; Oc-bristles present or absent; P-bristles w e a k , parallel,
diverging or absent; 0-2 pairs of F-bristles, if present, c u r v i n g forward or b a c k w a r d ; inter-
frontal bristles s o m e t i m e s present i m m e d i a t e l y above the antennae; scattered interfrontal
setulae present; vibrissae absent to present. W i n g u n m a r k e d ; costa continuous; v e i n Sc
incomplete; crossvein B M - C u and cell c u p absent. Tibiae w i t h o u t dorsal preapical bristle.
Biology: T h e larvae h a v e b e e n found under the bark o f trees and have b e e n reared from
flower heads (inflorescences) and rotting or d r y organic matter; larvae o f Leiomyza have
b e e n associated w i t h fungi. T h e adults are often found o n flowers, o n l o w vegetation, near
sap e x u d i n g from tree w o u n d s , and relatively often o n w i n d o w s indoors, or in case of one
g e n u s (Leiomyza) near or o n m u s h r o o m s and bracket fungi i n shaded places. T h e
Mediterranean A. mahunkai is found near the sea, associated w i t h Inula crithmoides
( C o m p o s i t a e ) ; other Mediterranean species (Phlebosotera, s o m e Asteia) are often found
under Tamarix trees. A d u l t s of the species Astiosoma rufifrons D u d a is attracted to w o o d
ash after bonfires, especially during t h e e v e n i n g .
Identification references: C h a n d l e r 1978 (British Isles); M e r z i 9 9 6 d (Northern and
Central Europe); Stackelberg 19893s (former U S S R ) ; C M P D :
Papp 1998c; C A T : Papp 1 9 8 4 b .

Atelestidae (key couplet 7 7 , 7 9 - 8 0 ; fig. 165-166)

Systematics: L o w e r e r e m o n e u r a n B r a c h y c e r a ; s u p e r f a m i l y
Empidoidea; traditionally part o f the Empididae; in Europe 3
genera and 4 species. For a discussion about the p h y l o g e n y and
classification see Sinclair & C u m m i n g 2006.
Characters: M i n u t e to small (1.5-3.5 m m ) , black flies. Head
rather small; e y e s big, holoptic i n male; third antennal s e g m e n t
w i t h an arista o f 1-3 times its o w n length; m o u t h p a r t s relative
ly short, directed forward or s o m e w h a t d o w n . W i n g clear or
tinged, w i t h a stigma spot b e t w e e n apex o f v e i n s S C and R i ,
stigma spot s o m e t i m e s indistinct; cell d m present and v e i n M
forked (Meghyperus), or cell d m absent and v e i n M n o t forked 460 Atelestus pulicarius (Fallen),
(Atelestus, Nemedina); cell c u p n o r m a l l y larger than cells br male; Collin 1961
and b m , if n o t then C u A 2 c o n v e x and anal v e i n arched to w i n g
margin; alula and large anal lobe usually present; cell b m and alula absent in Nemedina.
Tibia of h i n d leg, and in s o m e cases first tarsal s e g m e n t as w e l l , s o m e t i m e s dilated and lat
erally compressed.
Biology: T h e b i o l o g y o f this small family is little k n o w n . T h e larvae are u n k n o w n . A d u l t s
o f Meghyperus and Atelestus m a i n l y occur in grassland or patches of herbs or o n leaves in
d e c i d u o u s w o o d l a n d s , often near w a t e r (in the Netherlands, M. sudeticus L o e w w a s col
lected t w i c e , in b o t h cases near a river: o n a gravel b a n k in the G e u l and o n a sandbar in the
Slinge; V a n A a r t s e n , pers. c o m m . 2005); adults o f Nemedina have b e e n collected along
b r o o k s . B o t h sexes fly in s w a r m s . Because o f the a n a t o m y o f their mouthparts, the adults
are p r e s u m e d to be flower visitors (Sinclair 8t C u m m i n g 2006).
Identification references: C h a n d l e r 1981 (Nemedina); Chvala
1983 ( r e v i s i o n ) ; C o l l i n 1961 (British Isles); G o r o d k o v 8t
K o v a l e v 1 9 8 9 ( f o r m e r U S S R ) ; Sinclair & P a p p 2 0 0 4
(Nemedina); Sinclair & C u m m i n g 2 0 0 6 (classification);
C M P D : n o contribution; C A T : C h v a l a 1989b.

Athericidae (key couplet 4 8 ;fig.9 5 , 1 0 5 - 1 0 6 )

Systematics: L o w e r Brachycera; s u p e r f a m i l y Tabanoidea; 4
genera and about 10 species in Europe.
Characters: M e d i u m sized (7.5-10 m m ) , b r o w n i s h to black flies,
u s u a l l y w i t h banded markings o n t h e a b d o m e n . B o d y densely
set w i t h short erect bristles. Ocelli present; eyes in male near
ly holoptic, i n female dichoptic; antenna short, third s e g m e n t
reniform; m o u t h p a r t s p o w e r f u l , at m o s t as l o n g as t h e height 461 Eggcluster of Atherix ibis (Fabricius);
o f t h e head. W i n g l o n g and w i d e , often w i t h dark markings; Tournier in Sguy 1950

114
 veins R i and R2+3 e n d i n g i n the costa at the same point; cell m3 o p e n ; cell c u p closed; legs
relatively long; tibiae o f m i d and h i n d legs w i t h apical spurs; e m p o d i u m p u l v i l l i f o r m .
Biology: T h e larvae are aquatic and prey o n larvae of other water insects. Prior to pupation, they
m o v e to nearby land. A d u l t s are always encountered near running water, usually o n the v e g
etation close by. Eggs are deposited in suitable spots over the water, such as bridges and over
hanging branches. In some species, all females deposit their eggs together before they die.
These clusters o f eggs and dead flies m a y include u p to several tens of thousands of females.
The adults largely feed o n nectar b u t some species feed o n blood of humans and catde.
Identification references: N a g a t o m i 1 9 8 4 (genera); Nartshuk 1989b (former U S S R ) ;
Stubbs 8t Drake 2001 (British Isles); S t u c k e n b e r g 1973 (revision); T h o m a s 1997 (species);
C M P D : R o z k o s n y & N a g a t o m i 1997b; C A T : Majer 1988a.

Aulacigastridae (key couplet 115; fig. 249-250)

Systematics: Acalyptrate Brachycera; superfamily
O p o m y z o i d e a ; i n E u r o p e 1 g e n u s , Aulacigaster, w i t h 4
species.
Characters: Small (2-5 m m ) , m a i n l y dark coloured flies w i t h an
e x t e n s i v e grey p r u i n o s i t y , pale tarsi a n d red e y e s . A r i s t a
p u b e s c e n t or w i t h longer rays; ocelli present; O c - b r i s t l e s
reduced or absent; P-bristles absent; 2 pairs o f F-bristles, the
l o w e r pair c u r v i n g forward and slightly i n w a r d , the u p p e r pair
b a c k w a r d and further from the e y e m a r g i n than the l o w e r pair;
interfrontal bristles absent; vibrissae present. W i n g clear;
costa w i t h humeral and subcostal breaks; v e i n Sc thin b u t
complete, v e i n s Sc and R i reaching the costa separately, b u t
just before they m e r g e for a short distance or t o u c h each other,
462 Aulacigaster leucopeza (Meigen),
apical part o f v e i n Sc m o r e pallid; crossvein B M - C u absent; cell male; Papp 1973
c u p closed. Tibiae w i t h o u t dorsal preapical bristle.
Biology: T h e larvae o f Aulacigaster live in sap e x u d i n g from w o u n d s in d e c i d u o u s and
coniferous trees, w h e r e they feed o n m i c r o - o r g a n i s m s . T h e adults are usually found near
the larval sites b u t are also attracted to other fermenting fluids or substrates; they are
k n o w n to hibernate in sheltered places i n forests.
Identification references: Kassebeer 2001 (species); Papp 1998a (revision); Stackelberg
198931 (former U S S R ) ; C M P D : Papp i 9 g 8 d ; C A T : Papp i984f.

Axymyiidae (key couplet 21; fig. 56)

Systematics: N e m a t o c e r a ; s u p e r f a m i l y A x y m y i o i d e a ; i n
Eastern Europe 1 species, Mesaxymyia kerteszi (Duda).
Characters: M e d i u m sized to large (9-12 m m ) , robust, dark,
Bibio-like species. Ocelli c o n s p i c u o u s , e y e s d i v i d e d l o n g i t u d i
nally in t w o b y a distinct suture; male holoptic; antenna c o m
pact, w i t h 14-17 segments. W i n g darkened, w i t h anal lobe and
characteristic venation. Legs relatively short; tibiae w i t h o u t
apical bristles or spurs.
Biology: T h e larvae o f this small family live i n tunnels in m o i s t 463 Axymyia turcata McAtee,
w o o d of tree trunks l y i n g o n the ground. U p to 150 tunnels per male; Wood 1981 (North America)
metre o f trunk length h a v e b e e n recorded. T h e y are b o r e d from
the outside inward, each m i n e b e i n g inhabited b y a single larva. Pupation takes place
w i t h i n the trunk as w e l l . A d u l t s are found in w o o d l a n d , in particular in shaded spots
alongside rivers and in l o w l a n d s . O n s u n n y days, t h e y fly in s w a r m s about 2 metres a b o v e
the ground.
Identification references: M a m a e v 1989c (former U S S R ) ; C M P D : K r i v o s h e i n a 2000;
C A T : M a m a e v & K r i v o s h e i n a 1986c.
BibiOtlidae (key couplet 28; fig. 63-64)
Systematics: Nematocera; superfamily Bibionoidea; traditional
ly this family includes the Pleciidae and Hesperinidae as w e l l ;
in Europe 2 genera, Bibio and Dilophus, and about 47 species.
Characters: S m a l l to large (2-15 m m ) , r o b u s t l y b u i l t
N e m a t o c e r a . U s u a l l y densely set w i t h fine hairs; b o d y colour
pale b r o w n to black, in s o m e cases (partially) red or yellow.
Ocelli c o n s p i c u o u s ; in male e y e s holoptic and consisting o f
t w o separate parts, in female eyes dichoptic and simple; anten
na inserted rather l o w , w i t h 6-12 segments. W i n g w i d e , w i t h
distinct anal lobe; w i n g clear or darkened. Tibia o f fore leg u s u
ally s w o l l e n , in Bibio w i t h a l o n g , stout spur and an apical p r o
jection; in Dilophus w i t h several r o w s o f short, stout spurs.
Biology: T h e larvae are largely found in the topsoil o f m o i s t 464 Bibio Johannis (Linnaeus),
grasslands and w o o d l a n d and in decaying w o o d . T h e y appear to male; Freeman & Lane 1985
feed mainly on rotting plant matter; in s o m e cases also o n living
roots or stems, thereby causing damage to valuable crops that can be substantial if they are
present in large numbers. Densities u p to 37,000 larvae per square metre have been record
ed. T h e adults are mainly found in spring, usually present for only a few w e e k s every year,
sometimes in v e r y large numbers. T h e males often fly or hover together in swarms. T h e
adults, those of Dilophus in particular, often visit flowers as well.
Identification references: D u d a 1930 (Lindner); Freeman & Lane 1985 (British Isles);
Haenni 1982 (Dilophus); Krivosheina 1989c (former U S S R ) ; Mikolajczyk 1977 (Poland);
T o m a s o v i c 2000 (Belgium); Z e e g e r s 1 9 9 7 , 1 9 9 8 (Netherlands);
C M P D : S k a r t v e i t i 9 9 7 ; C A T : K r i v o s h e i n a 1986b.

Blephariceridae (key couplet 19; fig. 54-55)

Systematics: Nematocera; superfamily Blephariceroidea; 5 g e n
era and about 38 species in Europe.
Characters: Small to large (3-15 m m ) , slender, delicate, dull
b r o w n to grey coloured, l o n g - l e g g e d N e m a t o c e r a . Ocelli pres
ent; e y e s often c o m p r i s i n g t w o differently faceted parts;
antenna c o m p a c t , w i t h 13-15 s e g m e n t s , s o m e t i m e s less. W i n g
w i t h a grid o f secondary v e i n s and a c o n s p i c u o u s anal lobe.
Biology: T h e larvae live in torrential streams or their splash
z o n e s and in waterfalls in hill and m o u n t a i n areas. T h e larvae
feed o n algae, bacteria and organic matter t h e y scrape from
r o c k y substrates. T h e adults are often found near the larval
habitats. W h e n resting, they often h a n g b y their fore legs,
u s i n g leaves, projecting rocks, bridges etc. as their p o i n t o f
attachment. T h e females prey on small insects like
E p h e m e r o p t e r a , Plecoptera, E m p i d i d a e (Diptera) or e v e n 465 Liponeura brevirostris Loew,
male; Sguy 1940
Blephariceridae. In s o m e cases the male is p r o v i d e d w i t h l o n g
m o u t h p a r t s used to feed on nectar, b u t m o s t males have
reduced m o u t h p a r t s .
Identification references: Frutiger 8t Jolidon 2000 (larvae,
p u p a e , C e n t r a l E u r o p e ) ; N a r t s h u k 1989a (former U S S R ) ;
C M P D : C o u r t n e y 2000; C A T : Z w i c k 1992.

Bolitophilidae (key couplet 36; fig. 81)

Systematics: Nematocera; superfamily Sciaroidea; traditionally
part o f the M y c e t o p h i l i d a e ; in Europe 1 g e n u s , Bolitophila,
w i t h about 36 species.
Characters: Small to m e d i u m sized (4-7 m m ) , delicate, greyish
b r o w n N e m a t o c e r a w i t h l o n g , slender legs and a b d o m e n and a
characteristic w i n g venation (e.g. basal position o f crossvein
B M - C u ) . Head and e y e s small; ocelli present; antenna conspic 466 Bolitophila cinerea Meigen,
u o u s l y l o n g and slender, s o m e t i m e s w i t h l o n g v e r t i c i l s . female; Hutson et al. 1980
 T h o r a x m u c h vaulted. W i n g clear or w i t h banded markings along the crossveins; anal lobe
w e a k or absent. C o x a e elongate; tibiae w i t h apical bristles or spurs.
Biology. T h e b i o l o g y o f this small family is little k n o w n . T h e larvae all d e v e l o p in terrestri
al m u s h r o o m s . T h e adults are m a i n l y found i n m o i s t s u r r o u n d i n g s and alongside w a t e r i n
w e l l w o o d e d areas.
Identification references: Chandler & R i b e i r o 1995 (Atlantic islands); H u t s o n et al. 1980
(British Isles); Kurina & Schacht 2003 (synopsis identification references); Stackelberg
i g 8 9 g (former U S S R ) ; A . I . Z a i t z e v (Eastern Europe); C M P D : Soli et al. 2000; C A T :
Plassmann 1988.

Bombyliidae (key couplet 61;fig.128-130)

Systematics: L o w e r m u s c o m o r p h B r a c h y c e r a ; s u p e r f a m i l y
Asiloidea; traditionally this family includes the
M y t h i c o m y i i d a e ; s o m e 52 genera and about 3 4 0 species in
Europe.
Characters: M i n u t e to large (1-20 m m ) flies, often thorax and
a b d o m e n covered w i t h fine hairs or in part scale-like hairs.
B o d y usually broad, bee-like b u t s o m e t i m e s slender; colour
b r o w n , grey, black or w i t h a c o n s p i c u o u s pattern o f contrast
ing colours; in s o m e cases the b o d y is d e v o i d o f hair c o v e r i n g
and shining. Eyes large, male often holoptic; antenna w i t h 3-4
segments, the third s e g m e n t largest and t h e fourth m a k i n g u p
the flagellum, there m a y also b e a v e r y small transparent style.
467 Anthrax anthrax (Schrank),
Mouthparts w i t h o u t or (usually) w i t h a l o n g , sucking p r o male; Tth 1977
boscis. W i n g occasionally clear b u t m o s t often w i t h a conspic
u o u s pattern; v e i n s R2+3 and R 4 often strongly c u r v i n g forward; cell b e l o w discal cell
(cell cuai) o p e n ; cell c u p usually o p e n ; w i n g s held h o r i z o n t a l l y w h e n at rest. Legs l o n g
and slender; tibiae w i t h apical bristles or (rarely o n m i d leg) spurs; e m p o d i u m bristle-like
or absent.
Biology: T h e larvae are parasitoids or predators o f eggs, larvae and pupae o f other insects
and e g g masses o f spiders. M a n y species parasitise bees in w h i c h case t h e female deposits
her eggs in or near the nest o f the bees. T h e first instar enters t h e nest and feeds o n the
contents o f the cells, including the b e e larvae. O t h e r species parasitise solitary w a s p s ,
antlions, pupae o f m o t h s and beetles in t h e soil or are predatory o n e g g s o f grasshoppers.
A d u l t s usually feed o n nectar and t h e females o n pollen. In a f e w species, the short-lived
adults do n o t feed at all. B o m b y l i i d a e are generally g o o d fliers and m a n y are able to h o v e r
at the same spot i n front o f a flower in order to feed w h i l e h o v e r i n g . Species n o t p o s s e s s
ing a l o n g proboscis often settle o n the flowers t h e m s e l v e s . T h e adults are m a i n l y active
on w a r m , s u n n y days and prefer o p e n , d r y habitats.
Identification references: V a n der G o o t & V a n V e e n 1996 ( N o r t h w e s t e r n Europe);
Greathead & Evenhuis 2001 (key genera); O l d r o y d 1969; Stubbs 8t Drake 2001 (British
Isles); T o t h 1977 (Hungary); Trojan 1967 (Poland); V.F. Z a i t z e v
1989b (former U S S R ) ; C M P D : Greathead 8t E v e n h u i s 1997;
C A T : V.F. Z a i t z e v i 9 8 g d ; Evenhuis 8t Greathead 1 9 9 9 .

(Borboridae): n a m e u s e d i n t h e past for Sphaeroceridae.

(Borboropsidae): part o f the H e l e o m y z i d a e .

Brachystomatidae (key couplet 61;fig.131)

Systematics: L o w e r e r e m o n e u r a n B r a c h y c e r a ; s u p e r f a m i l y
Empidoidea; separated from t h e Empididae b y Sinclair 8t
C u m m i n g 2006; except for Brachystoma, t h e family k e y s o u t
here under the Empididae b y lack o f suitable easy to u s e char
acters for inclusion in t h e k e y ; in Europe 13 species in 5 genera:
Brachystoma (2), Gloma (1), Heleodromia (7),
Pseudoheleodromia (1) and Trichopeza (2).
Characters: Small to m e d i u m (2-7 m m ) , slender, y e l l o w i s h to 468 Heleodromia immaculata
dark flies. Head small w i t h relatively large eyes, in s o m e cases Haliday, male; Collin, 1961

117
 male holoptic; antenna usually w i t h 3 s e g m e n t s of w h i c h the third is the largest and bears
a l o n g usually apical arista or style; m o u t h p a r t s stout, projecting d o w n w a r d s t o w a r d the
fore legs. W i n g clear or infscate; v e i n R4+5 forked except in Heleodromia; cell d m a l w a y s
present. Legs strong, l o n g and slender; fore legs s o m e w h a t raptorial.
Biology: T h e larvae are a s s u m e d to prey o n other insects, although n o n e are k n o w n ;
a s s u m e d to live in d a m p soil and likely not aquatic. T h e adults are predators, m a i n l y on
flying Dptera. Flies are associated w i t h m a t u r e forests, s o m e s w e p t from w e t depres
sions. A e r i a l m a t i n g s w a r m s are not k n o w n .
Identification references: Collin 1961 (British Isles); G o r o d k o v & K o v a l e v 1989 (former
U S S R ) ; Sinclair 8t C u m m i n g 2006 (classification); W a g n e r 1985 (Heleodromia); C M P D :
n o contribution; C A T : Chvla 8t W a g n e r 1989.

Braulidae (key couplet 207; fig. 4 3 9 - 4 4 0 )

Systematics: A c a l y p t r a t e Brachycera; superfamily Carnoidea;
in Europe 1 g e n u s , Braula, w i t h 3 species.
Characters: M i n u t e to small (1.2-2.5 m m ) , dorsoventrally flat
tened, reddish to dark b r o w n flies, w i t h the b o d y e v e n l y c o v
ered b y rather l o n g bristles. W i n g s , halteres and scutellum
absent. Eyes reduced to a pale spot j u s t b e h i n d the s u n k e n
antenna; ocelli absent. T h o r a x small, closely adpressed to the
a b d o m e n . Legs relatively l o n g and strong; tarsal c l a w s trans
f o r m e d into a c o m b o f over 20 teeth, c u r v i n g i n w a r d .
Biology: T h e larvae and adults live as c o m m e n s a l s in the nests o f
b e e s . T h e larvae feed o n honey, p o l l e n and organic debris. T h e
469 Braula coeca Nitzsch,
adults attach t h e m s e l v e s to the b o d y o f the q u e e n (rarely o f a female; Sguy 1950
w o r k e r ) , absorbing food and liquids from the m o u t h o p e n i n g
of the bee. T h e o n l y species o f i m p o r t a n c e to b e e - k e e p i n g is
Braula coeca N i t z s c h . It w a s n o t a real nuisance b u t nevertheless has disappeared from
m a n y countries o n account o f measures taken against Varroa m i t e s w h i c h cause severe
d a m a g e in b e e h i v e s .
Identification references: D o b s o n 1999 (British Isles); r s i Pal 1966 (revision);
Stackelberg i98gax (former U S S R ) ; C M P D : Papp I9g8g; C A T : Papp 1 9 8 4 ^

Calliphoridae (key couplet 1 8 5 - 1 8 7 , 1 9 0 ; fig. 1 2 , 1 4 , 1 7 9 , 4 0 1 ,

409,411,413,417-418,421-423)
Systematics: Calyptrate Brachycera; superfamily Oestroidea; in
Europe s o m e 22 genera and about 115 species.
Characters: T h e family m o s t probably does n o t represent a nat
ural systematic g r o u p i n g and hence is n o t easy to characterise
unequivocally. It includes small to large (4-16 m m ) , usually
robust flies. M a n y species have a greenish or bluish metallic
lustre, w i t h s o m e silvery, y e l l o w i s h , or g o l d e n p r u i n o s i t y in a
variable pattern; in s o m e species the b o d y colour is greenish
y e l l o w , p o l i s h e d black, occasionally w i t h o u t metallic sheen.
Head, b o d y and legs often w i t h strong bristles, in Pollenia t h o
rax c l o t h e d w i t h y e l l o w c u r l y hairs. A r i s t a u s u a l l y l o n g 470 Calliphora uralensis Villeneuve,
p l u m o s e in at least the basal t w o - t h i r d s , rarely pubescent. T h e male; Stackelberg 1956
shape o f the l o w e r calypter is strongly variable, ranging from
w e l l rounded-triangular to elliptic; w i n g a l w a y s w i t h v e i n M i c u r v i n g forward; cell r4+5
o p e n or closed at the w i n g margin, rarely w i t h a v e r y short petiole. Meral bristles a l w a y s
present and s u b s c u t e l l u m distinctly absent, that is, the part b e l o w the scutellum is flat,
n o t c o n v e x ; it is usually separated from the s c u t e l l u m b y a m e m b r a n o u s part; posterior
spiracle c o v e r e d b y an anterior and posterior lappet, usually o f unequal size w i t h the p o s
terior lappet usually f o r m i n g an o p e r c u l u m .
Biology: M a n y species d e v e l o p in carrion o f v a r i o u s origin. Eggs are frequently deposited
o n dead vertebrates, m a k i n g s o m e calliphorids i m p o r t a n t in forensic e n t o m o l o g y . T h e
larvae can be: true obligate agents of myiasis in vertebrate animals (e.g. Lucilia bufonivora
M o n i e z o n amphibia) and h u m a n s (Dermatobia, in the Tropics); n e c r o p h a g o u s to faculta-

118
 tive agents o f myiasis (Calliphora, Cynomya, Lucilia, e.g. L. sericata (Meigen) i n sheep,
Phormia, Protophormia); parasites o f earthworms and other w o r m s (Bellardia, Onesia,
Polleniinae), snails and slugs (Angioneura, Eurychaeta, Melinda, Melanomyd), or egg clus
ters o f g r a s s h o p p e r s (Stomorhina); or parasites s u c k i n g b l o o d from bird nestlings
(Protocalliphord). T h e adults are often found o n flowers, plant detritus, carrion and d u n g .
T w o species, Pollenia rudis (Fabricius) a n d Protophormia terraenovae (Robineau-
D e s v o i d y ) s o m e t i m e s occur indoors in large n u m b e r s , l o o k i n g for a shelter to survive the
winter.
Identification references: C r o s s k e y 8t Lane 1 9 9 3 ( m e d i c a l l y i m p o r t a n t g e n e r a ) ;
Erzinclioglu 1996 (British Isles); G r u n i n 1989c (former U S S R ) ; Huijbregts 2002 (survey
Netherlands); R o g n e s 1991 ( N o r t h w e s t e r n Europe), 2002 (Israel); C M P D : R o g n e s 1998;
C A T : S c h u m a n n 1986.

(Calobatidae): part o f the Micropezidae.

Camillidae (key couplet 95; fig. 191, 207, 209-210)

Systematics: Acalyptrate Brachycera; superfamily Ephydroidea; in
Europe 1 g e n u s , Camilla, w i t h 8 species.
Characters: Small (2-3.5 m m ) , slender, dark and lustrous flies.
Arista w i t h l o n g rays above and shorter rays b e l o w ; ocelli present;
Oc-bristles present; P-bristles c o n v e r g i n g ; 2 pairs o f F-bristles, the
l o w e r pair c u r v i n g forward, the u p p e r pair b a c k w a r d ; scattered
interfrontal setulae present; vibrissae present. W i n g u n m a r k e d ;
costa w i t h b o t h humeral and subcostal breaks; v e i n Sc incomplete;
crossvein B M - C u absent; cell c u p o p e n . Femur o f fore leg w i t h a
spine o n its ventral side, b e y o n d the m i d d l e ; o n l y tibia o f m i d leg
w i t h dorsal preapical bristle.
Biology: T h e b i o l o g y o f this s m a l l f a m i l y is little k n o w n .
Presumably the larvae feed o n organic matter (dung and detritus);
they have b e e n collected in soil at the entrances o f rabbit b u r r o w s
and in the nest o f a field v o l e (Microtus agrestis (Linnaeus)). T h e Camilla glabrata Collin, female;
4 7 1

P a p p 1 9 7 3
adults are frequently found o n flowers, o n l o w v e g e t a t i o n or o n
bare soil, near the entrance o f b u r r o w s , and are often found o n
windows.
Identification references: B e u k 8t D e Jong 1 9 9 4 (Netherlands); Papp 1985 ( r e v i e w ) ;
Stackelberg I98gay (former U S S R ) ; C M P D : Papp 1998k; C A T : Papp 19840.

Campichoetidae (key couplet 100; fig. 208,223-225)

Systematics: A c a l y p t r a t e Brachycera; superfamily Ephydroidea;
traditionally part o f the Diastatidae; in Europe 7 species i n 2 g e n
era, Campichoeta (6) and Euthychaeta (1).
Characters: Small (2.5-4 m m ) flies, b o d y colour dark w i t h head and
legs partially y e l l o w . A r i s t a s h o r t p u b e s c e n t t o m o d e r a t e l y
p l u m o s e ; ocelli present; Oc-bristles present; P-bristles c o n v e r g
ing; 2-3 pairs o f F-bristles, 1 pair c u r v i n g forward, the other pair(s)
backward; interfrontal bristles absent; vibrissae present. W i n g
partially tinged, in particular along the costa and w i t h a central
pale spot, or w i n g c o m p l e t e l y tinged b r o w n i s h to greyish; costa
w i t h a subcostal break; v e i n Sc incomplete; crossvein B M - C u pres
ent; cell c u p closed. Tibiae w i t h dorsal preapical bristle.
Biology: T h e larvae b e l o n g i n g to this family are u n k n o w n ; they
have b e e n suggested to be saprophaeous in rotten w o o d b u t this
1 c- j * j L r j 1 472 Campichoeta punctum Meiqen,
has n o t been confirmed. T h e adults are found o n l o w vegetation in g p a p p 1g 7

w o o d l a n d areas; Campichoeta is found in m o i s t w o o d l a n d s in par

ticular.
Identification references: C h a n d l e r 1986 (British Isles), 1987 (revision); C M P D : C h a n d l e r
1998b; C A T : Papp 1984P (as subfamily o f Diastatidae).

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Canacidae (key couplet 100; fig. 238a-b)
Systematics: Acalyptrate Brachycera; superfamily
Carnoidea; in Europe 4 species in 2 genera, Canace (3) and
Xanthocanace (1).
Characters: M i n u t e to small (1.6-5 m m ) , robust, y e l l o w to
greyish black, pruinose flies, generally w i t h c o n s p i c u o u s
w h i t i s h to greyish markings. Head large w i t h small anten
na; arista bare to pubescent; ocelli present; Oc-bristles
present; P-bristles absent b u t often replaced b y diverging
pseudopostocellar bristles; 2-5 pairs o f F-bristles, c u r v i n g
o u t w a r d ; interfrontal bristles present; gena w i t h 1 or more
u p c u r v e d bristles; vibrissae present. W i n g u n m a r k e d ;
costa w i t h a subcostal break; v e i n Sc parallel to v e i n R i and
m e r g i n g w i t h that v e i n just before the costa; crossvein
B M - C u present; cell c u p closed. Tibiae w i t h o u t dorsal 473 Canace salonitana Strobl, male;
preapical bristle. Canzoneri & Meneghini 1983
Biology: A l l the European species live i n the tidal z o n e
w h e r e t h e y feed o n algae and organic matter.
Identification references: C a n z o n e r i & M e n e g h i n i 1983 (Italy); Mathis 1982 (Canace)
Mathis & Freidberg 1982 (Xanthocanace); Stackelberg i 9 8 g a w (former U S S R ) ; C M P D
Mathis 1998; C A T : C o g a n 1984a.

Canthyloscelidae (key couplet 24; fig. 61-62)

Systematics: N e m a t o c e r a ; superfamily Scatopsoidea; t h e
families Canthyloscelidae and S y n n e u r i d a e are often c o n
sidered o n e family, g i v e n various n a m e s ; in N o r t h e r n ,
Eastern and Central Europe 2 species: Hyperoscelis eximia
(Boheman) and H. veternosa M a m a e v & K r i v o s h e i n a .
Characters: S m a l l t o m e d i u m s i z e d (2.5-9 m m ) , s t o u t
N e m a t o c e r a . B o d y colour b r o w n to black w i t h less dark
spots. Ocelli present b u t m e d i a l ocellus small or absent;
eyes large, holoptic and n a r r o w l y separated b e l o w the
antennae; antenna w i t h 12-16 segments, t h e terminal seg
m e n t elongate. W i n g slightly tinged, i n particular along
the costa and at t h e w i n g tip. Legs relatively short and
stout; hind leg w i t h apical half o f femur s w o l l e n and tibia
curved.
Biology: A l m o s t n o t h i n g is k n o w n about the b i o l o g y o f t h e
t w o European species o f Hyperoscelis. T h e larvae h a v e
b e e n found in moist, rotting w o o d o f s t u m p s and fallen 474 Hyperoscelis veternosa Mamaev &
tree trunks. T h e adults are v e r y rare (at least in collections) Krivosheina; Haenni 1997
and it is a s s u m e d that t h e y occur especially i n old u n d i s
turbed w o o d l a n d s , i n close v i c i n i t y o f the breeding sites o f
the larvae.
Identification references: A n d e r s s o n 1982 ( S w e d e n ) ;
H u t s o n 1 9 7 7 ( r e v i s i o n ) ; K r i v o s h e i n a 1 9 8 9 b (former
U S S R ) ; C M P D : H a e n n i 1997b; C A T : M a m a e v &
K r i v o s h e i n a 1986b.

Carnidae (key couplet 127, 131, 215; fig. 272a-b, 286-289,

449)
Systematics: Acalyptrate Brachycera; superfamily
Carnoidea; 3 genera and about 4 0 species in Europe.
Characters: M i n u t e to small (1-2.5 m m ) , m o r e or less l u s
trous black flies. Arista pubescent; ocelli present; O c - b r i s
tles present; P-bristles diverging, parallel, c o n v e r g i n g , P-
bristles absent i n Camus; 4 pairs o f F-bristles, l o w e r 2 pairs 4 7
Carnus hemapterus Nitzsch, male en
5

shortwinged female; Seguy 1951

 c u r v i n g inward, u p p e r 2 pairs obliquely o u t - b a c k w a r d ; interfrontal bristles present; v i b -
rissae present. W i n g n o t marked; costa w i t h b o t h h u m e r a l and subcostal breaks; v e i n Sc
incomplete; crossvein B M - C u absent, crossvein D M - C u absent in Camus; cell c u p o p e n
or absent, in Camus the w i n g m i g h t b e shed off near the base. Tibia w i t h o u t dorsal preapi-
cal bristle.
Biology: T h e larvae o f m o s t species d e v e l o p in all sorts o f organic matter o f animal origin,
in particular in droppings and d u n g , carrion, and nests o f birds; o n l y a f e w species can
develop in other decaying m e d i a , i n c l u d i n g rotten fungi or plant matter. A d u l t s usually
occur o n or near the breeding substrates, b u t visit flowers as w e l l , w i t h a preference for
umbellifers. T h e species o f Camus live exclusively in the nests o f birds, especially those
breeding in b u s h e s , trees and in (tree)holes, w h e r e the larvae feed o n nest debris. T h e
adults emerge from the pupa w i t h n o r m a l w i n g s , fly about until t h e y have found another
suitable nest and then shed their w i n g s . T h e y attach t h e m s e l v e s to the nestlings and feed
on the flakes and m o i s t u r e o f their skin, and they are suspected to feed o n b l o o d as w e l l .
Identification references: G r i m a l d i 1977 (Camus); H e n n i g 1972 (review); C M P D : Papp
1998b; C A T : Papp 1984k.

Cecidomyiidae (key couplet 16, 2 2 , 1 9 3 , 201-202; fig. 4 9 , 5 7 ,

424.437)
Systematics: Nematocera; superfamily Sciaroidea; s o m e 280
genera an about 1 6 4 0 species in Europe.
Characters: Minute to small (0.5-3 m m ) , rarely larger (up to 8
m m , w i n g length u p to 15 m m ) , delicate N e m a t o c e r a . Eyes
holoptic w i t h the exception o f a f e w genera w i t h reduced
w i n g s ; m o u t h p a r t s reduced; antenna in general c o n s p i c u o u s l y
long, usually w i t h 12-14 segments, s o m e t i m e s less, s o m e t i m e s
more, up to 4 0 ; antennal segments bearing differently-shaped
sensoria, in s o m e Porricondylinae and in all C e c i d o m y i i n a e
these are thread-like, in the supertribe C e c i d o m y i i d i the sen
soria (some or most) in the shape o f l o n g l o o p s ; ocelli present
(in the Lestremiinae) or absent. W i n g usually clear, in a f e w
species w i t h a pattern; n u m b e r o f longitudinal v e i n s reduced;
costa usually w i t h a break just b e y o n d v e i n R5. Legs long, tibi
ae lacking apical bristles.
Biology: T h e larvae and adults can be found in a large array o f
habitats. T h r e e m a i n groups can be distinguished w i t h respect 476 Contarinia tritici (Kirby), male;
Balachowsky & Mesnil 1935
to the feeding m o d e o f the larvae: 1) larvae living in, and feed
ing on, m u s h r o o m s , bracket fungi, d y i n g and d e c a y i n g w o o d ,
or other organic substrates (soil, litter, plant remains, etc.); 2) larvae that cause damage to
higher plants, either living free, in galls, or as leaf-miner; 3) larvae p r e y i n g o n i n v e r t e
brates, in particular o n (larvae of) small insects. O f the three subfamilies, Lestremiinae,
Porricondylinae and C e c i d o m y i i n a e , o n l y representatives o f the last cause the formation
of galls. Several o f these species can cause severe damage in agriculture and horticulture.
Identification references: Berest & M a m a e v 1989 (genera Lestremiinae); Jaschhof 1998
(Lestremiinae); M a m a e v 1989b (former U S S R ) , 1 9 9 0 (genera Porricondylinae), 1991
( H o l o n e u r i n i ) ; M h n 1 9 6 6 - 1 9 7 1 (Lindner); N i j v e l d t 1 9 6 9
(species o f e c o n o m i c importance); C M P D : Skuhrav 1997;
C A T : S k u h r a v 1986; G a g n 2 0 0 4 ; S k u h r a v et al. 2006
(Denmark).

Ceratopogonidae (key couplet 18; fig. 53)

Systematics: Nematocera; superfamily C h i r o n o m o i d e a ; s o m e
30 genera and about 590 species i n Europe.
Characters: Minute to small (1-5 m m ) , slender to robust, often
dark coloured N e m a t o c e r a , m o s t capable o f p i e r c i n g (e.g.
Dasyhelea n o t capable), allied to the C h i r o n o m i d a e , b u t less
slender and w i t h broader w i n g s . Ocelli absent; antenna w i t h 477 Culicoides pulicaris (Linnaeus);
13-15 segments w h i c h in the male are m o r e or less pubescent, female; Sguy 1951

121
 W i n g usually clear, in species o f s o m e genera w i t h a pattern; usually the R-veins short
ened, enclosing 1 or 2 small cell(s), and m o r e strongly d e v e l o p e d than the other veins. Legs
simple, s o m e t i m e s w i t h ventral spines o n femora, especially o f the fore leg, spines usual
ly m o r e d e v e l o p e d in females.
Biology: T h e larvae o f m o s t genera are aquatic, living in w e l l s , brooks, rivers, lakes (record
ed as deep as 43 metres b e l o w the surface), p o n d s , p o o l s , w a t e r in rot holes in trees, in leaf
axils and other small b o d i e s o f w a t e r w i t h a h i g h content o f organic matter; in the foam at
the surface o f aquariums, etc.; larvae o f s o m e species are k n o w n from saline waters.
Larvae o f other genera inhabit m o r e terrestrial habitats such as w e t m e a d o w s , peat b o g s ,
fenland, m o i s t soil, leaf litter, m o s s , e.g. Sphagnum, under tree bark, in rotting roots and
bracket fungi, in e x u d i n g tree sap, in d r y i n g horse m a n u r e and c o w dung, in ants nests.
T h e larvae feed o n a diversity o f food sources; s o m e feed o n m i n u t e organic particles or
m i c r o - o r g a n i s m s , others are predators o f insect larvae, w o r m s and small invertebrates.
T h e adults o f m o s t species are largely found at a distance from the larval habitats. A d u l t s
can occur in large n u m b e r s , in particular w h e n e v e n i n g approaches, but m a n y species, e.g.
those o f Leptoconops, are m a i n l y active b y day. T h e males often fly in s w a r m s . A d u l t s o f
b o t h sexes feed o n flower nectar and h o n e y d e w . In s o m e cases, e.g. Dasyhelea, this is the
o n l y food source, b u t usually the females feed o n b o d y fluids o f insects; those o f
Leptoconops, Culicoides and Forcipomyia (Lasiohelea) feeding o n the blood of mammals
including h u m a n s . T h i s b l o o d serves as a source o f protein for e g g production. Despite
their tiny size, m a n and animals alike can suffer substantially from ceratopogonids, on
account o f the n u m b e r o f ceratopogonids b e i n g so vast and because w h i l e feeding on
b l o o d t h e y can transmit diseases.
Identification references: B o o r m a n 1993 (genera feeding o n blood); R e m m 1989 (former
U S S R ) ; S z a d z i e w s k i e t a l . 1997 (genera); W i r t h & G r o g a n 1988 (World); C M P D : B o o r m a n
1997; C A T : R e m m 1988; Borkent 8t W i r t h 1 9 9 9 .

Chamaemyiidae (key couplet 1 1 9 , 1 6 0 ; fig. 257a-b, 3 4 4 - 3 4 6 )

Systematics: Acalyptrate Brachycera; superfamily
L a u x a n i o i d e a ; t r a d i t i o n a l l y t h i s f a m i l y i n c l u d e s the
Cremifaniidae; 9 genera and about 110 species in Europe.
Characters: M i n u t e to small (1-5 m m ) , usually silvery grey flies,
in s o m e cases w i t h b r o w n , grey or black markings on the head,
thorax and a b d o m e n . Arista bare to pubescent; ocelli present;
O c - b r i s t l e s p r e s e n t , in s o m e cases c u r v i n g b a c k w a r d , or
absent; P-bristles c o n v e r g i n g or reduced to absent; 0-3 pairs o f
F-bristles, if present, c u r v i n g b a c k w a r d ; especially in the large
genus Leucopis dorsal bristles o f the head largely absent; scat
tered interfrontal setulae present; vibrissae present or absent,
often a series o f vibrissa-like bristles present. W i n g u n m a r k e d ,
in s o m e cases costal area darkened; costa continuous; v e i n Sc
c o m p l e t e ; crossvein B M - C u present; cell c u p closed. Tibiae
w i t h o u t dorsal preapical bristle. S J P ^ W ? ' P Y S T I 9 M A ( M E I 9 E N )
'
Biology: T h e larvae prey o n aphids, scale insects and their allies,
e.g. gall-building Psyllidae. S o m e species have b e e n artificially
introduced as biological control agents against pest w o o l l y aphids, especially in conifer
o u s forests. T h e adults are found especially in grassland habitats and near the larvae w h e r e
flies of e.g. the g e n u s Leucopis feed o n h o n e y d e w . T h e adults of this genus are able to w a l k
s i d e w a y s and b a c k w a r d s w i t h ease, probably an adaptation enabling t h e m to avoid the
ants trying to chase off or kill e v e r y t h i n g that c o m e s near 'their' aphids.
Identification references: Tanasijtshuk 1986, i 9 8 g d (former U S S R ) , 1992 (genera);
T a n a s i j t s h u k & B e s c h o v s k i 1991 (Eastern E u r o p e ) ; C M P D : M c L e a n 1998a; C A T :
Tanasijtshuk 1984a.
Chaoboridae (key couplet 14; fig. 3 9 , 4 6 )
Systematics: Nematocera; superfamily Culicoidea; 3 genera and
9 species in Europe.
Characters: Small to m e d i u m sized (2-10 m m ) , delicate, pale
yellow, b r o w n or grey Nematocera, r e s e m b l i n g m o s q u i t o e s
(Culicidae). A n t e n n a e , w i n g s , legs and a b d o m e n l o n g and
slender. Ocelli absent; antenna a l w a y s w i t h 15 s e g m e n t s ,
p l u m o s e in the male; m o u t h p a r t s p r o t r u d i n g b u t m u c h less
than in m o s q u i t o e s . U p p e r part o f thorax w i t h f e w to m a n y
bristles b u t w i t h 2 pairs o f bare stripes. W i n g n a r r o w ; v e n a t i o n
as in Culicidae.
Biology: Larvae w i t h a remarkably transparent b o d y , especially
in Chaoborus, g i v i n g rise to the n a m e of p h a n t o m larvae. T h e y
are aquatic and prey on small invertebrates. T h e larvae o f
Cryophila, Mochlonyx a n d Chaoborus (Shadonophasma)
occur in t e m p o r a r y puddles, those o f Chaoborus (Chaoborus),
Chaoborus (Peusomyia) and Mochlonyxin various permanent 479 Chaoborus crystallinus (De Geer),
male; Saether 1997b
standing waters (lakes, p o n d s , s w a m p s , peat b o g s , etc.). T h e
adults are largely found near the larval habitats and often fly in
swarms.
Identification references: Saether 1997a (species), 2002 (species); Stackelberg i g 8 9 d (for
m e r U S S R ) ; C M P D : Saether 1997b; C A T : W a g n e r 1990b.

Chironomidae (key couplet 18,197; fig. 52,433-434)

Systematics: N e m a t o c e r a ; s u p e r f a m i l y c h i r o n o m o i d e a ; in
Europe s o m e 180 genera and about 1190 species.
Characters: M i n u t e to m e d i u m sized (1-10 m m ) N e m a t o c e r a
w i t h long, slender w i n g s , a b d o m e n and legs. Ocelli absent;
antenna generally w i t h 13 to 17 segments w i t h c o n s p i c u o u s l y
long verticils in the male. W i n g usually clear, in s o m e cases
w i t h dark markings; anterior v e i n s m o r e strongly d e v e l o p e d
than posterior ones.
Biology: N o t w i t h s t a n d i n g a considerable n u m b e r o f species o f
w h i c h the larvae are terrestrial, the larvae o f the majority o f
the species are aquatic or semi-aquatic. T h e y inhabit a large
array o f habitats ranging from the pristine to the m o s t severe
ly polluted and including brackish, m a r i n e and saline habitats.
The larvae feed o n algae, small or larger particles o f organic
matter and o n plankton; s o m e are carnivorous (Tanypodinae 480 Microtendipes nervosus (Staeger),
male; Seguy 1951
and s o m e others). T h e c o m p o s i t i o n o f the larval fauna is a
major factor in ascertaining the quality o f the water. T h e
adults, likewise, occur in a w i d e range of habitats, largely those
near water. In s o m e cases, the males fly in c o n s p i c u o u s l y large
swarms.
Identification references: Klink 8t Moller Pillot 2003 (larvae;
N o r t h w e s t e r n E u r o p e ) ; L a n g t o n &. V i s s e r 2003 (pupae;
N o r t h w e s t e r n Europe); Lindegaard 1997 (subfamilies, tribes);
Pasini 8t Ferrarese 1998 (Italy); Pinder 1978a, 1978b (British
Isles); Shilova 1989 (former U S S R ) ; W i e d e r h o l m 1983 (lar
vae), 1986 (pupae), 1989 (adults); W i l s o n & R u s e 2005 (pupae
British Isles); C M P D : Saether et al. 2000; C A T : A s h e &
Cranston 1990.

Chiropteromyzidae (key couplet 98; fig. 2i6a-b)

Systematics: Acalyptrate Brachycera; superfamily
Sphaeroceroidea; traditionally part o f the H e l e o m y z i d a e ; in
Europe 2 species: Chiropteromyza wegelii Frey and Neossos 481 Chiropteromyza wegelii Frey, male;
original (drawing by Dick Langerak)
nidicola Frey.

123
Characters: M i n u t e (1.2-2 m m ) , black, metallic lustrous flies. Arista bare to pubescent; ocel
li present; Oc-bristles present; P-bristles c o n v e r g i n g ; 2 pairs o f F-bristles, the l o w e r pair
c u r v i n g o u t w a r d ; interfrontal setulae present; vibrissae present, in Chiropteromyza w i t h
a 5-6 stronger bristles o n t h e gena b e h i n d the vibrissa. W i n g u n m a r k e d , w h i t i s h ; costa
thin in front and after crossvein H and w i t h a subcostal break; v e i n Sc incomplete, m e r g
ing apically w i t h v e i n R i ; crossvein B M - C u present; cell c u p closed. Dorsal preapical bris
tle present, s o m e t i m e s w e a k e r o n the tibiae o f fore and h i n d legs. T h e original description
o f C. wegelii Frey, 1952, m e n t i o n s that preapical bristles are absent. O n our request, Pekka
V i l k a m a a , curator o f the Finnish M u s e u m o f Natural H i s t o r y at Helsinki has examined
the t y p e material o f C. wegelii. H e has ascertained that the preapical bristles are present
(see also Haenni 1988).
Biology: T h e b i o l o g y o f this small family is little k n o w n . T h e adults have b e e n reared from
nests o f birds and from the excrement of bats.
Identification references: C M P D : Papp 19981; C A T : S o o s 8t Papp 1984.

Chloropidae (key couplet 1 3 0 , 1 6 6 , 214; fig. 240, 282-285, 361-

362,448)
Systematics: A c a l y p t r a t e Brachycera; superfamily Carnoidea;
s o m e 65 genera and about 395 species in Europe.
Characters: M i n u t e to m e d i u m sized (1-5 m m . s o m e t i m e s u p to
8 m m ) , stout to slender flies, usually w i t h a c o n s p i c u o u s l y
large ocellar triangle and f e w bristles. O f t e n y e l l o w i s h or
greenish w i t h black, b r o w n or red markings; s o m e t i m e s b o d y
entirely or i n part s h i n i n g black or dusted grey. Arista bare to
pubescent, the arista itself s o m e t i m e s enlarged; ocelli present;
Oc-bristles present or reduced; P-bristles parallel, c o n v e r g i n g
or reduced; F-bristles usually reduced, s o m e t i m e s several for
w a r d , o u t w a r d , or b a c k w a r d c u r v e d pairs present; dorsal
region o f head usually closely packed w i t h smaller setulae; 482 Camarota curvipennis (Latreille),
vibrissae present in s o m e cases, u s u a l l y reduced or absent. male; Balachowsky & Mesnil 1935
Proepisternum w i t h a sharp anterior carina. W i n g generally
clear, i n o n l y a f e w species w i t h a pattern; costa virtually a l w a y s w i t h a subcostal break;
v e i n Sc incomplete; crossvein B M - C u absent; v e i n C u A i usually w i t h a flexure halfway
cell b m + d m ; often v e i n R4+5 and s o m e t i m e s also v e i n M i c u r v i n g t o w a r d the costa; cell
c u p o p e n or absent. Tibiae w i t h o u t dorsal preapical bristle.
Biology: T h e larvae o f the C h l o r o p i d a e generally feed o n l i v i n g plant or animal tissues or o n
dead organic matter (insect frass, rotting plant material, m u s h r o o m s and bracket fungi,
carrion, nests o f birds, etc.). S o m e feed o n root aphids or o n eggs of spiders and grasshop
pers. S o m e w o o d l a n d species live i n d e c a y i n g w o o d or b e l o w the bark o f trees. Larvae
reeding o n vegetable material prefer t h e stems and flowers o f grasses, reeds and sedges
w h e r e they m a y cause galls; a n u m b e r o f these g a l l - f o r m i n g species can cause damage to
agriculture. T h e adults are encountered chiefly in o p e n habitats such as m e a d o w s and
marshland, a l t h o u g h s o m e species are t y p i c a l o f w o o d l a n d .
S e v e r a l s p e c i e s v i s i t f l o w e r s ; f r o m late s u m m e r o n w a r d
Thaumatomyia notata (Meigen) m a y o c c u r indoors, w h e r e it
seeks refuge for the winter.
Identification references: A n d e r s s o n 1977 (genera); C o l l i n
1945 (Oscinellinae British Isles); Ismay 1 9 9 9 (Chloropinae
British Isles); Nartshuk 1987 (general); Nartshuk et al. 1989
(former U S S R ) ; C M P D : I s m a y 8t N a r t s h u k 2 0 0 0 ; C A T :
Nartshuk 1984c.

Chyromyidae (key couplet 119,125; fig. 256,270)

Systematics: Acalyptrate Brachycera; superfamily
Sphaeroceroidea; 3 genera and about 6 0 species in Europe.
Characters: M i n u t e to m e d i u m sized (0.5 -8 m m ) , usually y e l
l o w to pallid flies w i t h y e l l o w bristles and hairs, in s o m e cases
.11 11 1 1. 1 1 1 1 483 Aphaniosoma nunqaricum Soos;
w i t h b r o w n , grey or black markings o n head, thorax and female-Papp 1981

124
 a b d o m e n . Arista bare to pubescent; ocelli present; Oc-bristles present; P-bristles usually
present, converging; 2-6 pairs o f F-bristles, rarely absent, t h e l o w e r m o s t pair s o m e t i m e s
c u r v i n g i n w a r d ; scattered interfrontal setulae present; vibrissae present b u t w e a k . W i n g
unmarked; costa w i t h a subcostal break, v e i n Sc i n c o m p l e t e or c o m p l e t e , parallel to R i u p
to the costa or apical part pallid; crossvein B M - C u present, cell c u p closed. Tibiae w i t h o u t
dorsal preapical bristle.
Biology: T h e larvae are u n k n o w n b u t are b e l i e v e d to d e v e l o p i n a v a r i e t y o f organic detri
tus, such as nests and excrement o f birds, m a m m a l b u r r o w s , bat d u n g , rotting w o o d ,
b e l o w the bark o f trees, etc. A d u l t s are found o n f l o w e r s , shrubs and grasses; in d r y habi
tats, b u t also along river banks, in saltmarshes, i n caves, and often indoors o n w i n d o w s .
Species o f Aphaniosoma and Gymnochiromyia preferentially occur i n saltmarshes and
semi-arid habitats and are frequent o f f l o w e r i n g vegetation; Aphaniosoma species can
also be found o n Tamarix trees along t h e Mediterranean sea coast, often in v e r y large
numbers; Chyromya species s e e m to b e associated w i t h o p e n d e c i d u o u s w o o d l a n d .
Identification references: A n d e r s s o n 1971 ( S w e d e n ) ; Ebejer 1998a (Aphaniosoma), 1998b
(Gymnochiromyia), 2005 (Italy); M e r z 1998a (genera Europe, species Switzerland); S o o s
1981 (Central Europe); Stackelberg l g S g a q (former U S S R ) ; C M P D : W h e e l e r 1998; C A T :
Soos 1984I.

ClllSiidae (key couplet 8 8 , 1 1 4 ; fig. 189-190, 245-246)

Systematics: Acalyptrate Brachycera; superfamily
O p o m y z o i d e a ; 5 genera and about 15 species in Europe.
Characters: M i n u t e to m e d i u m sized (1.5-8 m m ) , slender flies.
B o d y dull to lustrous, y e l l o w to dark b r o w n or black, often
w i t h markings; near the e y e often silvery w h i t e . Head small,
arista pubescent or w i t h longer rays; ocelli present; Oc-bristles
present to absent; P-bristles diverging or absent; 2-5 pairs of F-
bristles c u r v i n g b a c k w a r d , t h e l o w e r m o s t pair in s o m e cases
c u r v i n g i n w a r d ; interfrontal bristles present to absent; vibris
sae present. W i n g clear to s o m e w h a t infuscated, often the api
cal portion darker or crossveins w i t h dark bands; costa w i t h a
subcostal break, v e i n Sc complete; crossvein B M - C u present,
cell c u p closed. Legs all y e l l o w or partially darker; tibiae w i t h
or w i t h o u t dorsal preapical bristle. 484 Clusia flava (Meigen), male;
S o 6 s 1 9 8 1
Biology: T h e larvae o f t h e f e w species o f w h i c h s o m e t h i n g is
k n o w n live in rotting w o o d o f d e c i d u o u s trees, largely i n
m i n e s o f other insects. A d u l t s are m a i n l y found in d a m p d e c i d u o u s forests near dead or
d y i n g trees, w i t h males displaying courtship b e h a v i o u r o n trunks and s t u m p s . A d u l t s are
not frequently netted b u t at proper locations t h e y are frequently collected in Malaise
traps.
Identification references: S o o s 1981 ( C e n t r a l E u r o p e ) ;
Stackelberg ig89ak (former U S S R ) ; S o o s 1987 (genera); Stubbs
1982 (British Isles); C M P D : Sasakawa 1998; C A T : Soos 1984L

(Clythiidae): n a m e used in the past for Platypezidae.

Cnemospathidae (key couplet 98; fig. 2isa-c)

Systematics: Acalyptrate Brachycera; superfamily
Sphaeroceroidea; traditionally part o f t h e H e l e o m y z i d a e ; i n
E u r o p e 1 s p e c i e s , Prosopantrum flavifrons ( T o n n o i r &
Malloch), o f S o u t h A m e r i c a n origin, k n o w n from the English
and G e r m a n coast.
m m
Characters: Small (2.5-4 ) flies; b o d y dark w i t h a b r o w n -
grey pruinosity. A n t e n n a and part o f t h e head y e l l o w to
orange; arista v e r y short pubescent; ocelli present; Oc-bristles
present; P-bristles converging; 2 pairs o f F-bristles, t h e l o w e r
pair c u r v i n g o u t w a r d ; s t r o n g v i b r i s s a e p r e s e n t . W i n g
unmarked, w i t h a y e l l o w i s h h u e ; costa w e a k e n e d just past

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 crossvein H , w e a k and w i t h a subcostal break; v e i n Sc i n c o m p l e t e , apically m e r g i n g w i t h
v e i n R i ; crossvein B M - C u present; cell c u p closed. Tibiae w i t h strong dorsal preapical
bristle.
Biology: T h e b i o l o g y o f this small family is little k n o w n . P.flavifrons, o f S o u t h A m e r i c a n
origin, is established o n the south coast o f England (Essex and D e v o n ; Ismay & Smith
1994; C o l e 1996) and in G e r m a n y ( W e s t Friesian Islands, w h e r e the species w a s collected
in d r y d u n e areas near colonies o f gulls; Stuke &. M e r z 2005).
Identification references: Ismay & S m i t h 1 9 9 4 ; C M P D : Papp 19981.

Coelopidae (key couplet 92; fig. 202-203)

Systematics: A c a l y p t r a t e B r a c h y c e r a ; s u p e r f a m i l y
S c i o m y z o i d e a ; in Europe 3 species in 2 genera, Coelopa
(2) and Malacomyia (1).
Characters: Small to m e d i u m sized (2.5-9 m m , usually 4 -
7 m m ) ; robust, especially in Coelopa s o m e t i m e s densely
bristly or hairy, flattened and dark coloured. Eyes small,
arista bare to pubescent; ocelli present; Oc-bristles pres
ent; P-bristles parallel or c o n v e r g i n g ; 2 pairs o f F-bris-
tles, c u r v i n g o u t w a r d ; scattered interfrontal setulae
present; vibrissae absent, b u t strong bristles present near 486 Coelopa pilipes Haliday, male;
the vibrissal angle; in s o m e cases m a n y l o n g , hair-like Sguy1934
bristles present. W i n g u n m a r k e d ; costa c o n t i n u o u s ,
v e i n Sc c o m p l e t e ; crossvein B M - C u present; cell c u p closed. Legs usually densely hairy;
tibiae w i t h dorsal preapical bristle.
Biology: C o e l o p i d a e generally inhabit coastal habitats w h e r e the larvae largely d e v e l o p in
rotting s e a w e e d w a s h e d u p b y the sea. Larvae and adults can b e found t h r o u g h o u t the
year.
Identification references: Burnet i 9 6 0 (Coelopa); Stackelberg 1989Z (former U S S R ) ;
C M P D : D . K . M c A l p i n e 1998b; C A T : G o r o d k o v 1984b.

Coenomyiidae (key c o u p l e t 4 3 , 5 5 ; fig. 8 9 , 8 9 a , 117,117a)

Systematics: Lower Brachycera; superfamily
Xylophagoidea; in Europe 1 widespread species,
Coenomyiaferruginea (Scopoli).
Characters: T h e o n l y European species, Coenomyiafer
ruginea is a rather large (14-20 m m ) and robust fly, of light-
to dark b r o w n coloration , s e t w i t h l o n g golden y e l l o w hairs
on head and thorax, the abdomen being shorter pubescent
and more lustrous. Head small; eyes holoptdc in the male;
antenna w i t h 10 segments and tapering to a point. W i n g
w i t h a yellow tinge; cell m3 open; cell cup m u c h narrowing
to the w i n g margin but nearly always open. Scutellum
w i t h a pair o f short spine-like projections. Legs relatively
short; tibiae w i t h apical brisdes; e m p o d i u m pulvilliform. 487 Coenomyia ferruginea (Scopoli),
Biology: T h e larvae prey o n insect larvae in the upper female; Sguy! 926
layer o f soil containing m u c h organic matter, for e x a m
ple particles o f dead w o o d , and have b e e n found in dead w o o d itself as w e l l . T h e adults
m a i n l y o c c u r in w o o d l a n d areas w h e r e they feed o n nectar and h o n e y d e w . In s o m e cases
t h e y e m i t a penetrating odour.
Identification references: V a n der G o o t 1985 ( N o r t h w e s t e r n Europe); Nartshuk i 9 8 g d
(former U S S R ) ; C M P D : R o z k o s n y & N a g a t o m i 1997a; C A T : Majer ig88e.

Conopidae (key couplet 6 7 , 1 3 4 , 1 5 2 ; fig. 1 3 8 , 1 4 3 - 1 4 5 , 299-300, 333)

Systematics: A c a l y p t r a t e Brachycera; superfamily C o n o p o i d e a ; s o m e 14 genera and about
85 species in Europe. \~) U Q
Characters: Small to large (3-20 m m , u s u a l l y 5-15 m m ) , slender to stout flies, often lus
trous, w i t h a black and y e l l o w , w a s p - l i k e colour pattern or w i t h reddish b r o w n markings;
m o u t h p a r t s usually c o n s p i c u o u s l y l o n g and geniculate. Arista bare; ocelli present or

126
 absent (Conopinae); Oc-bristles small or absent; P-, F-, inter-
frontal bristles and vibrissae absent. W i n g usually clear, in
s o m e cases w i t h dark markings, e.g. along the costa; costa
continuous; v e i n Sc complete; crossvein B M - C u present, cell
cup closed. Tibiae w i t h (Myopinae) or w i t h o u t dorsal preapi-
cal bristle.
Biology: C o n o p i d a e are parasitoids, i n particular o f b u m b l e
bees, bees and w a s p s in w h i c h the female deposits a single
egg in the a b d o m e n . T h i s happens i n flight w h i l e the c o n o p i d
female holds the a b d o m e n o f her v i c t i m in a fixed position
u s i n g special structures o n her o w n a b d o m e n . T h e larva
develops inside, to the d e t r i m e n t of the host, pupating w h e n
the host dies. T h e adults feed o n nectar and frequently visit
flowers (for m a t i n g p u r p o s e s as well). Species w i t h v e r y l o n g
m o u t h p a r t s visit flowers w i t h d e e p - l y i n g nectar, such as
Lamiaceae (Labiatae). T h e adults prefer w a r m and d r y areas 0 0 . . . . ... . ,
. , \ n, . i n 4 8 8 Physocephala chrysorrhoea (Meiqen),
and can be round flying near vegetation rich in f l o w e r s , near | . seguy 1950
m a e

b l o o m i n g heather and o n trees and shrubs in flower. T h e

females often fly s l o w l y t h r o u g h the vegetation searching for
a suitable host. M a n y species s h o w a striking resemblance to bees, w a s p s and their allies.
Identification references: B a n k o w s k a 1979 (Poland); K o r m a n n 2002 (Central Europe);
R i v o s e c c h i 1 9 9 6 (Italy); S m i t h 1 9 6 9 b ( B r i t i s h Isles); V a n V e e n 1 9 8 4 ( B e l g i u m ,
Netherlands); Z i m i n a 1989 (former U S S R ) ; C M P D : n o contribution; C A T : C h v a l a &
S m i t h 1988.

(Cordiluridae): n a m e used i n the past for Scathophagidae.

(Corynoscelidae): n a m e used in the past for Canthyloscelidae.

Cremifaniidae (key couplet 160; fig. 347-348)

Systematics: Acalyptrate Brachycera; superfamily
Lauxanioidea; traditionally part o f the C h a m a e m y i i d a e ; 2
species in Europe: Cremifania nigrocellulata C z e r n y and C.
lanceolata Papp.
Characters: M i n u t e to small (1.5-2.6 m m ) , grey or g r e y i s h
b r o w n flies. Arista pubescent, in s o m e cases arista o f aberrant
shape; ocelli present; Oc-bristles present; P-bristles absent,
but a pair o f diverging bristles present o n the h i n d part o f the
ocellar triangle; 2 pairs o f F-bristles, c u r v i n g obliquely out-
b a c k w a r d ; scattered interfrontal setulae present; vibrissae
absent. W i n g pale grey, w i n g base, subcostal cell or subcostal
and costal cells partially darkened; costa continuous; v e i n Sc 489 Cremifania nigrocellulata Czerny,
complete; crossvein B M - C u present; cell c u p closed. Tibiae male; Papp 1998h
w i t h o u t dorsal preapical bristle.
Biology: In o n e o f the t w o European species, v i z . , C. nigrocellu
lata, the b i o l o g y is w e l l k n o w n ; its larvae live o n balsam w o o l
ly aphids o f the family A d e l g i d a e , in coniferous w o o d s . T h i s
species, like s o m e species o f the C h a m a e m y i i d a e , w a s intro
duced in N o r t h A m e r i c a for the biological control o f adelgid
pest species.
Identification references: C M P D : Papp 1998I1; C A T :
Tanasijtshuk 1984b.

Cryptochetidae (key couplet 147; fig. 322)

Systematics: Acalyptrate Brachycera; superfamily Carnoidea;
in Europe 1 genus, Cryptochetum, w i t h 3 species.
Characters: Small (2-4 m m ) , stout, black or metallic blue l u s
trous flies w i t h a strikingly large third antennal s e g m e n t lack 490 Cryptochetum grandicorne Rondani,
ing the arista. Ocelli present; O c - , P- and F-bristles absent; male; Hennig 1937b

127
 scattered interfrontal setulae present; vibrissae absent. W i n g unmarked, costa w i t h b o t h
h u m e r a l and subcostal breaks; v e i n Sc c o m p l e t e u p to the costa, b u t pallid, hardly visible;
crossvein B M - C u absent; cell c u p closed. Tibiae w i t h o u t dorsal preapical bristle.
Biology: T h e larvae b e l o n g i n g to this family are endoparasitoids in coccoids (scale insects)
living o n Scots fir and allies (Pinus) and herbs. T h e adults in s o m e cases c r o w d i n g togeth
er o n tree trunks, leaf stalks or leaves. T h o s e o f s o m e A s i a n species can be a nuisance since
they continually seek o u t the e y e s o f h u m a n s (eye gnats). O n e Australian species w a s
introduced to N o r t h A m e r i c a and e l s e w h e r e in order to keep d o w n the Australian coc-
coid, Icherya purchasi W i l l i s t o n ; this is o n e o f the best examples of successful biological
pest control.
Identification references: Stackelberg i g S g a t (former U S S R ) ; C M P D : Nartshuk 2000;
C A T : Nartshuk 1984b.

CllUcidae (key couplet 14; fig. 6a, 17-18,45)

Systematics: Nematocera; superfamily Culicoidea; 8
genera and about 105 species in Europe.
Characters: Small to medium sized (3-9 mm, rarefy up
to 15 m m ) , pale to dark coloured N e m a t o c e r a w i t h
slender w i n g s , delicate legs and a b d o m e n ; w i t h l o n g
m o u t h p a r t s , in the female piercing; w i n g s , legs and
often a b d o m e n (partially) covered b y scales. Head
w i t h elongate m o u t h p a r t s that are m u c h longer than
491 Aedes aegypti (Linnaeus),
the head; e y e s r e n i f o r m , dichoptic; ocelli absent;
female; Seguy 1950
antenna w i t h 15 segments, w i t h l o n g verticils in the
male, verticils shorter in the female.
Biology: T h e larvae largely d e v e l o p in s h a l l o w stagnant waters, predominantly in small or
t e m p o r a r y b o d i e s o f water. T h e y feed o n particles o f detritus, bacteria, algae, etc.; a f e w
species are predatory, m a i n l y o n other culicid larvae. T h e females feed o n the b l o o d in
order to obtain protein for the d e v e l o p m e n t o f their eggs. Feeding preferences vary from
o n e species to another and hosts include c o l d - b l o o d e d vertebrates b u t especially birds and
m a m m a l s , i n c l u d i n g h u m a n s . T h e Culicidae o c c u p y virtually all habitats and can occur
locally, or temporarily, in vast n u m b e r s . D u e to their b l o o d feeding habits they can be
such a nuisance that they interfere w i t h h u m a n activities and have a negative impact o n
the yield o f cattle breeding. A p a r t from this, they are vectors o f several serious diseases o f
w h i c h malaria is the best k n o w n and claims the greatest n u m b e r o f casualties.
Identification references: Becker 2003 (genera feeding o n b l o o d ) ; Cranston et al. 1987
(British Isles); D a h l 1997 (genera and subgenera); G u t s e v i c h 1989 (former U S S R ) ; Haren
&. V e r d o n s c h o t 1995 (Netherlands); H u a n g 2002 (genera); M o h r i g 1969 ( G e r m a n y ) ;
Schaffner et al. 2001 (Europe); Service 1993 (genera feeding o n b l o o d ) ; Skierska 1977
(Poland); S n o w 1 9 9 0 (British Isles); S t o j a n o v i c h & S c o t t 1995a (illustrated k e y
Fennoscandia), 1995b (European Russia); C M P D : M i n a f 2000a; C A T : M i n a f 1990; T o t h
2004 (Hungary).

Clirtonotidae (key couplet 93; fig. 204)

Systematics: Acalyptrate Brachycera; superfamily
Ephydroidea; in Europe 1 w i d e s p r e a d species, Curtonotum
anus (Meigen).
Characters: M e d i u m sized (5-7 m m ) , y e l l o w to pale grey flies
w i t h a s o m e w h a t hunchbacked thorax. Arista l o n g p l u m o s e ;
ocelli present; Oc-bristles present; P-bristles converging; 2
pairs o f F-bristles, the l o w e r pair c u r v i n g forward, the upper
pair b a c k w a r d ; interfrontal bristles or setulae absent; vibrissae
or vibrissa-like bristles present. W i n g unmarked; costa w i t h
humeral and subcostal breaks; v e i n Sc complete; crossvein B M -
C u absent; cell c u p closed. Tibiae w i t h dorsal preapical bristle.
Biology: Little is k n o w n about the b i o l o g y of this small family o f
3 genera. T h e adults are found especially in m o i s t areas (e.g. in 492 Curtonotum anus Meigen,
Israel along shores o f streams, Freidberg pers. c o m m . ) . T h e y female; Papp 1973

128
 are m a i n l y active at dusk, during the day t h e y hide in s h a d o w y spots such as animal bur
r o w s . T h e larvae o f n o n - E u r o p e a n species h a v e b e e n found i n e g g - p o u c h e s o f the migra
tory locust.
Identification references: Stackelberg 1989M) (former U S S R ) ; C M P D : Papp 1998]'; C A T :
Papp i g 8 4 q .

Cylindrotomidae (key couplet 7; fig. 26,34-37)

Systematics: Nematocera; superfamily T i p u l o i d e a ; tradi
tionally part o f the Tipulidae; 4 genera and 6 species in
Europe.
Characters: Large (11-16 m m ) , y e l l o w i s h to pale b r o w n i s h
Nematocera w i t h l o n g , slender antennae, w i n g s , legs
and a b d o m e n . Head w i t h a short rostrum; nasus absent;
antenna w i t h 16 segments; ocelli absent. U p p e r side o f
the thorax w i t h 3 w i d e dark stripes, or thorax m a i n l y
grey (Triogma); transverse suture distinct in m e d i a n
third o f thorax, fading o u t laterally. W i n g w i t h 2 anal
veins. A b d o m e n in male w i t h bipartite (Diogma) or tri
partite distiphallus, in female w i t h broad and short cer-
cus.
Biology: Larvae p h y t o p h a g o u s and, Cylindrotoma e x c e p t
ed, living o n terrestrial, semi-aquatic and aquatic m o s s
es. T h e larvae o f Cylindrotoma living o n various herbs
(Caltha, Viola, Stellaria, Anemone, etc.) in moist w o o d
land habitats w i t h a life m o d e similar to that o f caterpil
lars. T h e adults p r e d o m i n a n t l y in m a r s h y areas and
m o i s t w o o d l a n d , resting o n the vegetation or s l o w l y fly 493 Cylindrotoma distintissima (Meigen),
male; Papp & Schumann 2000
ing about near the sites w h e r e the larvae are found.
Identification references: B r i n k m a n n 1997 (genera);
Peus 1952 (Lindner); S a v c h e n k o 1989a (former U S S R ) ;
Stubbs 1 9 9 4 (British Isles); C M P D : n o contribution; C A T : S o o s 8t Oosterbroek 1992
Oosterbroek 2006.

(Cypselidae): n a m e used in the past for Sphaeroceridae.

(Cyrtidae): n a m e used in the past for Acroceridae.
(Dacidae): part o f the Tephritidae.

DiadOCidiidae (key couplet 37; fig. 82)

Systematics: Nematocera; superfamily Sciaroidea; tradi
tionally part o f the Mycetophilidae; i n Europe 1 genus,
Diadocidia, w i t h 5 species.
Characters: Small (3-4.5 m m ) , y e l l o w i s h b r o w n , delicate
Nematocera w i t h pale b r o w n to grey w i n g s w i t h a char
acteristic venation. Head and eyes small; e y e s dichoptic;
ocelli present; antenna w i t h 17 segments. T h o r a x vault
ed. Legs slender, coxae elongate; tibiae w i t h apical bris
tles or spurs. 494 Wing Diadocidia ferruginosa
Biology: T h e larvae feed o n m y c e l i u m , encapsulated in (Meigen); Soli et al. 2000
m u c o u s tubes in and u n d e r n e a t h rotting w o o d and in
m u s h r o o m s and bracket fungi. In s o m e cases the adults are found in and around h o l l o w
tree s t u m p s in great n u m b e r s .
Identification references: H u t s o n et al. 1980 (British Isles); Kurina 8t Schacht 2003 (syn
opsis identification references); Lastovka 8t Matile 1972 (review); Stackelberg 1989J (for
mer U S S R ) ; A . I . Z a i t z e v (Eastern Europe); C M P D : Soli et al. 2000; C A T : K r i v o s h e i n a
1988a.

129
Diastatidae (key couplet 100;fig.221-222)
Systematics: Acalyptrate Brachycera; superfamily
Ephydroidea; traditionally this family includes the
C a m p i c h o e t i d a e ; in Europe 1 genus, Diastata, w i t h 9 species.
Characters: Small (2.5-4 m m ) , stout, dark flies w i t h grey p r u -
inosity, head partially dull y e l l o w and lustrous silvery grey;
antennae and legs largely y e l l o w . Arista short to m e d i u m
p l u m o s e ; ocelli present; Oc-bristles present, P-bristles c o n
v e r g i n g ; 2 pairs o f F-bristles, t h e outer pair c u r v i n g forward,
the inner pair b a c k w a r d ; interfrontal bristles absent; vibrissae
present. W i n g w i t h various patterns o f dark markings, in
s o m e cases clear, b u t cell c a l w a y s darkened; costa w i t h b o t h
h u m e r a l and subcostal breaks and w i t h several small spines
a m o n g the c u s t o m a r y setulae; v e i n Sc i n c o m p l e t e ; crossvein
495 Diastata fuscula Fallen,
B M - C u present; cell c u p closed. Tibiae w i t h dorsal preapical male; Papp 1973
bristle.
Biology: T h e larvae of this small family are u n k n o w n ; they are t h o u g h t to b e saprophagous
because t h e adults are frequently encountered near litter and soil, in all t y p e s o f w o o d
land, marshes, b o g areas and alongside stagnant w a t e r s , s o m e t i m e s in great n u m b e r s .
Identification references: C h a n d l e r 1986 (British Isles), 1987 (revision); Stackelberg
i 9 8 g a z (former U S S R ) ; C M P D : Chandler 1998c; C A T : Papp 1984P.

Diopsidae (key couplet 155;fig.337)

Systematics: A c a l y p t r a t e Brachycera; superfamily D i o p s o i d e a ;
1 species, Sphyracephala europaea Papp 8t Foldvari, i n Central
Europe.
Characters: T h e single species o f European Brachycera w i t h
eyestalks, S. europaea, is small (3-4 m m ) . B o d y largely lustrous
black and covered w i t h grey, m e d i u m l e n g t h hairs; head par
tially y e l l o w . Arista bare; ocelli present; O c - and P-bristles
absent; 1 pair o f F-bristles, c u r v i n g b a c k w a r d ; interfrontal
bristles absent, interfrontal setulae present; vibrissae absent.
W i n g w i t h faint markings in t h e central part and at the tips; 496 Sphyracephala europaea (Papp &
costa c o n t i n u o u s ; v e i n Sc complete; crossvein B M - C u absent; Foldvari), male; Simova-Tosic &
cell c u p closed. Tibiae w i t h o u t dorsal preapical bristle. Stojanovic 1999
Biology: M o s t larvae o f Diopsidae are r o o t borers or miners,
especially in grasses, w h i l e s o m e species, probably i n c l u d i n g
S. europaea, live in, and feed o n , rotting organic matter. T h e adults prefer moist, s h a d o w y
sites, in particular alongside w a t e r s and m a y occur in great n u m b e r s . It is k n o w n that they
seek refuge in great n u m b e r s in caves and other sheltered places during winter. In O c t o b e r
1996 S. europaea w a s found in H u n g a r y (and Europe) for the first time. Hundreds of indi
viduals w e r e observed at the w a l l o f a h i g h river bank, m a i n l y at the entrances o f small
holes m a d e b y h y m e n o p t e r a n s . T h e species has also b e e n found in Serbia w h e r e in
N o v e m b e r 1999 several t h o u s a n d s of individuals w e r e observed sitting o n the ground o n
a h i g h b a n k o f loess along a river.
Identification references: Papp et al. 1997 (Hungary); Simova-Tosic 8t Stojanovic 1999
(Serbia); C M P D : Hilger 2000.

Ditomyiidae (key couplet 33;fig.76)

Systematics: Nematocera; superfamily Sciaroidca; traditionally
part o f the M y c e t o p h i l i d a e ; in Europe 2 genera, Ditomyia and
Symmerus, w i t h 2 species each.
Characters: M e d i u m s i z e d ( 6 - 8 m m ) , y e l l o w i s h b r o w n
N e m a t o c e r a w i t h l o n g and slender legs and a b d o m e n , and a
characteristic v e n a t i o n (including a l o n g v e i n R2+3). Head and
eyes small; e y e s a l m o s t t o u c h i n g each other or distinctly 4 9 Ditomyia fasciata (Melgen);
7 w i n g

dichoptic; ocelli present; antenna w i t h 15 segments, in s o m e Hutson et al. 1980

130
 cases laterally compressed a n d / o r l o n g pubescent. T h o r a x vaulted. W i n g clear or slightly
tinged (Symmerus) or w i t h transverse bands (Ditomyia); anal lobe w e a k or absent. C o x a e
elongate; tibiae w i t h l o n g apical bristles or spurs.
Biology: T h e larvae live in bracket fungi and in dead parts of d e c i d u o u s trees. T h e adults are
found in w o o d l a n d areas w h e r e m o s t o f their activity probably takes place at d u s k and in
the e v e n i n g .
Identification references: H u t s o n et al. 1980 (British Isles); Kurina 8t Schacht 2003 (syn
opsis identification references); Stackelberg 1989b (former U S S R ) ; A . I . Z a i t z e v (Eastern
Europe); C M P D : Soli et al. 2000; C A T : M a m a e v 8t K r i v o s h e i n a 1988a.

Dixidae (key couplet 13; fig. 4 3 - 4 4 )

Systematics: Nematocera; superfamily Culicoidea; in Europe 2
genera, Dixa and Dixella, and about 30 species.
Characters: Small to m e d i u m sized (3-5.5 m m ) , y e l l o w i s h to
b r o w n i s h grey Nematocera. Antennae, w i n g s , legs and
a b d o m e n l o n g and slender. Ocelli absent; antenna w i t h 16 seg
ments. W i n g usually clear, w i t h an anal lobe and in s o m e cases
characteristic p i g m e n t spots; v e i n R2+3 s t r o n g l y c u r v i n g .
Tibiae o f fore and m i d leg w i t h short spurs.
Biology: T h e larvae are found at or near the w a t e r surface w h e r e
they take o n a characteristic b o d y posture, a reversed U shape,
near the edge o f the w a t e r or just a b o v e it, h o l d i n g t h e m s e l v e s
up b y objects projecting from the w a t e r surface. Pupation also
takes place there. T h e y are filter feeders, l i v i n g o n m i c r o
organisms and tiny particles o f organic matter. Dixa larvae 4 9 8 D i x a s p e c f e m a | e ; W a g n e r 1 9 g 7

prefer r u n n i n g water, those o f Dixella are usually found in

stagnant water, w h i l e b o t h genera o c c u r i n s l o w - m o v i n g
water. T h e adults m a i n l y occur near the larval habitats w h e r e the males m a y fly i n
swarms.
Identification references: D i s n e y 1999 (British Isles); Stackelberg 1989c (former U S S R ) ;
W a g n e r 19976 (genera); C M P D : W a g n e r 1997b; C A T : R o z k o s n y 1990.

Dolichopodidae (key c o u p l e t 7 6 , 2 1 2 ; fig. 9 7 , 1 5 5 , 1 6 2 - 1 6 4 )

Systematics: L o w e r e r e m o n e u r a n B r a c h y c e r a ; s u p e r f a m i l y
Empidoidea; s o m e 6 0 genera and about 775 species in Europe.
For a discussion about the p h y l o g e n y and classification see
Sinclair 8c C u m m i n g 2006.
Characters: M i n u t e to m e d i u m sized (1-9 m m ) flies w i t h l o n g
and slender legs. B o d y slender, the smaller species stouter.
B o d y colour usually greenish metallic lustrous, in s o m e cases
dull y e l l o w , b r o w n or black. Eyes virtually a l w a y s dichoptic;
third antennal s e g m e n t largest and w i t h a usually long, apical
or dorsal arista; m o u t h p a r t s usually short and w i t h a w i d e
aperture adapted for sucking small prey. W i n g clear or tinged,
in s o m e cases partially strongly coloured, marked or w i t h dis
tinct spots; cells br, b m and c u p small, cell c u p a n d / o r the anal
v e i n m a y be absent; crossvein B M - C u absent; 2 v e i n s branch- 499 Dolichopus popuiaris Wiedemann,
ing from crossvein D M - C u g o in t h e direction o f the w i n g m a l e
; Verrall 1909
margin; the u p p e r o n e in s o m e cases strongly c u r v i n g or
forked into M i and M 2 . Legs, the tibiae in particular, usually w i t h l o n g bristles; in s o m e
genera legs raptorial.
Biology: T h e larvae prey o n small invertebrates and are found in particular in m o i s t soil, in
rotting organic matter, b e l o w tree bark, etc.; larvae o f Thrypticus m i n e the stalks o f
M o n o c o t y l e d o n s . T r u l y aquatic species are lacking b u t m a n y are semi-aquatic (living in
the water margin z o n e ) ; a restricted n u m b e r o f species d e v e l o p at t h e shores o f saline
inland bodies o f w a t e r or the intertidal z o n e along seashores. T h e adults occur in a w i d e
range o f habitats, for example near w a t e r or m o r e in the o p e n , e.g. in m e a d o w s and gar-

13 1
 dens. T h e y are often sitting h i g h o n their legs o n the ground or o n vegetation, o n tree
trunks, rocks, w a l l s etc.; s o m e species w a l k about o n the w a t e r surface. T h e adults, like
the l a r v a e , are p r e d a t o r s , l a r g e l y f e e d i n g o n s m a l l i n v e r t e b r a t e s . T h e s p e c i e s o f
Dolichopus, for example, prefer to prey o n the larvae o f Culicidae.
I d e n t i f i c a t i o n r e f e r e n c e s : D ' A s s i s Fonseca 1978 (British Isles); G r i c h a n o v 2004 (Sweden);
Meuffels 2001 (genera Netherlands); N e g r o b o v 8t Stackelberg 1989 (former U S S R ) ;
Parent 1938 (France); Sinclair & C u m m i n g 2006 (classification); Stackelberg & N e g r o b o v
1930-1979 (Lindner, incomplete); W e b e r 1989 (Central Europe); C M P D : n o contribution;
C A T : N e g r o b o v 1991.

(Dorylaidae): n a m e u s e d in the past for Pipunculidae.

DroSOphilidae (key couplet 9 9 , 126, 133, 220; fig.

217-220, 271, 297-298)
S y s t e m a t i c s : A c a l y p t r a t e Brachycera; s u p e r f a m i l y
Ephydroidea; s o m e 17 genera and about 120 species
in Europe.
C h a r a c t e r s : M i n u t e to m e d i u m sized (1.5-7 m m ) flies,
dull or lustrous w i t h b o d y colour v a r y i n g from y e l
l o w to b r o w n i s h black, often w i t h s o m e w h i t i s h
markings o n head and thorax a n d / o r a pattern o f
stripes o n thorax and a b d o m e n ; w h e n alive eyes
often clear red. A r i s t a usually w i t h l o n g rays, i n
500 Drosophila funebris (Fabricius);
s o m e cases pubescent; ocelli present, Oc-bristles
Seguy1951
present, except i n Acletoxenus; P-bristles usually
present, c o n v e r g i n g ; 3 pairs of F-bristles, 1 pair c u r v
ing forward or obliquely i n w a r d , 2 pairs b a c k w a r d ; scattered interfrontal bristles present;
vibrissae present. W i n g usually u n m a r k e d , in s o m e cases w i t h dark markings or exten
sively infuscated; costa w i t h b o t h h u m e r a l and subcostal breaks; v e i n Sc incomplete;
crossvein B M - C u usually absent, in s o m e cases present; cell c u p o p e n or closed. Tibiae
virtually a l w a y s w i t h dorsal preapical bristle, s o m e t i m e s small or absent (e.g. Cacoxenus).
B i o l o g y : T h e larvae o f m o s t species feed o n m i c r o - o r g a n i s m s in rotting organic matter such
as fruits, m u s h r o o m s , bracket fungi, cacti, etc. O t h e r larvae live in slime fluxes or exud
ing tree sap, are m i n e r s , or prey o n other insects (e.g. larvae o f Cacoxenus l i v i n g as para-
sitoids in nests o f solitary bees; Acletoxenus larvae are predators of larvae of Aleurodidae).
T h e adults are usually attracted b y the scent of m u s h r o o m s , bracket fungi, ripening or rot
ting organic matter, and are v e r y frequently found indoors. S o m e species are easy to rear
in large n u m b e r s , u s i n g artificial substrates. H e n c e D r o s o p h i l i d a e are m u c h used in scien
tific research in a large v a r i e t y o f fields.
I d e n t i f i c a t i o n r e f e r e n c e s : B a c h l i & Burla 1985
(Switzerland); B a c h l i et al. 2 0 0 4 (Denmark,
Scandinavia); S t a c k e l b e r g l g S g b a ; G o r n o s t a e v 2 0 0 1 (for
m e r U S S R ) ; C M P D : Bachli 1998; C A T : Bachli & Teresa
R o c h a Pite 1 9 8 4 .

Dryomyzidae (key couplet 145; fig. 314-316)

S y s t e m a t i c s : Acalyptrate Brachycera; superfamily
S c i o m y z o i d e a ; traditionally this f a m i l y i n c l u d e s the
H e l c o m y z i d a e and Heterocheilidae; in Europe 4 species
in 2 genera, Dryomyza (3) and Neuroctena (1).
C h a r a c t e r s : M e d i u m sized to large (5-18 m m ) , y e l l o w ,
orange, or b r o w n coloured, bristled or quite ' h a i r y ' flies.
A r i s t a bare t o p u b e s c e n t ; ocelli present; O c - b r i s t l e s
present, P-bristles parallel to diverging; 1-2 pairs o f F-
b r i s t l e s , c u r v i n g o b l i q u e l y o u t - b a c k w a r d ; scattered
interfrontal setulae present; vibrissae absent. W i n g clear - | Neuroctena anilis (Fallen),
5 0

or tinged and often w i t h d a r k b a n d s along the crossveins; male; Stackelberg 1956

132
 costa continuous; v e i n Sc complete; crossvein B M - C u present; cell c u p closed. Tibiae w i t h
dorsal preapical bristle.
Biology: T h e larvae d e v e l o p in rotting organic matter such as m u s h r o o m s and bracket
fungi, droppings, carrion, etc. T h e adults are m a i n l y found in moist, shaded places in
w o o d l a n d areas, usually o n putrefying matter, i n c l u d i n g carrion, decaying m u s h r o o m s ,
excrement and tree sap.
Identification references: Falk 2005 (British Isles); Stackelberg 19893a (former U S S R ) ;
C M P D : O z e r o v 1998; C A T : S o o s i 9 8 4 g .

(Eginiidae): part o f the Muscidae.

Empididae (key couplet 6 3 , 7 8 ; fig. 133-134,156,170,172)

Systematics: L o w e r e r e m o n e u r a n B r a c h y c e r a ; s u p e r f a m i l y
Empidoidea; traditionally this family includes the Atelestidae,
Brachystomatidae, H y b o t i d a e and Microphoridae; s o m e 23
genera and about 810 species in Europe. For a discussion about
the p h y l o g e n y and classification see Sinclair 8t C u m m i n g
2006.
Characters: M i n u t e to large (1-12 m m , s o m e t i m e s larger), slen
der to robust, pallid to dark flies, i n several genera w i t h p o w
erful, piercing mouthparts. Head small w i t h relatively large
eyes, in s o m e cases holoptic, often w i t h an incision at level o f
antennae; antenna usually w i t h 3 segments o f w h i c h the third
is the largest and bears a short or l o n g , usually apical arista or
style; m o u t h p a r t s short to strongly elongate, in the latter case
projecting forward or d o w n w a r d t o w a r d the fore legs. W i n g 502 Empis tesellata Fabricius, male;
clear or (partially) tinged, in s o m e cases w i t h a stigma spot, or Coiyer & Hammond 1968
w i t h a distinct w i n g pattern; v e i n R4+5 often forked; cell d m
almost a l w a y s present. Legs p o w e r f u l , usually l o n g and slen
der; in s o m e cases fore legs raptorial, adapted to catching and h o l d i n g o n to prey.
Biology: T h e larvae prey o n other insects, m a i n l y larvae o f Diptera living in the g r o u n d , o n
organic matter, or in decaying w o o d ; s o m e are aquatic. T h e adults are originally predators
as w e l l , m a i n l y o n flying Diptera, b u t s o m e species forage b y w a l k i n g about o n tree trunks
or prey o n aquatic insects, i n c l u d i n g those that h a v e just pupated near the shoreline. S o m e
adults visit flowers and partially or entirely feed o n p o l l e n and nectar. M a n y species f o r m
s w a r m s , the males offering a prey or a silken balloon, w h i c h m a y or m a y n o t contain a
prey, to the female as part o f the m a t i n g ritual. In Hormopeza large s w a r m s of b o t h sexes
have been o b s e r v e d in s m o k e o f w o o d fires.
Identification references: Bartak 1982 (Rhamphomyia Central Europe); C h v a l a 1 9 9 4
(Empis), 2005 (Hilara); C o l l i n 1961 (British Isles); V a n der G o o t 1990 (Empis Netherlands,
Belgium, L u x e m b o u r g ) ; V a n der G o o t et al. 2000 (Hilara Netherlands); G o r o d k o v 8t
Kovalev 1989 (former U S S R ) ; N i e s i o l o w s k i 1992 (Poland);
Sinclair 8c C u m m i n g 2006 (classification); W a g n e r i 9 9 7 g
(genera); C M P D : n o contribution; C A T : C h v a l a 8t W a g n e r
1989.

Ephydridae (key couplet 9 5 , 1 3 2 , 1 6 7 , 2 1 7 ;fig.211-213,290-293,

365-366,450)
Systematics: Acalyptrate Brachycera; superfamily
Ephydroidea; s o m e 6 0 genera and about 335 species i n Europe.
Characters: M i n u t e to m e d i u m sized (1-11 m m ) , usually dull,
dark coloured flies. Face often s w o l l e n and c o n v e x ; arista
s o m e t i m e s bare to pubescent b u t usually w i t h l o n g rays dor-
sally o n l y ; ocelli p r e s e n t ; O c - b r i s t l e s p r e s e n t , P - b r i s t l e s
absent, p s e u d o p o s t o c e l l a r - b r i s t l e s , i f p r e s e n t , parallel to
diverging; F-bristles pairs variable, c u r v i n g forward, o u t w a r d ,
or backward, in s o m e cases absent; scattered interfrontal setu-

133
 lae present or absent; vibrissae present. W i n g usually u n m a r k e d , in s o m e cases darkened,
m a r k e d or spotted; costa w i t h h u m e r a l and subcostal breaks; v e i n Sc incomplete;
crossvein B M - C u absent; cell c u p o p e n or absent. O n l y tibia of m i d leg in s o m e cases w i t h
dorsal preapical bristle; in s o m e cases fore legs raptorial.
Biology: T h e Ephydridae usually inhabit aquatic, semi-aquatic and coastal or non-coastal
saline habitats w h e r e t h e y m a y be found in large n u m b e r s . T h e larvae feed o n m i c r o
organisms such as bacteria, algae and yeasts; s o m e species prey o n insects and other small
aquatic life forms w h i l e others prefer rotting animal tissue, excrement, eggs o f spiders or
frogs; species feeding o n plant juices or m i n i n g in plants are k n o w n as w e l l . A d u l t s are
usually found, s o m e t i m e s in v e r y large n u m b e r s , o n or near the soil in h u m i d or w e t habi
tats w h e r e they mate, oviposit, or feed o n a w i d e variety o f organic matter like algae, h o n -
e y d e w , carrion, etc.; o t h e r w i s e , adults m a y b e found at s o m e distance from humidity, e.g.
o n tree trunks and in vegetation. S o m e species have b e e n recorded as pests in crops and
greenhouses.
Identification references: C a n z o n e r i 8t M e n e g h i n i 1983 (Italy); H o l l m a n n - S c h i r r m a c h e r
1988 ( I l y t h e i n a e ) ; N a r t s h u k i g 8 g i ( f o r m e r U S S R ) ; O l a f f s o n 1991 (Scatella,
Lamproscatella); Z a t w a r n i c k i 1997 (genera); C M P D : Mathis & Z a t w a r n i c k i 1998; C A T :
C o g a n 1984b; Papp 1 9 8 4 m (Risinae); Mathis 8t Z a t w a r n i c k i
1995-

(Erinnidae): n a m e used in the past for X y l o p h a g i d a e .

(Eurygnathomyiidae): part o f the Pallopteridae.

Fanniidae (key couplet 180; fig. 395)

Systematics: Calyptrate Brachycera; superfamily Muscoidea; 3
genera and about 82 species in Europe.
Characters: Small to m e d i u m sized (2-5 m m , s o m e u p to 9 m m )
flies, usually w i t h grey to black b o d y colour. O c c i p u t w i t h o u t
w h i t e or pallid, thin hair-like bristles; arista bare to pubescent;
interfrontal bristles absent. Meral bristles absent; l o w e r side o f
s c u t e l l u m bare i n its m i d d l e region; s u b s c u t e l l u m absent.
L o w e r calypter small. A p i c a l half o f v e i n Sc straight; v e i n M i
n o t strongly c u r v i n g ; anal v e i n n o t reaching the w i n g margin; 504 Fannia canicularis (Linnaeus),
v e i n A 2 and the anal v e i n w o u l d intersect i f A 2 w o u l d contin- e ; Stackelberg 1956
m a |

ue further. Tibia o f h i n d leg w i t h a s u b m e d i a n dorsal bristle.

Biology: T h e larvae are saprophagous and feed m o s t l y a m o n g d e c a y i n g organic materials.
M a n y are surface scrapers, feeding o n m i c r o - o r g a n i s m s o n the surface o f the substrate,
w h i c h include a w i d e range o f vegetable and animal detritus, m u s h r o o m s , nests or bur
r o w s of birds, m a m m a l s and insects, dead insects, m o l l u s c s and vertebrates, animal drop
p i n g s and d u n g , h u m a n faeces, etc. T h e adults occur m a i n l y in or near w o o d l a n d areas and
less i n m o r e o p e n habitats. T h e males o f several species fly i n
s w a r m s . S o m e species h a v e successfully adapted to the h u m a n
e n v i r o n m e n t and are frequently m e t w i t h i n d o o r s . G o o d
e x a m p l e s are Fannia canicularis (Linnaeus) (the lesser h o u s e
fly) and F. scalaris (Fabricius) (the latrine fly).
Identification references: C r o s s k e y & Lane 1993 (synanthrop-
ic species o f Fannia); D ' A s s i s Fonseca 1968 (British Isles);
Gregor 8 t R o z k o s n y 1995 (Central Europe); H e n n i g 1955-1964
(Lindner); R o z k o s n y et al. 1997 (revision); C M P D : Pont 2000;
C A T : Pont 1986a; R o z k o s n y et al. 1997.

(Fungivoridae): n a m e u s e d in the past for M y c e t o p h i l i d a e .

Gasterophilidae (key couplet 173;fig.376)

Systematics: Calyptrate Brachycera; superfamily Oestroidea;
t r a d i t i o n a l l y part o f t h e O e s t r i d a e ; i n E u r o p e 1 g e n u s ,
Gasterophilus, w i t h 6 species. 505 Gasterophilus pecorum (Fabrius);
Characters: M e d i u m sized to large flies (9-16 m m ) . B o d y and Gamin 1969

134
 legs covered w i t h soft, hair-like bristles, g i v i n g t h e m the aspect o f b u m b l e b e e s or bees.
M o u t h o p e n i n g small; m o u t h p a r t s r u d i m e n t a r y ; antenna small. W i n g w i t h v e i n M i
rather straight, slightly c u r v i n g b a c k w a r d , reaching the w i n g m a r g i n b e y o n d the w i n g tip;
l o w e r calypter small. S u b s c u t e l l u m absent; m e r o n lacking strong bristles along its h i n d
margin near the posterior spiracle. Female w i t h a l o n g ovipositor w h i c h can b e folded
b e l o w the a b d o m e n in rest.
Biology: T h e larvae live as parasites in the s t o m a c h and intestine o f horses and d o n k e y s .
T h e eggs are deposited directly o n the h o s t ' s skin and the first larval instar penetrates the
skin and w o r k s its w a y inside the animal, t h e y first migrate to the m o u t h and later to the
stomach and intestine. T h e larva leaves the h o s t along w i t h the e x c r e m e n t and pupates in
the soil. T h e adults possess reduced m o u t h p a r t s , they do n o t feed and are f o u n d near
horses and stables.
Identification references: C o l w e l l et al. 2 0 0 4 (general); G r u n i n 1969 (Lindner), 1989a
(former U S S R ) ; C M P D : M i n a f 2000b; C A T : S o o s 8t M i n a f 1986a.

Helcomyzidae (key couplet 146; fig. 317-320)

Systematics: Acalyptrate Brachycera; superfamily
Sciomyzoidea; traditionally part o f the D r y o m y z i d a e ; 2
species in Europe: Helcomyza ustulata Curtis and H. mediter-
ranea (Loew).
Characters: M e d i u m sized to large (6-11 m m ) , stout, p r e d o m i
nantly grey pruinose flies, densely c o v e r e d w i t h fine setae.
Head large w i t h relatively small eyes; arista bare to pubescent;
ocelli present; Oc-bristles reduced or absent; P-bristles paral
lel to s o m e w h a t d i v e r g i n g ; 1-2 pairs o f F-brisdes, c u r v i n g out
w a r d ; scattered interfrontal setulae present; vibrissae absent.
W i n g u n m a r k e d , crossvein often accompanied b y dark bands;
costa continuous; v e i n Sc complete; crossvein B M - C u present;
cell cup closed. Tibiae w i t h dorsal preapical bristle.
Biology: T h e species o f this small family occur along the coast
where the adults are often found on the beach, making a short 506 Helcomyza ustulata Curtis,
flight when disturbed. T h e larvae live in s e a w e e d w a s h e d u p male; McAlpine 1998c
on the shore and w h i c h has dried to a greater extent than the
seaweed in w h i c h C o e l o p i d a e , Heterocheilidae or Orygma
species are l i v i n g .
Identification references: Stackelberg i 9 8 g y (former U S S R ) ;
C M P D : D . K . M c A l p i n e 1998c; C A T : G o r o d k o v 1984a.

(Heleidae): n a m e used in the past for C e r a t o p o g o n i d a e .

Heleomyzidae (key couplet 92, 117,220; fig. 1 0 , 2 0 1 , 2 5 3 - 2 5 4 )

Systematics: Acalyptrate Brachycera; superfamily
S p h a e r o c e r o i d e a ; t r a d i t i o n a l l y t h i s f a m i l y i n c l u d e s the
C h i r o p t e r o m y z i d a e , C n e m o s p a t h i d a e and T r i x o s c e l i d i d a e ;
s o m e 23 genera and about 150 species in Europe.
Characters: R o b u s t , m i n u t e to large (1.2-12 m m ) , pallid y e l l o w
to dark grey or b r o w n , in s o m e cases reddish flies. Arista bare
or p l u m o s e ; ocelli present; Oc-bristles present; P-bristles c o n
verging; 1 or 2 pair(s) o f F-brisdes, c u r v i n g o u t w a r d or back
w a r d ; interfrontal bristles absent, interfrontal setulae present
or absent; vibrissae present. W i n g clear, tinged or w i t h a pat
tern, especially along the crossveins; costa w i t h a subcostal
break; v e i n Sc complete; crossvein B M - C u present; cell c u p
closed. Tibiae usually w i t h dorsal preapical bristle w h i c h m a y 507 Eccoptomera microps (Meigen),
be small or lacking in the Borboropsinae and in Oldenbergiella. male; Seguy 1950

Biology: T h e larval habitats o f the v a r i o u s subfamilies can be

summarised as f o l l o w s : In the H e l e o m y z i n a e a n d
H e t e r o m y z i n a e the larvae m a i n l y feed o n rotting organic mat-

13 5
 ter and can be found in carrion, excrement, m u s h r o o m s , bracket fungi, animal b u r r o w s ,
nests o f birds, etc. T h e larvae of the Suilliinae h a v e chiefly b e e n found in m u s h r o o m s and
bracket fungi w h i l e Suillia lurida (Meigen) is a w e l l - k n o w n pest in onions and garlic. T h e
adults are often found near the larval habitats, e.g. in w o o d l a n d areas, b u t also in caves and
in or near nests of birds and m a m m a l b u r r o w s .
I d e n t i f i c a t i o n r e f e r e n c e s : G o r o d k o v 1989b (former U S S R ) ; Papp 1998I (Nidomyia); Soos
1981 (Central Europe); W i t h e r s 1978 (Suillia); C M P D : Papp 1998!; C A T : G o r o d k o v 1984c.

( H e l o m y z i d a e ) : n a m e used in the past for H e l e o m y z i d a e .

Hesperinidae (key couplet 26,203; fig. 65-66)

S y s t e m a t i c s : Nematocera; superfamily Bibionoidea; tra
ditionally part o f the Bibionidae; in Europe (Balkan) 1
species, Hesperinus imbecillus (Loew).
C h a r a c t e r s : A slender, dark coloured, small to m e d i u m
sized (4-6 m m ) species w i t h l o n g antennae, legs and
a b d o m e n . Ocelli c o n s p i c u o u s ; antennal insertion near
l o w e r m a r g i n o f the face; antenna w i t h typically elon
gate third s e g m e n t . W i n g darkened, l o n g i n the male,
short in t h e female w h e r e it reaches to halfway d o w n the
s e c o n d abdominal s e g m e n t . Tibiae w i t h apical bristles or
spurs.
B i o l o g y : Little is k n o w n about t h e b i o l o g y o f this small
family. T h e larvae live in rotting w o o d o f d e c i d u o u s 508 Hesperinus imbecillus (Loew),
trees. male; Sguy 1951
I d e n t i f i c a t i o n r e f e r e n c e s : Stackelberg 1 9 8 9 m (former
U S S R ) ; C M P D : K r i v o s h e i n a 1997b; C A T : K r i v o s h e i n a
& M a m a e v 1986.

Heterocheilidae (key couplet 146;fig.321)

S y s t e m a t i c s : Acalyptrate Brachycera; superfamily
S c i o m y z o i d e a ; traditionally part o f the D r y o m y z i d a e ; in
E u r o p e , a l o n g N o r t h Sea a n d Baltic S e a , 1 s p e c i e s ,
Heterocheila buccata (Fallen).
C h a r a c t e r s : M e d i u m sized (6-9 m m ) , stout, largely grey
ish b r o w n species, densely covered w i t h fine setae.
Head large w i t h small eyes; arista bare to pubescent;
ocelli present; Oc-bristles present; P-bristles m o r e or
less parallel; 3 pairs o f F-bristles, c u r v i n g o u t w a r d ; scat
tered interfrontal setulae p r e s e n t ; v i b r i s s a e absent. 509 Heterocheila buccata (Fallen),
W i n g u n m a r k e d ; costa continuous; v e i n Sc complete; male; McAlpine 1998d
crossvein B M - C u present; cell c u p closed. Tibiae w i t h
dorsal preapical bristle.
B i o l o g y : T h e sole representative o f this family in Europe,
Heterocheila buccata, is a species of coastal habitats. T h e
larvae d e v e l o p in s e a w e e d w a s h e d u p o n the shore; t h e
adults are usually found near t h e larval b r e e d i n g sites.
I d e n t i f i c a t i o n r e f e r e n c e s : Stackelberg i 9 8 9 y (former
U S S R ) ; C M P D : D . K . M c A l p i n e I9g8d; C A T : G o r o d k o v
1984b (as part o f the H e l e o m y z i d a e ) .

Hilarimorphidae (key couplet 57;fig.122)

S y s t e m a t i c s : L o w e r m u s c o m o r p h Brachycera; superfam
ily A s i l o i d e a ; traditionally part o f the Rhagionidae; 2
species in Europe: Hilarimorpha singularis Schiner and
H. tristis Egger.
C h a r a c t e r s : S m a l l to m e d i u m sized (2-7 m m ) , stout, 510 Hilarimorpha spec, male; Webb
b r o w n or black coloured flies r e s e m b l i n g species o f 1974

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 Hilara (Empididae), w i t h pubescence and a characteristic venation. Male holoptic; anten
na short, w i t h 4 distinct segments of w h i c h the third is the largest, and w i t h a small apical
style; m o u t h p a r t s small. W i n g clear or tinged, i n s o m e cases w i t h dark markings along
vein R i ; discal cell absent; v e i n s R4+5 and M1+2 forked in m o r e or less similar w a y s ; cell
cup closed. Legs w i t h o u t bristles or spurs and w i t h o u t e m p o d i u m .
Biology: T h e larvae o f this small family are u n k n o w n . T h e adults w e r e chiefly collected
near bodies of water, for example near p u d d l e s in cart tracks, at o p e n sandy spots near fen-
land, or o n w i l l o w s alongside creeks.
Identification references: Lindner 1925 (species); Nartshuk 1989c (former U S S R ) ; W e b b
1974 (revision); C M P D : N a g a t o m i 1997c; C A T : Majer ig88d.

Hippobosddae (key couplet 1 6 9 , 209; fig. I74a-b, 370-371,

438,442-443)
Systematics: Calyptrate Brachycera; superfamily
Hippoboscoidea; 11 genera and about 30 species in Europe.
Characters: Small to m e d i u m sized (2.5-10 m m ) flies. M o s t
species are dorsoventrally flattened as an adaptation to their
ectoparasitic m o d e o f life. Head closely adpressed t o thorax
and eyes w e l l - d e v e l o p e d . M o s t species possess w e l l - d e v e l
oped w i n g s and are able fliers; s o m e have reduced w i n g s or
shed their w i n g s after finding a suitable host. W i n g venation
511 Lipoptena cervi (Linnaeus),
reduced and concentrated at the w i n g base and anterior region. female; Hutson 1984
Legs short and p o w e r f u l ; coxae o f m i d legs and usually also
hind legs far apart; tarsi w i t h h o o k - l i k e c l a w s , enabling these insects to cling to the hairs
or feathers o f their hosts.
Biology: A l l species are ectoparasites o f birds or m a m m a l s and feed o n b l o o d . T h e larvae
develop inside the a b d o m e n o f the female fly and are deposited o n e b y one, i m m e d i a t e l y
followed b y pupation, generally o n the g r o u n d or in the soil. M o s t species visit a limited
n u m b e r o f host species b u t s o m e species are found o n species o f birds sharing the same
habitat. Flightless species can find n e w hosts b y contact b e t w e e n hosts or t h r o u g h infec
tion at nest sites.
Identification references: B o r o w i e c 1984 (Poland); Biittiker 1 9 9 4 (Switzerland); G r u n i n
1989b (former U S S R ) ; H u t s o n 1984 (British Isles); Maa 1963 (genera); S c h u u r m a n s
S t e k h o v e n jr. ck. V a n d e n Broek 1969 (Netherlands); T h e o d o r 8t O l d r o y d 1 9 6 4 (Lindner);
C M P D : n o contribution; C A T : Soos &. Hurka 1986.

Hybotidae (key couplet 77, 80, 212; fig. 9 6 , 154, 1 6 7 - 1 6 9 , 1 7 1 ,

173.446)
Systematics: L o w e r e r e m o n e u r a n B r a c h y c e r a ; s u p e r f a m i l y
Empidoidea; traditionally part o f the Empididae; s o m e 30 g e n
era and about 4 4 0 species in Europe. For a discussion about the
p h y l o g e n y and classification see Sinclair 8t C u m m i n g 2006.
Characters: M i n u t e to m e d i u m sized (1-6 m m , rarely larger)
flies, usually dark b u t colour variable, y e l l o w to black. Eyes rel
atively large, dichoptic or holoptic, w i t h an incision near the
antennae; antenna short or w i t h a large third s e g m e n t and l o n g
arista; m o u t h p a r t s v a r y i n g b e t w e e n short ( O c y d r o m i i n a e ) to
long and directed straight or obliquely d o w n w a r d (Hybotinae)
or d i r e c t e d d o w n w a r d ( T a c h y d r o m i i n a e ) . W i n g clear or
tinged, in s o m e cases w i t h markings, along the costa and
around or in b e t w e e n v e i n s R i and R2+3 in particular; cell d m
absent (Tachydromiinae) or present. Legs long, slender and 512 Synechus muscarius (Fabricius),
often partially adapted to catching and h o l d i n g prey; femora o f male; Chvla 1983
fore legs a n d / o r m i d legs s w o l l e n in the T a c h y d r o m i i n a e w h i l e
in the other subfamilies those o f the h i n d legs are s w o l l e n .
Biology: T h e larvae o f only a f e w species are k n o w n . T h e y live in soil, rotting w o o d , organ
ic matter, d u n g , etc., and prey o n small invertebrates. T h e adults prey o n small insects and
other a r t h r o p o d s , c a t c h i n g their p r e y i n f l i g h t ( H y b o t i n a e , O c y d r o m i i n a e , s o m e

137
 Tachydromiinae) or w h i l e w a l k i n g or r u n n i n g about o n the ground, o n tree trunks, leaves,
etc. (most Tachydromiinae). O n l y a f e w species visit flowers to feed o n nectar and pollen.
S w a r m i n g has b e e n observed in the O c y d r o m i i n a e .
Identification references: C h v a l a 1975 ( T a c h y d r o m i i n a e ) , 1983 ( H y b o t i n a e ) , 1989a
(Platypalpus); C o l l i n 1961 (British Isles); V a n der G o o t 1 9 9 0 (Hybos Netherlands,
B e l g i u m , L u x e m b o u r g ) ; G o r o d k o v 8c K o v a l e v 1989 (former U S S R ) ; Papp & F o l d v a r i 2 0 0 1
(Central Europe); Sinclair 8t C u m m i n g 2006 (classification); C M P D : n o contribution;
C A T : C h v a l a &. K o v a l e v 1989.

(Hyperoscelididae): n a m e u s e d i n the past for Canthyloscelidae.

Hypodermatidae (key couplet 174; fig. 377-378)

Systematics: Calyptrate Brachycera; superfamily Oestroidea;
traditionally part o f the Oestridae; 4 genera and 8 species in
Europe.
Characters: Large to v e r y large (10-22 m m ) flies. B o d y w i t h scant
short bristles or b o d y and legs w i t h soft pile o f hairs, thereby
resembling bees or bumblebees. M o u t h o p e n i n g small, m o u t h -
parts rudimentary; antenna small; the ends o f the ptilinal
suture, b e l o w the antennae, n o t strongly c u r v e d t o w a r d each
other. V e i n M i w i t h part b e y o n d crossvein D M - C u gradually
c u r v i n g or m o r e or less straight t o w a r d the costa; cell 14+5 o p e n
or closed at the w i n g margin, w i t h o u t a petiole; l o w e r calypter
large. Subscutellum narrow, s w o l l e n ; m e r o n usually w i t h bris
tles along its h i n d margin, near the posterior spiracle.
Biology: T h e larvae live as s u b c u t a n e o u s endoparasites in m a m - 5 1 3
Hypoderma diana Brauer; Grunin
1 9 6 5
mals, especially deer and cattle b u t also occasionally m a m m a l s
(e.g. horses, guinea pigs). M o s t species are specialised in o n e or
o n l y a f e w h o s t species. T h e eggs are deposited o n the skin o f the host and the first instar
larva penetrates the skin. S e c o n d and third instar larvae accumulate o n the host's back and
f o r m large warbles. Just prior to pupation the fully g r o w n larva leaves the host through
the skin, pupating in the soil. T h e adults h a v e reduced m o u t h p a r t s and do n o t feed. T h e y
are often found sitting o n w a l l s , plants, etc., or near their host. T h e species that are of eco
n o m i c importance to cattle breeding have b e e n virtually exterminated during the second
half o f the 20th century.
Identification references: C o l w e l l et al. 2 0 0 4 (general); G r u n i n 1964-1969 (Lindner),
19896 (former U S S R ) ; Zeegers 1992 (Netherlands); C M P D : M i n a f 2 0 0 o d ; C A T : Soos 8t
M i n a f 1986c.

(Itonididae): n a m e u s e d in the past for C e c i d o m y i i d a e .

Keroplatidae (key couplet 35; fig. 77-80)

Systematics: Nematocera; superfamily Sciaroidea, traditionally
part o f the M y c e t o p h i l i d a e ; s o m e 16 genera and about 110
species in Europe.
Characters: M e d i u m to large (4-15 m m ) , slender and delicate to
stout, in s o m e cases c o n s p i c u o u s l y coloured Nematocera w i t h
characteristic v e n a t i o n (e.g. c r o s s v e i n R - M absent). Head
small; m o u t h p a r t s s o m e t i m e s elongated; ocelli present; eyes
dichoptic; antenna w i t h 16-17 segments; antenna ranging from
relatively short, consisting o f laterally c o m p r e s s e d segments
to v e r y slender and longer than the body. T h o r a x vaulted.
W i n g relatively w i d e , clear or w i t h markings or crossbands, in
Macrocera w i t h a striking anal angle. C o x a e elongate; tibiae
w i t h apical bristles or spurs.
Biology: T h e larvae are usually found in m o i s t and dark spots in 514 Cerotelion lineatus (Fabricius),
w o o d l a n d . Keroplatinae larvae often occur u n d e r n e a t h brack- male; Matile 1990

138
 et fungi w h e r e they construct a w e b to collect spores ( m y c o p h a g o u s species) or catch
small invertebrates (predatory species). T h e latter group includes species p r o v i s i o n i n g the
strands of their w e b w i t h acid fluids in order to kill their prey. Macrocera larvae are found
in rotting w o o d , u n d e r n e a t h tree bark and b e l o w tree trunks. T h e y are covered i n a thin
layer o f m u c u s and feed o n animal detritus or p r e y u p o n small invertebrates. T h e adults
are found near the larval habitats; s o m e species visit flowers w h e r e t h e y feed o n nectar;
species o f Macrocera are often found o n the u p p e r side o f leaves, standing h i g h o n their
legs and h o l d i n g their w i n g s slightly spread out.
I d e n t i f i c a t i o n references: Chandler & R i b e i r o 1995 (Atlantic islands); H u t s o n et al. 1980
(British Isles); Kurina 8s. Schacht 2003 (synopsis identification references); Matile 1 9 9 0
(revision, excluding Orfeliini); Stackelberg 1989, 1989k (former U S S R ) ; A . I . Z a i t z e v
(Eastern Europe); C M P D : Soli et al. 2000; C A T : K r i v o s h e i n a 8s. M a m a e v 1988b.

(Larvaevoridae): n a m e used in the past for Tachinidae.

Lauxaniidae (key couplet 91,138; fig. 186-187,193,200,303-304)

Systematics: Acalyptrate Brachycera; superfamily
Lauxanioidea; s o m e 18 genera and about 160 species i n Europe.
Characters: Small to m e d i u m sized (2-7 m m ) , often rather
p l u m p , dull or partially lustrous flies; b o d y colour v a r y i n g
from y e l l o w to b r o w n or black, or w i t h a c o m b i n a t i o n o f
colours. Arista bare, pubescent, or w i t h l o n g rays; ocelli pres
ent; Oc-bristles present; P-bristles c o n v e r g i n g , i n rare cases
parallel (e.g. a m o n g a f e w s p e c i m e n s o f Cnemacantha mus
caria (Fallen)); 1-2 pairs o f F-bristles, c u r v i n g b a c k w a r d , the
l o w e r pair i n s o m e cases c u r v i n g inward; interfrontal bristles
absent; vibrissae absent b u t s o m e genera, e.g. Trigonometopus,
w i t h strong bristles near the vibrissal angle. W i n g usually
unmarked, in a n u m b e r o f species w i t h spots along the v e i n s ,
in s o m e cases entirely marked; costa c o n t i n u o u s ; v e i n Sc c o m
plete; crossvein B M - C u present; cell c u p closed. U s u a l l y all 515 Eusapromyza multipunotata (Fallen);
tibiae w i t h dorsal preapical bristle b u t in a f e w cases tibia o f male; Papp 1981
hind legs w i t h o u t such bristle.
Biology: A s far as k n o w n the larvae live o n m i c r o - o r g a n i s m s in
bracket fungi and m u s h r o o m s , in rotting vegetable matter, in
s o m e cases as miners or in nests o f birds. T h e majority o f species are found i n shaded
spots in m o i s t w o o d l a n d s , marshes, banks, etc. S o m e species of Homoneura, Minettia and
Calliopum are adapted to drier, m o r e o p e n habitats like dunes and grasslands.
Identification references: M e r z 2002 (genera Central Europe);
Shatalkin 2000 (species); Stackelberg i g S g a d (former U S S R ) ;
Stuckenberg i g 7 i (genera); C M P D : Papp 8c Shatalkin i g g 8 ;
C A T : Papp 1984a.

(Leptidae): n a m e used i n the past for Rhagionidae.

(Leptoconopidae): part o f the C e r a t o p o g o n i d a e .
(Leptogastridae): part o f the A s i l i d a e .
(Lestremiidae): part o f the C e c i d o m y i i d a e .
(Limnobiidae): n a m e used in the past for Limoniidae.

Limoniidae (key couplet 7 , 1 9 6 , i g 8 ;fig.31-32,430-431)

Systematics: Nematocera; superfamily Tipuloidea; traditional
ly part o f the Tipulidae; s o m e 7 0 genera and about 560 species
in Europe.
Characters: Pale y e l l o w to black, small to v e r y large (2-11 m m , a
single species u p to 30 m m ) Nematocera w i t h slender anten 516 Limonia phragmitidis (Schrank),
nae, legs and a b d o m e n . R o s t r u m usually short and w i t h o u t male; Papp & Schumann 2000

139
 nasus; ocelli absent; antenna usually w i t h 14-16 segments, in s o m e cases (Hexatoma) less
or (Ludicia) more. T h o r a x w i t h a V - s h a p e d transverse suture. W i n g w i t h 2 anal veins; in
s o m e cases w i n g w i t h characteristic m a r k i n g s or w i n g reduced in females or in b o t h sexes
(Chioned).
Biology: T h e larvae are generally found in various aquatic and semi-aquatic habitats. S o m e
species are terrestrial, living i n m o s s e s , m u s h r o o m s , bracket fungi, or in d y i n g or decay
ing w o o d . A f e w species are found i n brackish or saline habitats. M o s t larvae feed o n rot
ting vegetable matter, algae, m o s s e s , m u s h r o o m s , etc. T h e subfamily Limnophilinae has
m a n y species o f w h i c h the larvae prey o n insect larvae, o n w o r m s and other small inver
tebrates. In the subfamily Limoniinae m o s t larvae live in a tube m a d e o f strands they have
spun and in w h i c h t h e y also pupate. A d u l t s are found in several, usually m o i s t habitats,
often near those inhabited b y their larvae. Males often fly in s w a r m s .
Identification references: D i e n s k e 1987 (genera and subgenera); Geiger 1986 (Limoniinae
Switzerland); R e u s c h 8t O o s t e r b r o e k 1997 (genera and subgenera); Savchenko 1989c,
i 9 8 g d (former U S S R ) ; Stubbs 1 9 9 4 - 1 9 9 9 (British Isles); C M P D : n o contribution; C A T :
S a v c h e n k o et al. 1992; Oosterbroek 2006.

(Liriopeidae): n a m e used in the past for Ptychopteridae.

Lonchaeidae (key couplet 83;fig.182-183)

Systematics: A c a l y p t r a t e B r a c h y c e r a ; s u p e r f a m i l y
Tephritoidea; 8 genera and about 100 species in
Europe.
Characters: U s u a l l y stout, metallic g r e e n or b l u i s h
black, small to m e d i u m sized (3-6 m m ) flies. Head
large, h e m i s p h e r i c a l , s h o w i n g considerable sexual
d i m o r p h i s m w i t h frons narrower in the male; lunula
large; arista bare or w i t h shorter to longer pubescence;
ocelli present; Oc-bristles present; P-bristles w e a k ,
d i v e r g i n g ; 1 pair o f F-bristles, situated at the level o f 5 1 7 L o n c h a e a c h o r e a ( F a b r i c i u s ) j

the ocellar triangle and c u r v i n g b a c k w a r d ; scattered female; Colyer & Hammond 1968
interfrontal setulae present; vibrissae absent b u t i n
s o m e genera w i t h a series o f v i b r i s s a - l i k e bristles near the vibrissal angle. W i n g
u n m a r k e d ; cell c w i d e ; v e i n Sc complete; costa w i t h a subcostal break; crossvein B M - C u
present; cell c u p closed. Haltere dark b r o w n to black. Tibiae w i t h o u t dorsal preapical bris
tle or present o n l y o n tibia o f m i d legs. A b d o m e n short, w i d e , dorsoventrally flattened;
the female w i t h a lanceolate, i n c o m p l e t e l y retractable ovipositor.
Biology: T h e larvae of m o s t species are found in m o u l d e r i n g w o o d and underneath the bark
o f d y i n g or dead trees, w i t h several species associated w i t h certain tree species. In other
species the larval d e v e l o p m e n t takes place in rotting vegetable matter such as onions,
conifer seeds, fruits, vegetables, galls o n grasses or other substances. T h e larvae o f m o s t
species feed o n vegetable matter, a l t h o u g h s o m e are carrion eaters or prey o n the larvae o f
other insects, e.g. those o f beetles. Males congregate in s w a r m s in o p e n i n g s in forests.
Lonchaeidae are u s u a l l y less frequently encountered and in m a n y cases h a v e been
obtained b y rearing; h o w e v e r , they are b y n o m e a n s rare as is clear from the larger series
obtained b y b e e r / w i n e traps and Malaise traps.
Identification references: M a c G o w a n 8t R o t h e r a y 2000 (Lonchaea); J.F. M c A l p i n e 1 9 6 4
(revision), 1975 (Lonchaea); J.F. M c A l p i n e 8t Steyskal 1982 (genera); M o r g e 1959-1974
(review); Stackelberg 1989a! (former U S S R ) ; C M P D : n o contribution; C A T : K o v a l e v &
M o r g e 1984.

Lonchopteridae (key couplet 68; fig. 146-147)

Systematics: L o w e r Cyclorrhapha; superfamily Lonchopteroidea; in Europe 2 genera,
Lonchoptera and Neolonchoptera, w i t h about 13 species.
Characters: Small (2-5 m m ) , y e l l o w to b r o w n i s h black flies w i t h strong bristles and long,
subacute w i n g s . Eyes broadly dichoptic; third antennal s e g m e n t rounded, w i t h a long
arista; 1 pair o f strong, d i v e r g i n g bristles just above the antennae. W i n g clear to tinged,
rarely w i t h markings; venation characteristic and sexual dimorphic: in male anal v e i n

140
 reaching the hind margin, in the female e n d i n g in v e i n C u A i .
Biology: T h e larvae feed o n rotting vegetable matter, perhaps
including m u s h r o o m s and bracket fungi. T h e y can be found
a m o n g dead leaves, in plant debris and other d e c a y i n g materi
al in various habitats ranging from gardens to w o o d l a n d and
m e a d o w s . T h e adults feed o n nectar and are in s o m e cases
found in large n u m b e r s , often in m o i s t places alongside r u n
n i n g or stagnant w a t e r s , in w o o d l a n d s , m a r s h e s , s o m e species
in the tidal z o n e ; other species prefer drier habitats, s o m e are
ubiquitous and occur in a variety o f habitats from l o w l a n d s to
high m o u n t a i n areas. M o s t species are bisexual, b u t at least L.
bifurcata (Fallen) is parthenogenetic in m o s t areas of its nearly
J r
_ , J
.\ ' 518 Lonchoptera lutea Panzer, male;
c o s m o p o l i t a n range. , S 6 g u y 9 5 0

Identification references: B a h r m a n n & B e l l s t e d t 1 9 8 8

( G e r m a n y ) ; B a r t a k i g 8 6 (Europe); D e Meijere 1906 (species);
S m i t h 1969a (British Isles); Stackelberg 19890 (former U S S R ) ; Vaillant 1989 (species),
2002 (species); C M P D : Bartak 1998; C A T : A n d e r s s o n 1991.

(Lycoriidae): n a m e u s e d in t h e past for Sciaridae.

(Macroceridae): part o f the Keroplatidae.
(Manotidae): part o f the M y c e t o p h i l i d a e .

Megamerinidae (key couplet 154; fig. 335-336)

Systematics: Acalyptrate Brachycera; superfamily D i o p s o i d e a ;
in E u r o p e 1 w i d e s p r e a d s p e c i e s , Megamerina dolium
(Fabricius).
Characters: T h e o n l y European species, Megamerina dolium, is
m e d i u m sized (6-9 m m ) , elongate, black lustrous w i t h pale to
reddish femora. Head small and spherical; arista pubescent;
ocelli present; O c - , P-, F-, interfrontal bristles and vibrissae
absent. W i n g pale b r o w n , u n m a r k e d ; costa c o n t i n u o u s ; v e i n
Sc c o m p l e t e ; c r o s s v e i n B M - C u p r e s e n t ; cell c u p c l o s e d .
Femora o f hind legs s w o l l e n , w i t h 2 r o w s o f spines o n the
l o w e r side; tibiae w i t h o u t dorsal preapical bristle.
Biology: T h e larvae live under the bark o f d y i n g or dead decidu
ous trees and are predatory or n e c r o p h a g o u s o n t h e larvae o f
other insects. T h e adults are largely found near the larval habi
tats, chiefly o n tree trunks and leaves. 519 Megamerina dolium (Fabricius),
female; Seguy 1934
Identification references: V a n der G o o t & V a n V e e n 1996
( N o r t h w e s t e r n Europe); Stackelberg 19891; C M P D : n o contri
bution; C A T : Nartshuk 1984a.

(Melusinidae): n a m e used in the past for Simuliidae.

Micropezidae (key couplet 158; fig. 339-341)

Systematics: Acalyptrate Brachycera; superfamily Nerioidea; 5
genera and about 22 species in Europe.
Characters: Strikingly slender flies o f small to large size (3-16
m m ) w i t h n a r r o w w i n g s and l o n g legs. Head small, r o u n d e d to
elongate; antenna small w i t h arista bare or pubescent; ocelli
present; Oc-bristles absent; P-bristles d i v e r g i n g or absent; 0-3
pairs o f F-bristles, i f present, c u r v i n g forward or b a c k w a r d ;
interfrontal brisdes absent; vibrissae absent. W i n g clear to
fumosely patterned; costa continuous; v e i n Sc c o m p l e t e , its
e n d i n g in t h e costa m a y be close to that o f v e i n R i ; crossvein
B M - C u present or absent (in t h e M i c r o p e z i n a e ) ; cell c u p
closed. Tibiae w i t h o u t dorsal preapical bristle. 520 Raineria calceata (Fallen), female;
Biology: With respect t o their biology, the European Mihalyi 1972

141
 Micropezidae can b e divided in three groups. Species of Micropeza have p h y t o p h a g o u s lar
vae feeding in the root nodules of l e g u m i n o u s plants and are associated w i t h open habitats;
species of Rainieria d e v e l o p in the rotting w o o d o f deciduous trees and are restricted to old
forests; the majority of the other species have saprophagous larvae living in decayed vege
tation, old manure or fungi and their adults usually prefer h u m i d forests along river and
creeks. A d u l t Micropezidae prey o n small insects and other invertebrates and can be found
on flowers, leaves, rotting fruit, excrement, etc.
Identification references: A n d e r s s o n 1989 (Scandinavia); V a n der G o o t 8t V a n V e e n 1996
( N o r t h w e s t e r n Europe); G r e v e & Nielsen 1991 ( N o r w a y ) ; M e r z 1997 (Switzerland);
O z e r o v 1991 (Calobatinae); Rohacek & Bartak 1990 (Central Europe); Stackelberg ig8gr,
1989s (former U S S R ) ; C M P D : n o contribution; C A T : Soos 1984a.

Micro phoridae (key couplet 75;fig.161)

Systematics: L o w e r e r e m o n e u r a n B r a c h y c e r a ; s u p e r f a m i l y
Empidoidea; traditionally part o f the Empididae, considered
part o f the D o l i c h o p o d i d a e b y Sinclair 8t C u m m i n g 2006; 4
genera and about 16 species in Europe.
Characters: Minute to small (1.5-3 m m ) , greyish to black flies.
Head and eyes relatively large; male holoptic; third antennal
segment elongate, w i t h terminal arista; m o u t h p a r t s s o m e w h a t
elongate, directed straight forward or obliquely d o w n w a r d .
W i n g clear or tinged, in s o m e cases w i t h a stigma spot; costa
surrounding the w i n g entirely; cell c u p small; anal v e i n short or
absent; cell d m present w i t h 3 v e i n s from cell d m to the w i n g
margin. Legs short, hind leg in s o m e cases compressed and
w i t h conspicuous w i d e n e d tibia and first tarsal segment.
521 Microphor holosericeus (Meigen),
Biology: T h e larvae o f this small family are little k n o w n ; they male; Collin 1961
are a s s u m e d to p r e y o n organisms l i v i n g in rotting organic
material in sandy soil in particular. T h e adults are predators b u t can also frequently be
found o n flowers w h e r e t h e y feed o n p o l l e n and nectar. T h e females o f the genus
Microphor are k n o w n to feed o n insects caught in w e b s o f spiders. B o t h males and females
fly i n s w a r m s . Species o f Schistosoma and Parathalassius prefer sandy habitats, coastal
areas and deserts.
Identification references: C h v a l a 1983-19883 (revision); Collin 1961 (British Isles); Gatt
2003 (Mediterranean Microphorella); G o r o d k o v & K o v a l e v 1989 (former U S S R ) ; Sinclair
8t C u m m i n g 2006 (classification); C M P D : n o contribution; C A T : Chvala 1989c.

MHichiidae (key couplet 127-128; fig. 273-278)

Systematics: A c a l y p t r a t e Brachycera; superfamily Carnoidea; 9
genera and about 4 5 species in Europe.
Characters: M i n u t e to m e d i u m sized (1-6 m m ) , usually dark
coloured, s o m e t i m e s lustrous black flies a n d / o r w i t h e x t e n
sive silvery dusting. M o u t h p a r t s s o m e t i m e s elongate; arista
bare to pubescent; ocelli present; Oc-bristles present; P-bris-
tles parallel or c o n v e r g i n g ; 2-7 pairs o f F-bristles, nearly a l w a y s
the l o w e r 2 pairs c u r v i n g i n w a r d , the u p p e r pairs c u r v i n g for
w a r d , o u t w a r d , or b a c k w a r d ; interfrontal bristles present, in
s o m e cases in distinct r o w s , in other cases m o r e scattered; v i b -
rissae or vibrissa-like bristles present, s o m e t i m e s v e r y thin.
W i n g u n m a r k e d ; costa w i t h humeral and subcostal breaks;
v e i n Sc incomplete; crossvein B M - C u usually present, in s o m e 522 Madiza glabra Fallen, male;
cases v a g u e or absent; cell c u p closed. Tibiae w i t h o u t dorsal Stackelberg 1956
preapical bristle.
Biology: T h e species in this family s h o w diverse m o d e s of life; m o s t species feed o n organ
ic matter or animal w a s t e , several species feed o n d u n g , sap e x u d i n g from tree w o u n d s ,
seeds, in nests of birds, in nests of ants and other social insects, etc. A d u l t s can be expect
ed in all sorts o f habitats, including indoors; they also regularly visit flowers, carrion and

142
 dung. A d u l t s o f s o m e species are k n o w n to attach t h e m s e l v e s to spiders and predatory
insects (e.g. Asilidae) in order to suck juices from t h e prey caught b y these animals.
Identification references: B r a k e 2 0 0 0 ( k e y W o r l d g e n e r a ) ; S a b r o s k y 1983
(Desmometopa); Stackelberg 1 9 8 9 8 V (former U S S R ) ; C M P D : Papp & W h e e l e r 1998; C A T :
Papp 1984J.

Musddae (key couplet 180,182-183; fig. 1 0 4 , 1 7 8 , 3 8 5 , 3 9 6 - 3 9 8 ,

407-408)
Systematics: Calyptrate Brachycera; superfamily Muscoidea;
s o m e 4 5 genera and about 575 species in Europe.
Characters: Small to large (2-18 m m ) flies, usually coloured grey
to black, in s o m e cases y e l l o w i s h b r o w n or w i t h an extensive
green to blue metallic lustre. M o u t h p a r t s w e l l d e v e l o p e d ;
o c c i p u t lacking w h i t e or pallid hairs; arista bare t o l o n g
p l u m o s e ; interfrontal bristles usually absent, i n s o m e cases
present in female (e.g. i n Azelia). S c u t e l l u m bare underneath;
subscutellum absent; meral brisdes virtually a l w a y s absent
although Eginia has a patch o f rather strong bristles and
Graphomya has a r o w o f rather w e a k bristles. W i n g w i t h api
cal half of v e i n Sc distinctly c u r v i n g ; v e i n M i straight or w i t h a
faint to distinct forward c u r v e ; cell r4+5 o p e n ; anal v e i n usual- 523 stomoxys calcitrans (Linnaeus
ly w e l l d e v e l o p e d , b u t n o t continuing to w i n g margin; v e i n A 2 female; Seguy 1951
and the anal v e i n w o u l d n o t intersect i f virtually continued
further (with the exception o f s o m e species o f Azelia). Tibia o f h i n d leg lacking a s u b m e -
dian dorsal bristle.
Biology: T h e f o l l o w i n g life m o d e s o f the larvae are k n o w n : carnivorous (Phaoniinae,
Mydaeinae, Coenosia, Limnophora, Lispe, Graphomya), saprophagous in decaying organ
ic matter (vegetable, animal, or excrement) w h e r e t h e third instar m a y or m a y n o t b e car
n i v o r o u s (Muscini, S t o m o x y n i , Hydrotaea) or, rarely, p h y t o p h a g o u s (Atherigona). T h e
larvae of Phaonia exoleta (Meigen) are k n o w n to live i n rot holes in trees w h e r e they feed
on the larvae o f Culicidae; those of Limnophora riparia (Fallen) live in r u n n i n g w a t e r and
the splash z o n e s along canals, larvae o f other genera are semi-aquatic (e.g. Limnophora,
Lispe, Graphomya). M a n y species h a v e adapted to h u m a n e n v i r o n m e n t s . T h e adults o f
m o s t species feed o n nectar b u t s o m e species are predators (Coenosiinae), feed o n b l o o d
( S t o m o x y n i ) or o n e x u d i n g (animal) w o u n d liquor (Hydrotaea); t h e latter t w o groups are
vectors of diseases. Musca autumnalis D e Geer is a species that can occur indoors in large
n u m b e r s from late s u m m e r o n w a r d , seeking shelter for t h e winter.
Identification references: C o u r i & P o n t 1 9 9 9 (genera C o e n o s i i n i ) ; C r o s s k e y 1993b
(species feeding o n b l o o d ) ; C r o s s k e y & Lane 1993 (synanthropic species o f Musca);
D ' A s s i s Fonseca 1968 (British Isles); Gregor et al. 2002 (Central Europe); H e n n i g 1955-
1964 (Lindner); K a b o s 1 9 6 4 (Netherlands); R o z k o s n y 8t Gregor 1997, 2 0 0 4 (species);
Stackelberg 1 9 8 9 ^ ; Oosterbroek 8c D e Jong 2005 (Eginia);
Z i m i n & E l b e r g i g 8 9 (former U S S R ) ; C M P D : n o contribution;
C A T : Pont 1986b.

(Musidoridae): n a m e used in the past for Lonchopteridae.

Mycetobiidae (key couplet 30; fig. 24,72)

Systematics: Nematocera; superfamily A n i s o p o d o i d e a ; tradi
tionally part of the A n i s o p o d i d a e ; i n Europe 4 species in 2 g e n
era, Mycetobia (3) and Trichomycetobia (1).
Characters: Small to m e d i u m sized (4-7 m m ) , dark coloured
Nematocera. Head small and almost spherical; ocelli present;
eyes almost holoptic; antenna w i t h 16 simple, u n i f o r m seg
ments. W i n g clear, w i t h anal lobe. Legs rather sturdy b u t long,
often less dark; coxae elongated; tibiae w i t h apical bristles or
spurs. 524 Mycetobia pallipes Meigen,
male; Krivosheina 1997d

143
Biology: The larvae usually occur in small groups and are found in moist habitats in rotting
or fermenting organic matter, such as underneath the bark or in wood of dead deciduous
and coniferous trees, in exuding tree sap, in rot holes filled with organic matter, etc. The
adults mainly occur near the larval habitats.
Identification references: Mamaev 1989a (former USSR); Pedersen 1971 (Scandinavia);
CMPD: Krivosheina I997d; C A T : Mamaev 8 t Krivosheina
1988b.

Mycetophilidae (key couplet 3 2 , 2 0 4 ;fig.75)

Systematics: Nematocera; superfamily Sciaroidea; traditionally
this family includes the Bolitophilidae, Diadocidiidae,
Ditomyiidae and Keroplatidae; some 7 0 genera and about 9 4 5
species in Europe.
Characters: Slender to relatively stout Nematocera, minute to
large (2-15 mm) but more usually small to medium sized (4-8
mm). The colour varies from pale, dull yellow to brown or black.
Head small; eyes dichoptic; 2 or 3 ocelli present; antenna usual
ly with 16 segments but sometimes segments close together
resembling a single third segment; mouthparts short, usually
half as long as the height of the head, but sometimes longer or
even extremely elongate as in Gnoriste. Thorax vaulted. Wing 5
Mycetophila fungorum (De Geer),
2 5

usually clear, in some cases tinged or with markings; venation ' Mihalyi 1972
f e m a l e

characteristic (vein CuAi and stem of veins Mi and M2 not con

nected or connected as far up as crossvein H); anal lobe absent or weak. Legs with elongate
coxae; tibiae with long apical spurs.
Biology: The larvae of most species feed on mushrooms, bracket fungi or mycelium in
moist, dead wood or in other moist organic substrates. They rarely are predators or feed
on organic detritus in nests of birds, on mosses or on liverworts (Hepaticae). Some larvae
live in a web they have spun themselves underneath tree trunks, bark, mushrooms and
bracket fungi, etc. The adults ("fungus gnats") occur chiefly in moist habitats, in wood
lands in particular although some species prefer rather different habitats like grassland,
marshes, moist heather, etc. They often are active in the evening while during the day they
hide in dark and shaded spots; some species are mainly found in caves and in burrows of
rodent. Some species visit flowers and are important as pollinators of certain plants.
Identification references: Chandler 8 t Ribeiro 1995 (Atlantic islands); Hutson et al. 1980
(British Isles, excluding Mycetophilinae); Kurina & Schacht 2003 (synopsis identification
references); Ostroverkhova 8 t Stackelberg 1989; Stackelberg 1989I (former USSR); A.I.
Zaitzev (Eastern Europe, 5 subfamilies), 2003 (Eastern Europe, Manotinae and
Mycetophilinae); CMPD: S0H et al. 2 0 0 0 ; CAT: Hackman et al. 1988; Krivosheina &
Mamaev 1988c (Manotinae).

Mydidae (key couplet 56; fig. 120-121)

Systematics: Lower muscomorph Brachycera; superfamily
Asiloidea; in Europe 4 genera and 7 species.
Characters: Large to very large (10-35 nam), dark coloured flies
with characteristic head and wing venation, and with bristles
on the legs only. Eyes dichoptic; head dorsally concave in
between the eyes, as in the Asilidae; antenna with 4 segments,
the terminal segment swollen. Wing clear or tinged, veins R 4 ,
R 5 and M curving forward, hence at least vein R 4 (in European
species also vein R5) ending in vein Ri; cell 1113 and cell cup
closed. Legs long and powerful, tibiae with apical spurs;
empodium bristle-like or absent.
Biology: The larvae prey on the larvae of Coleoptera living in
wood and the soil. The males in particular feed on nectar but
in some species the mouthparts are so much reduced that it is
likely that they are unable to feed at all. Mydidae mainly occur
in open, hot, dry, sandy habitats.

144
Identification references: Carles-Tolr 2006; CMPD: Richter
1997b; CAT: Richter 8t Zaitzev 1988.

MythCOmyiidae (key couplet 66; fig. 101,141-142)

Systematics: Lower muscomorph Brachycera; superfamily
Asiloidea; traditionally part of the Bombyliidae; 7 genera and
about 30 species in Europe.
m m
Characters: Minute to small (0.5-5 ) flies, body bare or with
short sparse hairs, mostly black often with a pattern of yellow
markings; very variable in shape from broad and compact to
slender and elongate. Eyes large, male sometimes holoptic;
antenna with four segments, the third segment largest and 527 Glabellula rctica (Zetterstedt),
with the fourth forming the flagellum, the bristle-like style female; Engel 1932
minute. Mouthparts elongate forming a sucking proboscis,
palps absent or minute. Wing usually clear and without markings; wing venation
reduced, costa incomplete, R2+3 unbranched, discal cell open or closed. Wings held ver
tically above the body when at rest. Legs slender; empodium brisde-like or absent.
Biology: Little is known about the biology, since only six species have been reared world
wide. Larvae of the European Glabellula rctica (Zetterstedt) have been found in the nests
of wood ants (Formica) and are presumed to be predatory. Elsewhere species have been
reared from grasshopper egg-pods and as a parasite of solitary bees. Adults are found rest
ing on flowers on which they feed and flying close by. Most species are found in warm arid
environments.
Identification references: Van der Goot 8t Van Veen 1996 (Northwestern Europe);
Greathead 8t Evenhuis 2001 (key genera); V.R Zaitzev 1989b (former USSR); CMPD:
Greathead 8t Evenhuis 1997 (as Mythicomyiinae); CAT: V F .
Zaitzev i98gd; Evenhuis 2002.

Nannodastiidae (key couplet 167; fig. 363-364)

Systematics: Acalyptrate Brachycera, not yet assigned to a
superfamily; in Europe 1 genus, Azorastia, with 3 species in
the Mediterranean region and the Azores.
Characters: Minute (0.7-1.3 mm), largely dull brown flies. Arista
pubescent or with short rays; ocelli present; Oc-bristles pres
ent; small, weak pseudopostocellar brisdes present, curving
forward or converging; 3 pairs of F-brisdes, curving backward;
vibrissae absent but weak bristles present along the lower mar
gin of the genae. Wing clear; costa with humeral and subcostal
breaks; vein Sc incomplete; crossvein BM-Cu absent;
crossvein DM-Cu present and vein Mi reaching the wing mar
gin; cell cup absent. Tibiae without dorsal preapical bristle. 528 Azorastia mediterranea Papp,
male; Papp 1980
Biology: The biology of this small family is very little known; all
species are found in coastal regions where they often occur in
caves and below overhanging cliffs. The larvae are presumed to
feed on organic detritus.
Identification references: Papp 8t Mathis 2001 (review);
CMPD: Papp & Mathis 1998.

Nemestrinidae (key couplet 49; fig. 107-109)

Systematics: Lower muscomorph Brachycera; superfamily
Nemestrinoidea; 6 genera and about 13 species in Europe.
Characters: Medium sized to very large (9-24 mm) flies devoid of
brisdes, body often covered with relatively long pubescence;
body colour variable, thorax and abdomen sometimes striped or
banded. Ocelli present; eyes dichoptic; mouthparts often elon
gate. Wing clear or tinged, large and long, with complete vena
tion; R-veins curving parallel to the hind margin, in some cases 5 2 t r i n u s reticulars Latreille,
9 N e m e s

with a large number of small cells; cell 1113 closed; cell cup open female; Verrall 1909
 or closed. Legs long and slender, lacking conspicuous bristles; e m p o d i u m pulvilliform.
Biology: T h e larvae are endoparasitoids o f grasshoppers or o f beetles o f the family
Scarabaeidae and play an i m p o r t a n t part i n regulating grasshopper populations. T h e y feed
o n the fats and b o d y juices o f their host. Larvae that are parasitoids o n grasshoppers are
c o n n e c t e d w i t h the air outside t h r o u g h a l o n g respiratory tube. T h e h o s t remains alive
until the larva reaches the fourth instar and e m e r g e s from the h o s t to spend the w i n t e r in
the soil. Pupation takes place i n the n e x t season. A d u l t s often h o v e r i n the air over f l o w
ers or in sunlit, o p e n spots; those w i t h elongate m o u t h p a r t s frequently visit flowers.
Identification references: B e r n a r d i 1973 (genera); P a r a m o n o v 1945-1956 (species);
R i c h t e r 1989a (former U S S R ) ; C M P D : R i c h t e r 1997a; C A T : Richter 1988.

(Nemopalpidae): part o f the Psychodidae.

(Neottiophilidae): part o f the Piophilidae.

Nycteribiidae (key couplet 208;fig.4 4 1 )

Systematics: C a l y p t r a t e B r a c h y c e r a ; s u p e r f a m i l y
H i p p o b o s c o i d e a ; 4 genera and about 15 species in
Europe.
Characters: M i n u t e to m e d i u m sized (1.5-5 m m ) , y e l
l o w i s h to b r o w n , spider-like, flattened flies w i t h o u t
w i n g s . Halteres present. H e a d small, n o t adpressed t o
the thorax b u t , i n resting position, b e n t b a c k w a r d o n 530 Phthiridium biarticulatum Hermann,
to thorax; eyes and ocelli small or absent. Legs l o n g female; Theodor 1975
w i t h s w o l l e n femora and tibiae; first tarsal s e g m e n t at
least as l o n g as all other tarsal s e g m e n t s c o m b i n e d .
Biology: T h e adults are ectoparasites o f bats, feeding o n their b l o o d . T h e larvae develop
inside the a b d o m e n of the female fly. Just prior to deposition the female leaves her bat h o s t
and glues the mature, almost pupated larva (prepupa) to a solid substrate near the resting
place o f the bats, after w h i c h the prepupa fairly rapidly forms a p u p a r i u m . Nycteribiidae
s h o w various degrees o f h o s t specificity, ranging from species-species associations,
t h r o u g h associations w i t h closely related hosts t o apparent absence o f any h o s t prefer
ence.
Identification references: H u t s o n 1984 (British Isles); S c h u u r m a n s S t e k h o v e n Jr. & V a n
d e n B r o e k 1969 (Netherlands); Stackelberg I 9 8 g b e (former U S S R ) ; T h e o d o r 1967 (revi
sion); C M P D : Hurka 1998a; C A T : Hurka & S o o s 1986a.

(Ochthiphilidae): n a m e u s e d in the past for C h a m a e m y i i d a e .

Odiniidae (key c o u p l e t 9 6 , 1 2 1 ; fig. 2 1 4 , 261-262)

Systematics: Acalyptrate Brachycera; superfamily
O p o m y z o i d e a ; 3 genera and about 1 4 species in Europe.
Characters: Small (2-5 m m ) , stout flies, b o d y u s u a l l y grey w i t h
dark m a r k i n g s , legs s o m e t i m e s banded. A r i s t a p u b e s c e n t ;
ocelli present; Oc-bristles present; P-bristles diverging; 3 pairs
o f F-bristles, the l o w e r pair c u r v i n g i n w a r d , the other pairs
b a c k w a r d ; interfrontal bristles absent; vibrissae present. W i n g
usually clear w i t h markings along the crossveins and near the
subcostal break; v e i n Sc incomplete; crossvein B M - C u pres
ent; cell c u p closed. Dorsal preapical bristle o n the tibiae often
thin, hair-like, i n s o m e cases similar to the other hair-like bris 531 Odinia maculata (Melgen), male;
tles. Papp 1978
Biology: T h e larvae usually live in association w i t h w o o d b o r i n g
insects and m a y b e found in tunnels and u n d e r n e a t h the bark of d e c i d u o u s and coniferous
trees. T h e earlier instars feed o n e x u d i n g juices and insect frass, the later instars prey o n
other insects l i v i n g i n w o o d . T h e adults live i n the v i c i n i t y of the larval habitats, in partic
ular near tree w o u n d s or o n w o o d - a s s o c i a t e d m u s h r o o m s and bracket fungi.
Identification references: Stackelberg 1989a} (former U S S R ) ; C M P D : Papp 1998c; C A T :
K r i v o s h e i n a 1984b.

146
Oestridae (key couplet 174; fig. 102,153, 379-383)
Systematics: Calyptrate Brachycera; superfamily Oestroidea;
traditionally this family includes the Gasterophilidae and
H y p o d e r m a t i d a e ; 4 genera and 8 species in Europe.
Characters: M e d i u m sized to large flies (9-18 m m ) . B o d y large
ly w i t h o u t bristles, nearly bare, or b o d y and legs covered in a
soft pilosity, m a k i n g these insects resemble bees or b u m b l e
bees. M o u t h o p e n i n g small, m o u t h p a r t s r u d i m e n t a r y ; anten
na small; ptilinal suture usually w i t h the ends c o n v e r g i n g
b e l o w the antennal insertions. L o w e r calypter large. V e i n M i
strongly c u r v i n g or w i t h an angle in the direction o f the w i n g 532 Cephenemyia trompe (Modeer);
margin or e n d i n g in v e i n R4+5; in the latter case cell r4+5 peti- Grunin 1966
olate. S u b s c u t e l l u m flat or strongly s w o l l e n ; m e r o n i n s o m e
cases w i t h bristles along its hind margin, near the posterior
spiracle (often difficult to see because o f the dense pilosity).
Biology: T h e larvae are parasites in the nostrils or nasal cavities
of m a m m a l s , especially sheep and goats b u t also occasionally antelopes, deer, camels and
horses. T h e host relation is rather specific. Oestridae are larviparous w h i c h m e a n s that
larvae (instead o f eggs) are deposited into the nostrils o f the host, usually from a distance.
T h e female hovers just in front o f the h o s t and ejects the larvae into the host's nostrils.
T h e larvae migrate to the nasal cavities and, in m o s t cases, later into the p h a r y n x . Full-
g r o w n larvae leave their h o s t t h r o u g h the nostrils or m o u t h o p e n i n g and pupate in the
soil. T h e adults h a v e reduced m o u t h p a r t s and do n o t feed. T h e y s e e m t o be rather inactive
insects and are relatively rarely observed. A c o n s p i c u o u s p h e n o m e n o n in this family is
hilltopping, i.e., the habit o f the adults seeking o u t h i g h places or landmarks, i n order to
raise the chances o f males and females m e e t i n g each other.
Identification references: C o l w e l l et al. 2 0 0 4 (general); G r u n i n 1966-1969 (Lindner),
i 9 8 g d (former U S S R ) ; Zeegers 1992 (Netherlands); C M P D : M i n a f 2000c; C A T : S o o s 8t
Minaf 1986b.

(Olbiogastridae): part o f the A n i s o p o d i d a e .

(Omphralidae): n a m e u s e d in the past for Scenopinidae.
(Oncodidae): n a m e used in the past for Acroceridae.

Opetiidae (key couplet 73; fig. 157-158)

Systematics: L o w e r Cyclorrhapha; superfamily Platypezoidea;
traditionally part o f the Platypezidae; in Europe 1 widespread
species, Opetia nigra Meigen.
Characters: A small (2-5 m m ) , slender, all dark species w i t h a
characteristic w i n g venation. Male w i t h holoptic eyes; antenna
w i t h elongate second and third segments, arista l o n g , termi
nal. W i n g darkened along the costa, in particular in the male;
v e i n Sc reaching the w i n g margin; crossvein D M - C u absent;
v e i n M w i t h a l o n g fork; small cell c u p n a r r o w i n g to a point;
anal lobe large in the male, virtually absent in the female. Legs
slender, simple.
Biology: T h e only European species, Opetia nigra, has b e e n
reared from a piece o f birch w o o d i n an advanced state o f
533 Opetia nigra Meigen, male;
decay. T h e ovipositor is peculiarly shaped, suggesting a spe Chandler 1998
cialised m o d e o f o v i p o s i t i n g and a distinctive larval habitat.
T h e adults often occur in w o o d e d areas b u t i n o p e n habitats as
w e l l . A d u l t males are often found o n leaves o f trees and b u s h e s , the m u c h less frequently
found females in l o w e r vegetation. It appears that nectar feeding m a y occur. T h e males
s o m e t i m e s congregate in s w a r m s .
Identification references: C h a n d l e r 2001 (revision); C M P D : C h a n d l e r 1998a; C A T :
Chandler 1991a.

147
Opomyzidae (key couplet 1 0 5 , 1 6 4 , 218; fig. 236-237, 358-360)
Systematics: Acalyptrate Brachycera; superfamily
O p o m y z o i d e a ; 3 genera and about 33 species in Europe.
Characters: Small (2-5 m m ) , slender, y e l l o w , b r o w n , reddish or
black coloured flies. Arista p u b e s c e n t or w i t h relatively l o n g
rays; ocelli present; Oc-bristles present; P-bristles absent or
diverging (in Anomalochaeta); 1 pair o f F-bristles, c u r v i n g
b a c k w a r d ; scattered interfrontal setulae present; vibrissae
absent although Geomyza w i t h a strong bristle near t h e v i b -
rissal angle. W i n g nearly a l w a y s w i t h an apical spot a n d / o r
darkly marked crossveins or marked all over; costa w i t h a sub
costal break; v e i n Sc usually i n c o m p l e t e b u t apical part s o m e
t i m e s visible as a thin line reaching the costa; crossvein B M - C u
present b u t usually i n c o m p l e t e ; cell c u p closed. Tibiae w i t h o u t
534 Opomyza florum (Linnaeus), male;
dorsal preapical bristle. Balachowsky & Mesnil 1935
Biology: In v i e w o f the larval food plants, i.e., grasses, the adults
are m a i n l y found in o p e n habitats. T h e larvae live in the stems o f grasses, a f e w species
b e i n g a pest in agriculture.
Identification references: C z e r n y 1928 (Lindner); Drake 1992, 1993 (British Isles); Soos
1981 (Central E u r o p e ) ; Stackelberg i 9 8 g a p (former U S S R ) ; Carles-Tolra 1993, 1 9 9 4
(Spain); V a n Zuijlen 1999 (Fallen collection); C M P D : Brunei 1998; C A T : Soos 1984k.

(Ortalidae): n a m e u s e d in t h e past for Ulidiidae: Otitinae.

(Otitidae): part o f the Ulidiidae.

Pachyneuridae (key couplet 29;fig.71)

Systematics: N e m a t o c e r a ; s u p e r f a m i l y P a c h y n e u r o i d e a ; i n
N o r t h e r n Europe 1 species, Pachyneurafasciata Zetterstedt.
Characters: Pachyneurafasciata is a large species (10-13 m m ) ,
r e s e m b l i n g Tipulidae o n account of its l o n g and slender anten
nae, w i n g s , legs and a b d o m e n . Ocelli p r o m i n e n t ; antenna
nearly as l o n g as the thorax, w i t h 17 segments. W i n g partly
tinged and w i t h a distinct stigma spot. Tibiae w i t h apical bris
tles.
Biology: Larvae a n d p u p a e o f t h e o n l y E u r o p e a n s p e c i e s ,
Pachynema fasciata, are found in rotting w o o d l y i n g o n the
ground, in particular in parts w i t h black f u n g o u s tissue. A d u l t s
o c c u r i n early s u m m e r o n tree trunks, p r e d o m i n a n t l y o n birch.
Identification references: Stackelberg i g 8 9 f (former U S S R ) ; 535 Head Pachyneura fasciata
C M P D : K r i v o s h e i n a 1997a; C A T : K r i v o s h e i n a & M a m a e v Zetterstedt; Krivosheina 1997a
1988a.

Pallopteridae (key couplet 1 4 4 , 1 6 2 ; fig. 311-313,352-353)

Systematics: A c a l y p t r a t e Brachycera; superfamily
Tephritoidea; 5 genera and about 23 species in Europe.
Characters: Small to m e d i u m sized (2.5-7 m m ) , slender flies
v a r y i n g in colour from pale y e l l o w to reddish, grey or black;
s o m e parts o f the head often silvery grey to w h i t e . Arista bare
or w i t h short rays, rays at m o s t t w i c e as l o n g as basal diameter
of arista; ocelli present; Oc-bristles present; P-bristles parallel
to s o m e w h a t diverging; 1 pair o f strong F-bristles, c u r v i n g
b a c k w a r d , s o m e t i m e s w i t h a w e a k e r F-bristle in front; scat
tered interfrontal setulae present, s o m e t i m e s almost invisible
or indeed lacking; vibrissae absent b u t usually a series o f fine
setulae present along t h e l o w e r m a r g i n o f the genae. W i n g 536 Palloptera quinquemaculata
often w i t h a pattern, s e l d o m clear; costa w e a k just b e y o n d (Macquart), female; Balachowsky &
crossvein H and w i t h a subcostal break; v e i n Sc complete; Mesnil 1935

148
 crossvein B M - C u present; cell c u p closed. Tibiae w i t h o u t dorsal preapical bristle, except
in Eurygnathomyia.
Biology: In s o m e species the larvae live under tree bark w h e r e they feed o n m y c e l i u m or
prey o n beetles (Carabidae, C e r a m b y c i d a e and Scolytidae). Larvae o f other species live in
plants, such as grasses, Apiaceae or the flower heads (inflorescences) o f Asteraceae
(Compositae); it is u n k n o w n w h e t h e r the species living in plants are really p h y t o p h a g o u s ,
carnivorous, or b o t h . T h e adults are generally found in shaded habitats, often near water.
D e p e n d i n g on the larval habitat, the adults are found o n flowers, often those o f umbellif-
ers, or on tree trunks and leaves.
Identification references: A n d e r s s o n 1 9 9 0 ( S w e d e n ) ; G r e v e 1993 ( N o r w a y ) ; J.F.
M c A l p i n e 1981c ( W o r l d genera); M o r g e 1963-1974 (review); Stackelberg 1989a]! (former
U S S R ) ; C M P D : M e r z 1998b; C A T : M o r g e 1984.

Pediciidae (key couplet 5 , 1 9 6 , 1 9 8 ; fig. 30,428-429,432)

Systematics: N e m a t o c e r a ; s u p e r f a m i l y T i p u l o i d e a ; u n t i l
recently part o f the Limoniidae w h i c h is traditionally included
in the Tipulidae; 5 genera and about 6 0 species in Europe.
Characters: Nematocera v a r y i n g from m e d i u m sized (5 m m in
Dicranota, subgenus Paradicranotd) to v e r y large (35 m m in
Pedicid). A n t e n n a e , w i n g s , legs and a b d o m e n l o n g and slender.
Ocelli absent; e y e s pubescent, w i t h o m m a t r i c h i a in b e t w e e n
the facets; antenna w i t h 12-17 s e g m e n t s . T h o r a x w i t h a V -
shaped transverse suture. W i n g w i t h 2 anal v e i n s , w i n g in
s o m e cases w i t h a characteristic pattern.
Biology: T h e larvae o f the g e n u s Ula feed o n m u s h r o o m s and
bracket fungi; all other larvae, as far as k n o w n , are aquatic to
semi-aquatic and prey o n w o r m s , insect larvae, mites and
other small invertebrates. Predatory larvae are m o b i l e and rap
idly m o v i n g ; because o f their massive head capsule they are
n o t able to s w a l l o w large prey as they are but h a v e to tear t h e m 537 Tricyphona immaculata (Meigen),
m a l e ; Pa &
to pieces. O f the three m o r e c o m m o n genera, the larvae o f P P Schumann 2000
Tricyphona are found in diverse w e t and semi-aquatic habitats
(from the shores of springs, streams and lakes to b o g s and forest litter in w e t places), those
of Dicranota tend to occur along and in r u n n i n g w a t e r s b u t d e v e l o p also at the shores o f
standing water b o d i e s ; larvae o f Pedicia are restricted to r u n n i n g w a t e r o f w e l l s , b r o o k s
and small rivers. Pupating takes place inside a silken tube. A d u l t s are found in an array o f
m o i s t habitats.
Identification references: D i e n s k e 1987; R e u s c h 8t Oosterbroek 1997 (genera and s u b g e n
era); S a v c h e n k o 1989c, i 9 8 g d (former U S S R ) ; Stubbs 1 9 9 4 - 1 9 9 9 (British Isles); C M P D : no
contribution; C A T : S a v c h e n k o et al. 1992; Oosterbroek 2006.

Periscelididae (key couplet 121; fig. 259-260)

Systematics: Acalyptrate Brachycera; superfamily
O p o m y z o i d e a ; in Europe 1 g e n u s , Periscelis, w i t h 4 species.
Characters: M i n u t e to small (1-5 m m ) , grey or dark coloured
flies, often w i t h banded legs. A n t e n n a w i t h second s e g m e n t
overhanging third segment; arista w i t h l o n g rays; ocelli pres
ent; Oc-bristles present; P-bristles diverging; 1 pair o f F-bris-
tles, c u r v i n g b a c k w a r d ; interfrontal bristles absent; a r o w o f
vibrissa-like bristles present. W i n g clear or w i t h infuscated
spots; costa continuous; v e i n Sc incomplete; crossvein B M - C u
present; cell c u p o p e n or closed. Tibiae w i t h o u t dorsal preapi
cal bristle.
Biology: T h e larvae o f this small family live in sap e x u d i n g
from d e c i d u o u s trees. T h e adults are often found near the lar
val sites and feed o n tree sap. Males m a y be seen h o v e r i n g
near sap runs.

149
Identification references: Bachli 1997 (Switzerland); Stackelberg ig89ae (former U S S R ) ;
C M P D : Mathis & Papp 1998; C A T : Papp 1984b.

(Petauristidae): n a m e u s e d in the past for Trichoceridae.

Phaeomyiidae (key couplet 143; fig. 310)

Systematics: Acalyptrate Brachycera; superfamily
S c i o m y z o i d e a , traditionally part o f the S c i o m y z i d a e ; in
Europe 1 g e n u s , Pelidnoptera, w i t h 3 species.
Characters: Small to large (3-11 m m ) , y e l l o w i s h b r o w n to b r o w n
coloured, slender flies w i t h b r o w n i s h tinged w i n g s . Arista
p u b e s c e n t ; ocelli p r e s e n t ; O c - b r i s t l e s p r e s e n t ; P-bristles
diverging; 2 pairs o f F-bristles, c u r v i n g b a c k w a r d ; interfrontal
bristles absent; vibrissae absent. W i n g tinged b r o w n , w i t h o u t
markings; costa continuous; v e i n Sc complete, crossvein B M - 539 Pelidnoptera nigripennis (Fabricius),
C u present; cell c u p closed. Tibiae w i t h dorsal preapical bristle. male; Rivosecchi 1992
Biology: O n e species i n this small family is k n o w n to b e a para-
sitoid o f D i p l o p o d a (millipedes) of the g e n u s Ommatoiulus. T h e female deposits her eggs
o n the millipede after w h i c h the y o u n g larva enters its host. U s u a l l y only o n e larva per
millipede w i l l emerge. T h e host remains alive until the larva is fully g r o w n and pupates in
the h o s t w h i c h , b y that time, is n o m o r e than an e m p t y shell.
Identification references: R e v i e r 8t V a n der G o o t 1989 ( N o r t h w e s t e r n E u r o p e ) ;
R i v o s e c c h i 1992 (Italy); Stackelberg 19893c (former U S S R ) ; C M P D : R o z k o s n y 1998b;
C A T : R o z k o s n y 8t Elberg 1984 (as subfamily o f S c i o m y z i d a e ) .

(Phlebotomidae): part o f the Psychodidae.

Phoridae (key couplet 6 9 , 210; fig. 1 4 8 - 1 4 9 , 4 4 5 )

Systematics: L o w e r Cyclorrhapha; superfamily Platypezoidea;
s o m e 35 genera and about 605 species in Europe.
Characters: M i n u t e to m e d i u m sized (0.5-6 m m ) flies, often
s o m e w h a t arched and m o r e or less stout, w i t h a characteristic
w i n g venation. Head, palps and legs usually w i t h strong, d e n
tate or feathered setae. T h e b o d y colour varies from black
t h r o u g h b r o w n , orange and y e l l o w to pale grey or pale w h i t e .
Eyes dichoptic; antenna w i t h a large, r o u n d e d or elongate third
s e g m e n t bearing a l o n g apical or s o m e w h a t dorsal arista direct
ed s i d e w a y s . W i n g clear or tinged, rarely w i t h markings; v e n a
tion reduced, the strong R - v e i n s e n d i n g in the costa about
halfway u p the w i n g ; the other v e i n s are w e a k e r and usually
f o l l o w a diagonal course, often parallel to each other. Legs t y p
ically w i t h stout femora; h i n d femora often s o m e w h a t lateral
540 Megaselia scalarls (Loew), female;
ly compressed. Disney 1983
Biology: Rather variable, s h o w i n g the greatest diversity o f all
D i p t e r o u s families. Larvae are found in several terrestrial habitats, in the nests o f social
insects, and in s o m e aquatic habitats, feeding o n organic detritus such as d u n g , carrion,
insect frass, dead snails, etc.; s o m e feed o n bracket fungi and m u s h r o o m s , o n m y c e l i u m
or o n living plants (among others as miners), t h e y m a y be predators or parasites of a w i d e
array o f organisms such as e a r t h w o r m s , snails, spiders and their eggs, centipedes, milli
p e d e s , insect larvae and pupae. M o s t adults feed o n nectar, h o n e y d e w and juice e x u d i n g
from fresh carrion and d u n g ; s o m e feed o n b o d y juices o f living beetle larvae and pupae,
others prey o n small insects. T h e adults are c o n s p i c u o u s o n account o f their fast and
s o m e w h a t abrupt m a n n e r o f l o c o m o t i o n . In s o m e species, the males fly in s w a r m s .
Identification references: D i s n e y 1983-1989 (British Isles); D i s n e y 1 9 9 4 (genera); V F .
Z a i t z e v 1989c (former U S S R ) ; C M P D : D i s n e y 1998; C A T : D i s n e y 1991.

(Phryneidae): n a m e used in the past for A n i s o p o d i d a e .

(Phthiriidae): part o f the B o m b y l i i d a e .

150
PiophiUdae (key couplet 114; fig. 247-248)
Systematics; Acalyptrate flies; superfamily Tephritoidea; s o m e
16 genera and about 30 species in Europe.
Characters: M i n u t e to m e d i u m sized (1.5-7 m m ) , y e l l o w or
b r o w n to black, in s o m e cases lustrous flies, s o m e species w i t h
a striking pale coloration of the face or w i t h m o r e or less e x t e n
sive y e l l o w or reddish coloured parts. A r i s t a bare to p u b e s
cent; ocelli present; Oc-bristles present; P-bristles d i v e r g i n g ,
rarely parallel; 0-4 pairs o f F-bristles, i f present c u r v i n g out
w a r d or b a c k w a r d , in Actenoptera the l o w e r 2 pairs c u r v i n g
inward; scattered interfrontal setulae present; vibrissae pres
ent. W i n g usually u n m a r k e d , in s o m e genera w i t h spots along
the crossveins a n d / o r at the ends o f the longitudinal v e i n s ;
costa w i t h a subcostal break; v e i n Sc c o m p l e t e ; crossvein B M -
541 Piophila casei Linnaeus, male;
C u virtually a l w a y s present; cell c u p closed. Tibiae w i t h o u t
Stackelberg 1956
dorsal preapical bristle.
Biology: T h e larvae o f m o s t species live in rotting w o o d , bracket fungi, or m u s h r o o m s ,
s o m e species live in nests o f birds w h e r e they feed o n the b l o o d o f the nestlings
(Neottiophilum), or in carrion, m a i n l y that o f larger vertebrates (Thyreophora and its
allies), or in substrates rich in protein. A w e l l - k n o w n representative o f the latter group is
the larva o f Piophila casei (Linnaeus), a pest in a large v a r i e t y o f animal products such as
cheese, dried fish, meat, etc. T h e adults are n o t v e r y active fliers and are m a i n l y found near
the larval habitats for oviposition or o n tree trunks and logs, possibly for mating.
Identification references: J.F. M c A l p i n e 1977 (genera); M e r z 1 9 9 6 b ( S w i t z e r l a n d ) ;
G o r o d k o v 1989a; Stackelberg i g S g a f , i 9 8 g a g (former U S S R ) ; C M P D : O z e r o v 2000; C A T :
Zuska 1984; Papp 1984c (Thyreophorinae); Soos 1 9 8 4 b (Neottiophilinae).

PipiMCUlidae (key couplet 65; fig. 139-140)

Systematics: L o w e r C y c l o r r h a p h a ; s u p e r f a m i l y
Syrphoidea; s o m e 14 genera and about 200 species in
Europe.
Characters: Small to large (2-12 m m ) , dark coloured
flies w i t h long, n a r r o w w i n g s and v e r y large e y e s
o c c u p y i n g nearly all o f the semi-spherical to nearly
entirely spherical head. A n t e n n a short, third seg
m e n t usually the largest, w i t h a dorsal arista or style;
m o u t h p a r t s small. W i n g clear or tinged b r o w n ; v e i n
M i and cell d m s o m e t i m e s absent; cell c u p elongate, 542 Pipunculus campestris Latreille,
male; Verrall 1909
e n d i n g acutely just before the w i n g margin. T h e
female has a lance-like ovipositor.
Biology: T h e larvae are p r e d o m i n a n t l y parasitoids o f leafhoppers and p l a n t h o p p e r s
(Hemiptera: A u c h e n o r r h y n c h a ) except Nephrocerus w h i c h has b e e n reared from adult
Tipulidae (Koenig 8t Y o u n g 2007). T h e large eyes of the adult p i p u n c u l i d s apparently play
an i m p o r t a n t role in locating a suitable host, as does the ability to h o v e r in o n e spot.
O n c e located, the female uses her lance-like o v i p o s i t o r to deposit an e g g directly inside
the host. More than o n e larva m a y be present in a single host. W h e n nearing c o m p l e t i o n
of d e v e l o p m e n t , the larvae eat the h o s t until o n l y an e m p t y shell remains. T h e larvae then
leave t h r o u g h the cuticle to pupariate in the soil.
Identification references: A l b r e c h t 1990 (Dorylomorpha); C o e 1966 (British Isles); V o n
der D u n k 1998 (Central Europe); Foldvari & D e M e y e r 2000 (Tomosvaryella); Jervis 1992
(Chalarus); Kehlmaier 2005 (tribe Eudorylini); D e M e y e r 1989 (Cephalops, Beckerias);
Tanasijtshuk 1989c (former U S S R ) ; C M P D : K o z a n e k et al. 1998; C A T : Tanasijtshuk 1988;
De M e y e r 1996.
Platypezidae (key couplet 74; fig. 159-160)
Systematics: L o w e r C y c l o r r h a p h a ; s u p e r f a m i l y
P l a t y p e z o i d e a ; traditionally this family includes the
O p e t i i d a e ; s o m e 12 genera and about 4 5 species i n
Europe.
Characters: M i n u t e to m e d i u m sized (1.5-6 m m ) ,
slender to stout flies, male and female often differ
ing in colour, the male in particular often all black;
one or b o t h sexes partially to all grey or y e l l o w , i n
s o m e cases w i t h orange, grey or silvery markings, or
a b d o m e n b i c o l o u r o u s . Eyes holoptic in the male;
antenna w i t h 3 s e g m e n t s , the third o f w h i c h is
largest and bears a l o n g , apical arista. W i n g clear or 543 Protoclythia modesta (Zetterstedt),
male; Verrall 1909
tinged, in s o m e genera the area along the w i n g mar
gin, in b e t w e e n and around v e i n s Sc and R i dark
ened; w i n g w i t h c o n s p i c u o u s anal lobe; anal v e i n reaching the w i n g margin; cell c u p end
ing in an acute angle, often elongate; in m o s t genera v e i n M forked; crossvein D M - C u vir
tually a l w a y s present (absent in Microsomia w h e r e crossvein R - M is also lacking). Legs rel
atively short, stout; first tarsal s e g m e n t of h i n d leg in the subfamily Callomyiinae l o n g and
cylindrical, usually s w o l l e n in the male, slender in the female, the subsequent tarsal seg
m e n t s cylindrical; in the subfamily Platypezinae the first tarsal s e g m e n t o f h i n d leg short,
laterally compressed, s u b s e q u e n t tarsal s e g m e n t s laterally compressed as w e l l , in particu
lar i n the female.
Biology: A s far as k n o w n the larvae d e v e l o p o n and in m u s h r o o m s and bracket fungi. T h e
adults are m a i n l y found in w o o d l a n d areas, w a l k i n g about o n leaves o f trees and shrubs
w h e r e t h e y feed o n h o n e y d e w and other organic matter o n the leaf surface. T h e males
s o m e t i m e s fly in s w a r m s ; those o f Microsania are k n o w n as ' s m o k e flies' since they
s w a r m in the s m o k e o f forest fires (and other fires as well). T h e larval m o d e o f life o f
Microsania is not k n o w n .
Identification references: Chandler 2001 (revision); Tanasijtshuk 1989b (former U S S R ) ;
C M P D : Chandler & Shatalkin 1998; C A T : C h a n d l e r 1991b.

Platystomatidae (key couplet 112,163; fig. 306,354-355)

Systematics: Acalyptrate Brachycera; superfamily
Tephritoidea; 2 genera, Platystoma and Rivellia, and about
20 species in Europe.
Characters: S m a l l t o large (3-11 m m ) , dark flies, i n
Platystoma w i t h b r o w n w i n g s w i t h n u m e r o u s hyaline
dots and stout, microtrichose b o d y and usually bright y e l
l o w ventral part o f a b d o m e n , in Rivellia w i t h dark banded
or spotted w i n g s and shiny, n a r r o w body. Head round to
distinctly flattened; arista bare to pubescent; ocelli pres
ent; Oc-bristles reduced or absent; P-bristles reduced or
absent; 1-2 pairs o f F-bristles; scattered interfrontal setulae 544 Platystoma seminationis (Fabricius),
male; Colyer & Hammond 1968
present; vibrissae absent b u t s o m e t i m e s vibrissa-like bris
tles present. C o s t a w i t h a h u m e r a l break, o t h e r w i s e c o n
tinuous; v e i n Sc complete, crossvein B M - C u present; cell
c u p closed. Tibiae w i t h o u t dorsal preapical bristle. Female w i t h oviscape, n o n retractable
basal s e g m e n t o f the ovipositor.
Biology: T h e larval habitats are rather diverse, m o s t species feeding o n rotting plant materi
als or o n roots in the soil, s o m e species living under the bark o f deciduous trees; larvae of
Rivellia feed o n the root nodules in legumes. T h e adults are also found in various habitats
such as w o o d l a n d , m o i s t m e a d o w s , dunes, etc., often resting o n tree trunks, stones and
leaves, in the latter case preferring the l o w e r side. T h e adults feed o n the juice e x u d i n g from
d u n g and carrion, o n nectar, h o n e y d e w , fruit and o n sap e x u d i n g from tree w o u n d s .
Identification references: C l e m e n t s 1 9 9 0 (British Isles); K o r n e y e v 2001 (genera); M e r z
1996c (Switzerland); Richter 1989c (former U S S R ) ; C M P D : D . K . M c A l p i n e 1998a; C A T :
Soos 19846.
Pleciidae (key couplet 27, 205; fig. 67-68)
Systematics: Nematocera; superfamily Bibionoidea, tradition
ally part o f t h e Bibionidae; in Europe 2 species, Penthetria
funebris M e i g e n (native, widespread) and P. heteroptera (Say)
(probably introduced, G e r m a n y ) .
Characters: T h e only native European species, Penthetria fune
bris, is m e d i u m sized (6-8 m m ) , B i i i o - l i k e , dark to black
coloured, w i t h short antennae, an elongate a b d o m e n and short
and n a r r o w w i n g s , especially so in the male. Ocelli p r o m i n e n t ;
eyes in the male almost t o u c h i n g and d i v i d e d into t w o parts;
in the female the e y e s are simple and dichoptic; antenna
inserted b e l o w the m i d d l e of the face, short, w i t h 11 segments.
W i n g tinged; v e i n R forked into R2+3 and R4+5; v e i n M forked
into M i and M2; anal lobe especially present in t h e female. Legs
545 Penthretia funebris (Meigen),
simple; tibiae w i t h distinct b u t relatively short apical spurs.
male; Seguy 1940
Biology: T h e larvae feed o n d e c a y i n g organic matter in the t o p -
soil o f d e c i d u o u s w o o d l a n d s , often near little streams or near tree trunks; t h e larvae o f P.
funebris h a v e b e e n found d e v e l o p i n g i n alder litter, the Nearctic P. heteroptera w a s found
at a c o m p o s t i n g facility in G e r m a n y w h e r e t h e larvae apparently d e v e l o p e d i n vegetable
refuse (Fitzgerald & W e r n e r 2004). T h e adults are usually rare b u t can occur in large n u m
bers locally; they d o n o t fly b u t w a l k o n the g r o u n d or crawl t h r o u g h t h e vegetation near
the larval habitats.
Identification references: D u d a 1930 (Lindner); Fitzgerald 8t W e r n e r 2 0 0 4 (key, r e v i e w ) ;
Mikolajczyk 1977 (Poland); Zeegers 1997, 1998 (Netherlands); C M P D : Skartveit 1997;
C A T : K r i v o s h e i n a 1986a.

Pseudopomyzidae (key couplet 133; fig. 294-296)

Systematics: Acalyptrate Brachycera; superfamily Nerioidea;
in Europe 1 w i d e s p r e a d species, Pseudopomyza atrimana
(Meigen).
Characters: A m i n u t e to small (1.7-2.5 m m ) , dark to black
coloured species. Arista pubescent; ocelli present; Oc-bristles
present; P-bristles converging; 3 pairs o f F-bristles, c u r v i n g
b a c k w a r d ; interfrontal bristles absent; vibrissae present.
W i n g u n m a r k e d , costa w i t h b o t h subcostal and h u m e r a l
breaks or a subcostal break and a humeral w e a k spot; v e i n Sc
incomplete; crossvein B M - C u absent; cell c u p closed. Tibiae
w i t h o u t dorsal preapical bristle.
Biology: T h e b i o l o g y o f this small family is little k n o w n . T h e
only European species, Pseudopomyza atrimana, occurs in 546 Head Pseudopomyza atrimana
w o o d l a n d habitats, and adults h a v e b e e n recorded to gather (Meigen); Chandler 1983
over rotting logs, a habitat similar to that o f an East A s i a n
species w h o s e larvae w e r e found l i v i n g under bark o f v a r i o u s
trees.
Identification references: C h a n d l e r 1983 ( B r i t i s h Isles);
Stackelberg i g 8 9 a n (former U S S R ) ; C M P D : D . K . M c A l p i n e &
Shatalkin 1998; C A T : K r i v o s h e i n a 1984c.

Psilidae (key couplet 150; fig. 325-327)

Systematics: Acalyptrate Brachycera; superfamily D i o p s o i d e a ;
6 genera and about 50 species in Europe.
Characters: Small to m e d i u m sized (2.5-10 m m ) , slender to
stout, y e l l o w to reddish, b r o w n or black coloured flies w i t h
few bristles. Head w i t h relatively small eyes, m o s t often w i t h
receding face; antenna small or w i t h third antennal s e g m e n t
c o n s p i c u o u s l y elongate; arista short or longer pubescent; 547 Psila merdaria (Collin), male; van der
ocelli present; Oc-bristles present; P-bristles diverging or Goot & van Veen 1996

153
 absent; 0-2 pairs o f F-bristles; scattered interfrontal setulae present; vibrissae absent.
W i n g u s u a l l y clear to slightly tinged, rarely marked, in s o m e cases along costa, the w i n g
tip or the crossveins w i t h dark band(s); w i n g w i t h a transverse w e a k e n i n g in basal half;
costa w i t h a subcostal break at s o m e distance from w h e r e v e i n R i m e e t s the costa; v e i n Sc
incomplete, its apical part transparent and m o r e or less perpendicular to the basal part;
crossvein B M - C u present; cell c u p closed. Tibiae w i t h o u t dorsal preapical bristle.
Biology: T h e larvae are nearly all p h y t o p h a g o u s . T h e y live in roots, tubers and stems o f a
range o f n o n - w o o d y herbs, a f e w species live under tree bark. Several species are pests in
agriculture and horticulture (such as the root fly Psila rosae Fabricius, living in several
crops nearly all over the w o r l d ; and Psila nigricornis M e i g e n w h i c h inhabits the roots o f
c h r y s a n t h e m u m s and other garden plants). T h e adults prefer a l o w , dense vegetation.
Identification references: V a n der G o o t & V a n V e e n 1996 ( N o r t h w e s t e r n Europe);
Stackelberg 1989W (former U S S R ) ; C M P D : Iwasa 1998b; C A T : S o o s 1984c.

Psychodidae (key couplet 1 2 ; fig. 4 2 - 4 2 3 )

Systematics: N e m a t o c e r a ; superfamily Psychodoidea; s o m e 4 0
genera and about 500 species in Europe.
Characters: Small to m e d i u m sized (2-6 m m ) , often stout, u s u
ally densely pubescent Nematocera, non-piercing except for
the subfamilies Sycoracinae and Phlebotominae. Ocelli absent;
in the Psychodinae, the largest and m o s t c o m m o n subfamily,
the eyes are reniform and w i t h an eye bridge, eyes circular in
the other subfamilies; antenna usually w i t h 12 to 16 segments,
the flagellomeres bottle- or barrel-shaped. W i n g hairy, w i t h 9-
10 longitudinal v e i n s and the R-fork and M-fork a l w a y s pres
ent, t h o u g h they m a y be incomplete basally; crossveins situat-
ed near the w i n g base or (partly) absent; w i n g usually acute, in 5 4 8 P n l e b o t o m u s p a p a t a s i ( S c o p o l i )

T r i c h o m y i i n a e , S y c o r a c i n a e and s o m e P s y c h o d i n a e w i n g male; Seguy 1950

rounded. W h i l e resting or w a l k i n g the w i n g s are hold above the
a b d o m e n like a roof, or besides the a b d o m e n in a broad V - s h a p e . Legs ranging from short
to elongate.
Biology: T h e larval habitats o f the five European subfamilies are rather diverse (and in s o m e
cases insufficiently k n o w n ) b u t truly aquatic species are lacking. T h e larvae o f the
P h l e b o t o m i n a e are k n o w n f r o m soil d u g u p f r o m r o d e n t b u r r o w s . T h o s e o f the
Sycoracinae are semi-aquatic in m o s s e s , spring b r o o k s , or waterfalls and fast streams
w h e r e t h e y l i v e o n r o c k s near the surface or in the s p l a s h z o n e . T h o s e o f the
T r i c h o m y i i n a e and probably the B r u c h o m y i i n a e live in rotting and dead w o o d o f decidu
ous trees. T h e larvae o f the Psychodinae occur in a w i d e range o f habitats: semi-aquatic,
water-filled tree holes, leaf axils, kitchen sink deposit catchers, w a t e r purification plants
(where t h e y can be a pest), or m o r e terrestrially in h u m u s and leaf litter, as miners in
leaves, in excrement, in public lavatories and toilets, in sand and m u d alongside water, in
m o s s , m u s h r o o m s and bracket fungi, in dead w o o d , etc. T h e adults have a short and errat
ic flight and are largely found in w o o d l a n d areas near water, in marshland, or near (indoor)
larval habitats. T h e y are p r e d o m i n a n t l y n o c t u r n a l and b y day they rest in dark places.
Certain species are regularly found o n w i n d o w panes at d u s k or occasionally s w a r m
r o u n d s i n k o v e r f l o w s . M o u t h p a r t s are n o n - f u n c t i o n a l e x c e p t in f e m a l e s o f the
Sycoracinae, w h i c h feed o n the b l o o d o f amphibians, and the Phlebotominae, the 'sand
flies' w h i c h feed o n h u m a n b l o o d and, in the tropics, can be i m p o r t a n t disease vectors.
S o m e Psychodinae do occasionally b e c o m e a nuisance if they, for example, breed in n u m
bers in domestic drains.
Identification references: Jung 1958 (Lindner: B r u c h o m y i i n a e ) ; Lane 1993
( P h l e b o t o m i n a e ) ; P e r f i l y e v 1989 ( f o r m e r U S S R ) ; S z a b o 1983 ( C e n t r a l E u r o p e ) ;
Tanasijtshuk 1989a (former U S S R ) ; T h e o d o r 1958 (Lindner: Phlebotominae); Vaillant
1971-1983 (Lindner: Psychodinae); W a g n e r 1982 (Trichomyiinae), i 9 9 7 d (subfamilies,
genera Psychodinae); W i t h e r s 1989 (British Isles); C M P D : W a g n e r 1997a; C A T : W a g n e r
1990a.
Ptychopteridae (key couplet n ; fig. 4 0 - 4 1 )
Systematics: N e m a t o c e r a ; s u p e r f a m i l y P t y c h o p t e r o i d e a ; i n
Europe 1 genus, Ptychoptera, w i t h about 15 species.
Characters: M e d i u m sized to large (7-15 m m ) , slender, black l u s
trous N e m a t o c e r a , often w i t h lighter m a r k i n g s o n thorax
a n d / o r a b d o m e n . A n t e n n a e , w i n g s , a b d o m e n and legs l o n g
and slender. Ocelli absent; antenna w i t h 15 to 2 1 segments.
T h o r a x w i t h deep, posteriorly directed transverse suture.
W i n g w i t h markings, in particular along the crossveins and
w h e r e v e i n s bifurcate; spurious v e i n present o n either side o f
c r o s s v e i n R - M a n d w i n g m e m b r a n e w i t h a d i s t i n c t fold
b e t w e e n v e i n s A i and Q 1 A 2 .
Biology: T h e larvae are aquatic to semi-aquatic, living in m u d d y
water, s h a l l o w p o o l s in marshes, Sphagnum p o o l s , or the mar
gins o f streams. T h e y filter-feed o n small organic particles. 549 Ptychoptera contaminata (Linnaeus);
T h e y are connected w i t h the air t h r o u g h a l o n g ventilation male; Dely-Draskovits 1983
tube at the e n d o f the a b d o m e n . T h e adults live in m a r s h y and
m o i s t habitats, near suitable substrates for t h e larvae as w e l l as quite s o m e distance from
the larval habitat.
Identification references: A n d e r s s o n 1997a (species); K r z e m i n s k i 1986 (Poland); Peus
1958 (Lindner); S t a c k e l b e r g 1989b (former U S S R ) ; D e l y - D r a s k o v i t s 1983 (Central
Europe); C M P D : R o z k o s n y 1997a; C A T : R o z k o s n y 1992.

PyrgOtidae (key couplet 148; fig. 323-324)

Systematics: Acalyptrate Brachycera; superfamily
Tephritoidea; in Central Europe 1 species, Adapsilia coarctata
Waga.
Characters: T h e o n l y European species, Adapsilia coarctata, is
m e d i u m sized (8-9 m m ) , b r i g h t l y to r e d d i s h y e l l o w , t h e
female w i t h a c o n s p i c u o u s l y l o n g ovipositor. Arista p u b e s
cent, ocelli absent, Oc-bristles small or absent, P-bristles small
and diverging or absent; 1-2 pairs o f small F-bristles; scattered
interfrontal setulae present; vibrissae absent. W i n g venation
as in fig. 324, the intensity and size o f the markings variable;
costa continuous, w i t h a w e a k spot near w h e r e v e i n Sc ends in
it; v e i n Sc nearly complete, the apical part near the costa b e i n g
pallid; crossvein B M - C u present; cell c u p closed. Tibiae w i t h
550 Adasilia coarctata Waga, male;
o u t dorsal preapical bristle. Seguy 1934
Biology: A s far as k n o w n , t h e larvae o f Pyrgotidae are endopar-
asitoids o f beetles o f the superfamily Scarabaeoidea. Similar to
w h a t is found in t h e C o n o p i d a e , an e g g is deposited inside the
a b d o m e n of the h o s t during flight. T h e adults appear m a i n l y to
be nocturnal, b u t they are found b y day as w e l l , for example
w h e n they visit flowers.
Identification references: Enderlein 1 9 4 2 (genera); H e n d e l
1933 (Lindner); K o r n e y e v 2 0 0 4 (genera); M e r z 1 9 9 6 c
(Switzerland); Stackelberg 1 9 8 9 X (former U S S R ) ; C M P D : n o
contribution; C A T : Soos i g 8 4 d .

Rachiceridae (key couplet 39;fig.85)

Systematics: L o w e r Brachycera; superfamily X y l o p h a g o i d e a ; in
S o u t h w e s t e r n Europe (France, Spain) 1 species, Rachicerus
tristis (Loew).
Characters: Rachicerus tristis, t h e o n l y European species, is
m e d i u m sized (6.5-8.5 m m ) , slender and dark coloured; legs
y e l l o w w i t h at least tarsal segments darkened; pubescence y e l - 551 Rachicerus varius Nagatomi,
l o w o n thorax, black o n a b d o m e n . Eyes dichoptic; antenna male; Nagatomi 1970 (Japan)

155
 extraordinary, w i t h 34-35 segments. W i n g w i t h a grey tinge; b o t h the cell m 3 and cell c u p
closed. Legs slender; tibiae w i t h short apical spurs; e m p o d i u m pulvilliform.
Biology: T h e b i o l o g y o f this small family is virtually u n k n o w n . T h e larvae live in rotting
w o o d , probably as predators.
Identification references: C M P D : N a g a t o m i 1997a; C A T : Majer 1988c.

Rhagionidae (key couplet 55; fig. 9 4 , 1 1 8 - 1 1 9 )

Systematics: L o w e r Brachycera; superfamily Tabanoidea; tra
ditionally this family includes the Athericidae and
Hilarimorphidae; 6 genera and about 85 species in Europe.
Characters: Small to large (2-20 m m ) flies w i t h l o n g w i n g s ,
legs and a b d o m e n . B o d y bare or partially pubescent, rarely
w i t h a f e w bristles. B o d y colour y e l l o w , y e l l o w i s h b r o w n or
black, often w i t h a b r o w n i s h , grey or dark pattern. In male
eyes generally nearly or entirely holoptic; third antennal seg
m e n t s w o l l e n and w i t h a dorsal or terminal style or arista;
m o u t h p a r t s usually fleshy. W i n g usually clear and w i t h a
dark stigma spot, in s o m e cases w i t h a distinct pattern; w i n g
v e n a t i o n usually w i t h 3 v e i n s from the discal cell in the 552 Rhagio scolopaceus (Linnaeus),
direction o f the w i n g margin; cell IT13 o p e n ; cell c u p o p e n or male; Oldroyd 1969
closed. Legs simple, apical spurs o n the tibiae variable, h o w
ever, m i d leg virtually a l w a y s w i t h 2 apical spurs; e m p o d i u m p u l v i l l i f o r m .
Biology: T h e larvae o f m o s t species breed i n m o i s t soil, s o m e are found in m o s s e s , liver
w o r t s , etc., w i t h the larvae of m a n y Chrysopilus species breeding inside decaying w o o d o f
d e c i d u o u s trees. T h e larvae prey o n insect larvae, w o r m s and other invertebrates; s o m e
species are k n o w n to feed o n organic detritus and carrion. T h e adults are generally less
abundant, b e i n g found in shaded habitats near the verges of w o o d l a n d s and near bodies o f
water. S o m e Rhagionidae, including the w e l l k n o w n Rhagio scolopaceus (Linnaeus), are
often seen o n leaves, tree t r u n k s and other vertically oriented objects. Symphoromyia
species are k n o w n to feed o n b l o o d of m a m m a l s . M o s t o f the other species feed o n nectar,
p o l l e n and h o n e y d e w in the adult stage; m a n y species are p r e s u m e d to prey as w e l l o n
small insects and other invertebrates.
Identification references: V a n der G o o t 1985 ( N o r t h w e s t e r n Europe); Nartshuk 1989b
(former U S S R ) ; O l d r o y d 1 9 6 9 ; Stubbs 8t D r a k e 2001 (British Isles); T h o m a s 1997
(species); C M P D : Majer 1997a; C A T : Majer 1988b.

Rhinophoridae (key couplet 185; fig. 4 0 0 , 4 0 3 - 4 0 4 , 4 1 0 , 4 1 2 ,

414-416)
Systematics: Calyptrate Brachycera; superfamily Oestroidea;
s o m e 11 genera and about 4 5 species in Europe.
Characters: Small to large (2-11 m m ) flies, often w i t h m a n y bris
tles; usually grey to black w i t h s o m e p r u i n o s i t y or b r o w n i s h
y e l l o w . Arista bare to p l u m o s e . W i n g in s o m e cases distinctly
marked; v e i n M i a l w a y s c u r v i n g forward; cell r4+5 s o m e t i m e s
petiolate. L o w e r calypter a l w a y s small, o v o i d , s p o o n - s h a p e d ,
erect, laterally e x t e n d e d (i.e., n o t r e c u m b e n t o n the thorax).
S u b s c u t e l l u m n a r r o w and distinct, o n l y w e a k l y c o n v e x ; meral
bristles a l w a y s present a l t h o u g h fewer i n n u m b e r and less 553 Phyto melanocephala (Meigen),
strong than in related families; posterior spiracle w i t h a lining male; Mihalyi 1986
of l o n g hair-like bristles, w i t h o u t lappets.
Biology: A s far as k n o w n , the larvae live as endoparasitoids in terrestrial w o o d l i c e
(Isopoda: Oniscidea). T h e e g g s are deposited o n trees, w a l l s , u n d e r n e a t h stones, etc., and
after h a t c h i n g the y o u n g larva actively searches for its host. Pupation takes place w i t h i n
the e x o s k e l e t o n o f the w o o d l o u s e w h i c h b y that time is n o m o r e than an e m p t y shell. T h e
adults are often found at sunlit spots, o n w a l l s , flowers, w i n d o w s , etc.
Identification references: Draber M o n k o 1989 (Poland); Herting 1961 (Lindner); M i h a l y i
1986 (Central Europe); Stackelberg I 9 8 g b f (former U S S R ) ; Zeegers 8t V a n V e e n 1993
(Netherlands); C M P D : Pape 1998b; C A T : H e r t i n g 1993.
(Rhyphidae): n a m e u s e d i n the past for A n i s o p o d i d a e .
(Risidae): part o f the Ephydridae.
(Sapromyzidae): n a m e used in the past for Lauxaniidae.

Sarcophagidae (key couplet 188-189; fig- 419-420)

Systematics: Calyptrate Brachycera; superfamily Oestroidea;
s o m e 30 genera and about 310 species in E u r o p e .
Characters: Small to v e r y large (3-22 m m ) , usually stout and
bristly flies. M o s t species grey to greyish black w i t h a charac
teristic, shifting, checkered pattern o n their a b d o m e n ; s o m e
species y e l l o w - g r e y , green-grey or a l m o s t entirely black l u s
trous, never metallic blue or green. A r i s t a bare to p l u m o s e .
W i n g usually hyaline, s o m e t i m e s infuscated anteriorly or var
iously o r n a m e n t e d w i t h dark markings; v e i n M i a l w a y s c u r v
ing forward, s o m e t i m e s e n d i n g i n v e i n R4+5 at the w i n g mar
gin or before, m a k i n g cell r.4+5 petiolate. L o w e r calypter u s u a l
ly broadly rounded-triangular, in a v e r y f e w cases smaller and 554 Sarcophaga carnaria (Linnaeus),
more or less r o u n d to elliptical. S u b s c u t e l l u m n a r r o w and flat, female; Oldroyd 1970
not s w o l l e n ; meral bristles a l w a y s present; posterior spiracle
covered b y an anterior and posterior lappet, usually the anterior lappet a n a r r o w fringe
and the posterior o n e f o r m i n g an o p e r c u l u m .
Biology: T h e life m o d e o f the larvae is m o s t variable: b r o o d parasitoids o f H y m e n o p t e r a
such as Sphecidae, Pompilidae and, in the M i l t o g r a m m i n a e , A p o i d e a ; feeding o n excre
m e n t (coprophagous larvae); inquilines p r e y i n g o n larvae and pupae o f b u m b l e b e e s and
social w a s p s ; feeding o n carrion (necrophagous larvae) o f fish, crustaceans, amphibians,
reptiles, birds and m a m m a l s ; n e c r o p h a g o u s w i t h all transitional stages into endopara-
sitism o n grasshoppers, beetles and caterpillars and pupae o f butterflies, o n true b u g s
(Hemiptera), sawflies and w o o d w a s p s ( H y m e n o p t e r a : S y m p h y t a ) , o n centipedes and
millipedes, o n e a r t h w o r m s and snails; parasitoids in pupae o f Lepidoptera and
H y m e n o p t e r a : S y m p h y t a , o n e g g clusters o f grasshoppers and spiders, o n e g g s of reptiles
(including those o f lizards); facultative and m a n d a t o r y agents o f myiasis in vertebrates
and m a n . A d u l t s can b e found in a w i d e range o f habitats; s o m e species prefer coastal
habitats. T h e males in particular often visit flowers.
Identification references: Pape 1987 ( N o r t h w e s t e r n Europe); P o v o l n y 8t V e r v e s 1997
(Central Europe); B . B . R o h d e n d o r f 1930-1982 (Lindner, m a i n l y M i l t o g r a m m i n a e ) , 1937,
1989 (former U S S R ) ; V e r v e s 1982-1993 (Lindner), 1 9 9 4 ( M i l t o g r a m m i n a e ) ; C M P D : Pape
1998a; C A T : V e r v e s 1986; Pape 1996; Pape 8c Carlberg 2001.

Scathophagidae (key couplet 177,179; fig. 393-394)

Systematics: Calyptrate Brachycera; superfamily Muscoidea;
s o m e 38 genera and about 160 species in Europe.
Characters: Small to large (3-12 m m ) , usually slender flies.
C o l o u r ranging from a dull y e l l o w i s h b r o w n to lustrous black
or y e l l o w , in s o m e species b i c o l o u r o u s . B o d y and legs often
w i t h m a n y bristles, s o m e t i m e s densely c o v e r e d b y fine hairs.
O c c i p u t usually w i t h s o m e to m a n y pale, l o n g hairs; arista bare
to p l u m o s e ; interfrontal bristles absent. W i n g usually clear,
s o m e t i m e s distinctly marked or darkened at the tip or along
the crossveins; anal v e i n l o n g , usually reaching the w i n g mar
gin. M e r o n w i t h o u t bristles along the h i n d margin, near the
posterior spiracle.
Biology: T h e b i o l o g y o f the species in this family varies w i d e l y . Scathophaga stercoraria (Linnaeus)
In m o s t species the larvae are p h y t o p h a g o u s ; s o m e are leaf- male; Stackelberg 1956
miners or living i n flower heads (inflorescences), seed cap
sules, stems, etc. In other species the larvae feed or prey o n small organisms in rotting
organic matter such as s e a w e e d , nesting material and d u n g ; or they prey o n e g g s of aquat
ic insects. T h e adults prey m a i n l y o n small soft b o d i e d insects and other organisms. S o m e
species are attracted to d u n g in considerable n u m b e r s . O n e o f the b e s t - k n o w n e x a m p l e s
 o f this family is Scathophaga stercoraria (Linnaeus). T h i s species, o f w h i c h the reddish
b r o w n , densely pilose males gather o n c o w d u n g , m a y b e o b s e r v e d nearly all year round.
Identification references: A n d e r s s o n 1997b (species); D e l y - D r a s k o v i t s 1981; G o r o d k o v
1989c (former U S S R ) ; Sifher 2003 (Central Europe); C M P D : D e Jong 2000; C A T :
G o r o d k o v 1986.

(Scatomyzidae): n a m e used in the past for Scathophagidae.

Scatopsidae (key couplet 23, 201; fig. 58-59,436)

Systematics: N e m a t o c e r a ; superfamily Scatopsoidea; s o m e 22
genera and about 100 species in Europe.
Characters: M i n u t e t o small (0.5-4 m m ) , stout Nematocera.
B o d y colour dull or lustrous, dark to black, occasionally w i t h
lighter spots or m o r e reddish. Ocelli present; e y e s reniform
and w i t h an e y e bridge present i n almost all species; antenna
short, w i t h 7-12 s e g m e n t s , the terminal s e g m e n t often s o m e
w h a t longer. W i n g short and w i d e , w i t h an anal lobe, clear or
slightly tinged; the R-veins w e l l d e v e l o p e d and e n d i n g far
short o f the w i n g tip, the other v e i n s weaker. Legs relatively
short and stout; tibiae w i t h o u t apical bristles or spurs.
Biology: T h e larvae live in rotting organic matter, in all sorts o f
substrates such as rotting plants, vegetables and fruits, in or
u n d e r n e a t h d e c a y i n g w o o d and bark, rot holes in trees, soil,
556 Ectaetia clavipes (Loew). fema-
leaf litter, m u s h r o o m s and bracket fungi, d u n g , carrion, ants le; Haenni 2004
nests, etc. T h e adults are found in a w i d e range o f habitats b u t
generally prefer m a r s h y and m o r e o p e n areas w h e r e t h e y often visit flowers. S o m e species
form s w a r m s consisting o f thousands o f individuals o n l o w vegetation and shrubs.
Identification references: A n d e r s s o n 1982 ( S w e d e n ) ; C o o k 1 9 6 9 - 1 9 7 4 ( r e v i s i o n ) ;
Freeman & Lane 1985 (British Isles); K r i v o s h e i n a 1989a (former U S S R ) ; C M P D : Haenni
1997a; C A T : K r i v o s h e i n a 8t H a e n n i 1986.

Scenopinidae (key couplet 5 9 , 6 3 ; fig. 125-126,132)

Systematics: L o w e r m u s c o m o r p h Brachycera; superfam
ily A s i l o i d e a ; i n E u r o p e 2 genera, Caenoneura a n d
Scenopinus, and about 16 species.
Characters: Small to m e d i u m sized (2-7 m m ) flies, usual
ly dark coloured, stout and s o m e w h a t dorsoventrally
flattened. Head and thorax pubescent, d e v o i d of bristles.
Eyes nearly a l w a y s holoptic in the male and consisting o f
t w o differently faceted parts; antenna usually oriented
d o w n w a r d , w i t h 3 segments; the third s e g m e n t is the
largest and i n s o m e cases split at the e n d a n d / o r bearing
a m i n u t e style. W i n g clear or tinged; v e i n M i c u r v i n g
w e l l forward, e n d i n g in the costa before the w i n g tip, or
in v e i n R 5 ; cell m 3 o p e n ; cell c u p closed. Legs relatively 557 Scenopinus fenestralis (Linnaeus),
short and p o w e r f u l , e m p o d i u m bristle-like or absent. female; Toth 1977
Biology: T h e larvae live in a v a r i e t y o f d r y substrates such
as soil, dust, nests of birds and other animals, tunnels m i n e d b y beetles and other organ
i s m s , holes in w o o d , under bark, etc., w h e r e t h e y prey o n small invertebrates, in particu
lar o n m i t e s and beetles. T w o species w i t h a w o r l d - w i d e distribution, Scenopinus fenes-
tralis (Linnaeus) and S. glahrifrons M e i g e n , are w e l l - k n o w n predators o f carpet beetles.
The adults feed o n nectar and visit f l o w e r s . A d u l t s are n o t often encountered in the w i l d
b u t m o r e regularly found indoors o n w i n d o w s or in r o o m s w i t h lots o f dust and d r y
organic matter (disused r o o m s , attics, church t o w e r s ) .
Identification references: Kelsey 1 9 6 9 (revision); N a r t s h u k i g 8 9 g (former U S S R ) ;
O l d r o y d 1969; S t u b b s & Drake 2001 (British Isles); C M P D : Krivosheina I997e; C A T :
Kelsey & S o o s 1989.

158
Sciaridae (key couplet 28, 31, 200, 204-205; fig. 69-70, 73-74,
435)
Systematics: N e m a t o c e r a ; s u p e r f a m i l y Sciaroidea; s o m e 28
genera and about 620 species in Europe. T h e genera
Heterotricha and Sciarosoma, m e n t i o n e d under k e y couplet 37,
are s o m e t i m e s tentatively placed in the Sciaridae.
Characters: Delicate, often black or rarely black and y e l l o w to
orange coloured Nematocera, m i n u t e to m e d i u m sized (1-6
m m , rarely u p to 10 m m ) . Ocelli present; e y e s usually w i t h an
eye bridge; antenna w i t h 16 simple s e g m e n t s . W i n g clear, in
s o m e cases tinged; w i n g v e n a t i o n rather constant: costa e n d -
ing in b e t w e e n R4+5 and M i , the only crossveins present are H
and R - M ; v e i n M forked at about half its l e n g t h ; C u A i and
C u A 2 w i t h a basal stem. Legs relatively l o n g , tibiae w i t h apical 5 5 8 L y c o r i e | | a s Q | a n ( w n n e r t z )

bristles or spurs. i m a n 1983

m a e ; F r e e

Biology: T h e larvae are largely terrestrial b u t restricted m a i n l y

to w e t habitats. T h e y are an i m p o r t a n t faunal e l e m e n t in soil and litter layers, b u t can also
be found in dead w o o d , excrement, m u s h r o o m s , bracket fungi, etc.; a n u m b e r o f species
d e v e l o p i n semi-aquatic habitats. Larvae o f Sciaridae m a i n l y feed o n m y c e l i u m . A limited
n u m b e r o f species feed o n l i v i n g plant tissues such as roots or, as miners, leaves and
stems, or live in m u s h r o o m s and bracket fungi. S o m e species are pests in forestry, horti-
culture and agriculture, m a i n l y in m u s h r o o m culture and in glasshouses. T h e larvae o f
s o m e species inhabit caves, others live in colonies o f social insects. Larvae m a y occur in
v e r y h i g h densities; massive processions o f 2 to 10 metres in l e n g t h , so-called 'army
w o r m s ' , are w e l l - d o c u m e n t e d . T h e adults are found in a w i d e range o f habitats as w e l l ,
mainly near the larval habitat. T h e y are often found indoors, in w h i c h case t h e y probably
hatched from larvae that d e v e l o p e d i n p o t t i n g soil o f indoor plants.
Identification references: Freeman 1983 (British Isles); G e r b a c h e v s k a y a 1989 (former
U S S R ) ; M e n z e l 2000 ( G e r m a n y ) ; M e n z e l 8t M o h r i g 2000 (revision); C M P D : M e n z e l 8t
M o h r i g 1997; C A T : Gerbachevskaya 1986; M e n z e l et al. 2006 (British Isles).

Sciomyzidae (key couplet 1 3 7 , 1 4 1 , 1 4 3 , 1 5 4 ; fig. 1 8 8 , 3 0 2 , 3 0 7 -

309.334)
Systematics: Acalyptrate Brachycera; superfamily
Sciomyzoidea; traditionally this family includes the
P h a e o m y i i d a e ; s o m e 2 4 genera and a b o u t 1 4 0 species i n
Europe.
Characters: Small to large (2-14 m m ) , generally slender flies
w i t h a b o d y colour v a r y i n g from black lustrous to dull grey,
b r o w n , reddish or y e l l o w . A n t e n n a s o m e t i m e s elongate; arista
pubescent or w i t h shorter or longer rays; ocelli present; O c -
bristles usually present, rarely (Sepedn) absent; P-bristles
d i v e r g i n g to parallel, s o m e t i m e s s l i g h t l y c o n v e r g i n g i n
Pteromicra; 1-2 pairs o f F-bristles, c u r v i n g b a c k w a r d , t h e
l o w e r pair s o m e t i m e s c u r v i n g i n w a r d ; interfrontal bristles
absent; interfrontal setulae s o m e t i m e s p r e s e n t ; v i b r i s s a e l , ^ ^ ^ ^ ! ! ^5
' ( 9 1 1 F a b r l c i u s
)'
r
. .... , ., . , . male; Rivoseccni 1992
absent. W i n g clear or w i t h c o n s p i c u o u s markings; costa c o n -
tinuous; v e i n Sc complete; crossvein B M - C u present; cell c u p
closed. Tibiae almost a l w a y s w i t h dorsal preapical bristle.
Biology: T h e larvae o f nearly all species feed o n snails or slugs, s o m e feeding o n t h e eggs o f
snails. M o s t o f the k n o w n larvae are semi-aquatic, s o m e are aquatic; a n u m b e r o f species
have terrestrial larvae. Larvae m a i n l y prey o n snails lacking an o p e r c u l u m ; in general,
aquatic larvae have to visit t h e surface frequently in order to breathe b u t s o m e species
preying o n bivalves have adapted to breathing under water. T h e larvae o f s o m e species
preying o n terrestrial snails and slugs are parasitoids that d e v e l o p inside a single snail or
slug; in other terrestrial species t h e penultimate instar emerges from the snail or slug it
d e v e l o p e d in and are as last instar predatory o n several snails. T h e adults are often found

159
 sitting o n the vegetation, their heads d o w n . D e p e n d i n g o n t h e larval habitat, they are
often found near water, in m a r s h y vegetation, b u t also in w o o d l a n d or e v e n d r y o p e n
habitats.
Identification references: R e v i e r & V a n d e r G o o t 1989 ( N o r t h w e s t e r n E u r o p e ) ;
R o z k o s n y 1 9 8 4 (Fennoscandia and D e n m a r k ) , 1987 (review); i 9 9 7 d (genera), 2002
(species); Stackelberg 19893c (former U S S R ) ; Vals 1989 (France, Southern Europe);
C M P D : R o z k o s n y 1998a; C A T : R o z k o s n y & Elberg 1984.

(Scopeumstidae): n a m e used in t h e past for Scathophagidae.

Sepsidae (key couplet 91, 113, 138, 156; fig. 1 9 4 - 1 9 9 ,

244.305.338)
Systematics: A c a l y p t r a t e B r a c h y c e r a ; s u p e r f a m i l y
S c i o m y z o i d e a ; 9 g e n e r a a n d a b o u t 50 s p e c i e s i n
Europe.
Characters: Small to m e d i u m sized (2-6 m m ) , slender,
ant-like flies w i t h relatively f e w bristles or hairs. B o d y
colour generally black, i n s o m e cases lustrous a n d / o r
w i t h a distinct pattern o f silvery p r u i n o s i t y o n t h e side
of the thorax. In general, w a l k i n g about o n vegetation
w h i l e w a v i n g their w i n g s . A n exception is Orygma luc- 560 Sepsis cynipsea (Linnaeus),
tuosum M e i g e n , a robust, hairy, s o m e w h a t flattened male; Oldroyd 1970
fly. T h e Sepsidae, including Orygma, are distinct from
all other A c a l y p t r a t e flies o n account o f the presence o f 1 or m o r e bristles at the p o s -
teroventral m a r g i n o f the posterior spiracle. Head rounded, arista bare; ocelli present; O c -
bristles present; P-bristles d i v e r g i n g or absent; 0-3 pairs o f F-bristles, i f present c u r v i n g
o u t w a r d or (obliquely) b a c k w a r d ; interfrontal bristles absent, interfrontal setulae present
in Orygma; vibrissae or vibrissa-like bristles u s u a l l y present. W i n g clear or tinged, often
w i t h a dark spot or c l o u d i n g at the w i n g tip; costa c o n t i n u o u s ; v e i n Sc complete; crossvein
B M - C u present; cell c u p closed. In the male femur and tibia o f fore legs often w i t h extru
sions, spurs, teeth and t h e like. Tibiae usually w i t h dorsal preapical bristle, b u t in several
genera small or absent (e.g. Meroplius, Nemopoda, Themira).
Biology: T h e larvae o f m o s t Sepsidae largely d e v e l o p i n d u n g , excrement, rotting organic
matter such as dead snails and insects, m u s h r o o m s and bracket fungi, in nests o f birds, in
soil rich i n organic c o m p o u n d s , in c o m p o s t , etc. T h e adults are generally o b s e r v e d near
the larval habitats, usually in s o m e w h a t o p e n habitats such as m e a d o w s , along water
courses, forest margins and clearings in w o o d l a n d areas. S o m e species occur in major
s w a r m s o n l o w vegetation and shrubs, w a v i n g their w i n g s . T h e s e s w a r m s can persist for
over a m o n t h and are often accompanied b y a certain odour. A l l this does n o t account for
Orygma luctuosum w h i c h is a seaside species, to b e found under cast u p seaweed.
Identification references: Pont 1979 (British Isles); D z e r o v 2003 (Russia); Pont 8t Meier
2002 (Europe); Stackelberg 19893b (former U S S R ) ; C M P D : Meier 8t Pont 2000; C A T :
Z u s k a & Pont 1984; O z e r o v 2005.

Similliidae (key couplet 17; fig. 50-51)

Systematics: N e m a t o c e r s ; superfamily C h i r o n o m o i d e a ;
8 genera s n d about 230 species in Europe.
Characters: B l o o d s u c k i n g , m i n u t e to m e d i u m sized (1.2-
6 m m ) , stout, usually dark coloured Nematocera. Eyes
holoptic in the male, dichoptic in the female; ocelli
absent; antenna short and stout, tapering t o w a r d t h e
end, w i t h 11, rarely 9 segments. W i n g short and w i d e ,
w i t h a distinct anal lobe; w i n g clear, in s o m e cases
slightly tinged; costa and R - v e i n s stronger than t h e
other v e i n s ; v e i n C u A 2 usually c u r v i n g like an elon
gate, inverted S. Legs short and stout. 561 Simulium ornatum Meigen,
Biology: T h e larvae are found in several t y p e s of r u n n i n g female; Seguy 1951

16o
 water, attached to a substrate like stones, m e t a l objects, plastic bags, plants, branches, etc.,
at or above the b o t t o m . T h e y live o n m i c r o - o r g a n i s m s and small organic particles they fil
ter from the water. Pupation takes place inside a c o c o o n . T h e adults feed o n nectar and
plant juices, females o f a n u m b e r o f species feed o n the b l o o d o f b o t h c o l d - b l o o d e d and
w a r m - b l o o d e d vertebrates and are k n o w n as 'black flies'. Feeding takes place in daylight,
mainly during the m o r n i n g and e v e n i n g h o u r s . In the tropics, b u t also in the tundra, taiga
and s o m e East European regions, Simuliidae can occur in such vast n u m b e r s as to inter
fere w i t h , or entirely prevent, h u m a n activity.
Identification references: Bass 1998 (larvae and pupae British Isles); C r o s s k e y 1993a (gen
era feeding o n b l o o d ) ; D a v i e s 1968 (British Isles); Jedlicka et al 2 0 0 4 (pupae Central
Europe); Jankovsky 2002 (former U S S R ) ; Jensen 1997 (genera); Lechthaler &. Car 2 0 0 4
(larvae pupae Central and W e s t e r n Europe); R i v o s e c c h i 1978 (Italy); R u b t z o v 1 9 5 9 - 1 9 6 4
(Lindner), 1989 (former U S S R ) ; Y a n k o v s k i i 2003 (former U S S R ) ; C M P D : Jedlicka &
Stloukalova 1997; C A T : R u b t z o v & Y a n k o v s k y 1988.

(Siphonellopsidae): part o f the Chloropidae.

(Solvidae): n a m e used in the past for X y l o m y i d a e .
(Spaniidae): part o f the Rhagionidae.

Sphaeroceridae (key couplet 85,213;fig.1 8 4 - 1 8 5 , 4 4 7 )

Systematics: Acalyptrate Brachycera; superfamily
Sphaeroceroidea; s o m e 4 0 genera and about 260 species in
Europe.
Characters: M i n u t e to m e d i u m sized (0.7-5.5 m m ) , stout, u s u
ally dull b r o w n to black coloured flies, characterised b y pres
ence o f vibrissae in c o m b i n a t i o n w i t h short, s w o l l e n first
tarsal s e g m e n t o f the h i n d leg. Arista bare to l o n g pubescent,
ocelli virtually a l w a y s present; Oc-bristles present or absent;
P-bristles c o n v e r g i n g or absent; 1-2 pairs o f F-bristles, c u r v i n g
o u t w a r d ; interfrontal bristles present, partially i n r o w s ori
ented i n w a r d ; vibrissae present. W i n g almost a l w a y s w i t h o u t
markings; costa w i t h breaks at the base, a subcostal break and
in s o m e cases a h u m e r a l break as w e l l ; v e i n Sc reduced;
crossvein B M - C u present or absent; cell c u p o p e n or closed. 5 6 2
Leptocera fontinalis Fallen, female;
P a p p 1 9 7 3
Tibiae w i t h or w i t h o u t dorsal preapical bristle.
Biology: T h e larvae live in a w i d e range of rotting organic mate
rials and presumably feed o n m i c r o - o r g a n i s m s . T h e y can b e found in d u n g , manure
d u m p s , leaf litter, organic matter w a s h e d u p o n the shore, m u s h r o o m s , bracket fungi, rot
ting vegetable matter, in caves, nests and other places inhabited b y animals. T h e adults are
mediocre fliers. T h e y can be found o n d e c a y i n g matter o f plant and animal origin, prefer
ably in d a m p habitats; relatively m a n y species live in caves or animal b u r r o w s . A f e w
species occur near h u m a n settlements and, i n case o f m a s s occurrence in rotting sub
strates or i n manure near h u m a n d w e l l i n g s , they m a y b e c o m e a nuisance because t h e y
usually also c o m e indoors. M a n y species l i v i n g in m a n u r e and d u n g are associated w i t h
hoofed animals and often form large p o p u l a t i o n s o n pastures and d u n g heaps near stables.
The family s h o w s a h i g h percentage o f species w i t h reduced w i n g s , or n o w i n g s at all.
Identification references: Nartshuk 1989b (former U S S R ) ; Pitkin 1988 (British Isles);
C M P D : Rohacek 1998c; C A T : Rohacek et al. 2002; C A T : Papp 1984L

161
Stenomicridae (key couplet 104; fig. 230-231)
Systematics: Acalyptrate Brachycera; superfamily
O p o m y z o i d e a ; traditionally part o f the Aulacigastridae or
Periscelididae; i n Europe 1 g e n u s , Stenomicra, w i t h 3 species.
Characters: M i n u t e (1-2 m m ) , delicate, y e l l o w to grey flies w i t h
inner vertical bristles c u r v i n g forward. Arista w i t h l o n g rays;
ocelli present w i t h the ocellar triangle situated m o r e or less in
m i d d l e o f frons; O c - b r i s t l e s absent; P-bristles small a n d
diverging or absent; 2 pairs o f F-bristles, l o w e r pair c u r v i n g
i n w a r d ; interfrontal bristles absent; vibrissae and a series o f
vibrissa-like bristles present. W i n g clear or w i t h tinged spots;
costa w i t h a subcostal break; v e i n Sc incomplete; crossvein
B M - C u absent; crossvein D M - C u present or absent; cell c u p
o p e n . Tibiae w i t h o u t dorsal preapical bristle.
Biology: T h e b i o l o g y o f this small family is little k n o w n . T h e
larvae probably d e v e l o p i n the w a t e r - h o l d i n g leaf bases o f
plants, particularly m o n o c o t y l e d o n s . T h e adults o f s o m e species often occur near water or
at m o i s t places and are closely associated w i t h t u s s o c k s o f large Carex species, others also
w i t h Typha, Scirpus or Cyperus; the species Stenomicra soniae Merz &. Rohacek may be
associated w i t h umbellifers. A s in s o m e C h a m a e m y i i d a e , the adults are able to w a l k side
w a y s and b a c k w a r d w i t h the same ease as forward.
Identification references: C o l l i n 1 9 4 4 , I r w i n 1982 (British Isles); M e r z 8t Rohacek 2005
(species); Stackelberg 198930 (former U S S R ) ; C M P D : Mathis 8t Papp 1998; C A T : Papp
1984^

( S t o m o x y d i d a e ) : part o f the Muscidae.

Stratiomyidae (key couplet 4 2 , 5 4 , 7 0 ;fig.8 8 , 1 0 3 , 1 1 6 , 1 5 0 )

Systematics: L o w e r Brachycera; superfamily Stratiomyoidea;
s o m e 27 genera and about 1 4 0 species in Europe.
Characters: Small to v e r y large (2-25 m m ) flies o f slender to
stout shape, in s o m e cases flattened. B o d y colour often strik
ing: usually dark w i t h c o n s p i c u o u s w h i t e , y e l l o w , or greenish
patterns, s o m e t i m e s w i t h blue to green metallic lustre. Eyes
usually holoptic in male; antenna w i t h 7-10 s e g m e n t s , v a r y i n g
b e t w e e n rather u n i f o r m to basal s e g m e n t s s w o l l e n and termi
nal s e g m e n t s m a k i n g u p the style; m o u t h p a r t s often short, in
s o m e cases elongate. In m o s t species s c u t e l l u m w i t h spine 564 Sargus flavipes Meigen,
like e x t e n s i o n s . W i n g usually clear, in s o m e species w i t h male; Rozkosny 1982
markings or partially m u c h darkened; v e n a t i o n characteristic,
e.g. R - v e i n s in front part o f the w i n g w i t h v e i n R5 almost a l w a y s e n d i n g w e l l before the
w i n g tip; discal cell d i a m o n d - s h a p e d , small to v e r y small; cell m 3 o p e n ; cell cup closed.
Legs simple, almost a l w a y s w i t h o u t bristles, in a f e w species tibia of the m i d leg w i t h a sin
gle apical bristle; e m p o d i u m p u l v i l l i f o r m .
Biology: C o n t r a r y to m o s t other l o w e r flies, the larvae o f the Stratiomyidae are n o t preda
tors or parasitoids. T h e y are aquatic to terrestrial and m a i n l y feed o n algae and rotting
organic matter, in m o s t cases o f vegetable origin. T h e aquatic habitats range from stagnant
to r u n n i n g water, m o i s t rock faces, o n w a t e r vegetation, marshes, saline e n v i r o n m e n t s
along the seaside or inland. Terrestrial larvae often o c c u r u n d e r n e a t h leaves and other rot
ting vegetable matter, in the topsoil, in s o m e cases in d u n g , in ants nests and b e l o w the
bark o f trees. T h e adults feed o n nectar and h o n e y d e w . T h e y are frequent visitors to flow
ers and are also found sitting o n leaves o f herbs, shrubs, or trees. Preferred habitats include
w o o d l a n d , the v i c i n i t y o f water, dunes or other coastal habitats.
Identification references: B r u g g e 2002 (Netherlands); Nartshuk 1989c (former U S S R ) ;
O l d r o y d 1969 (British Isles); R o z k o s n y 1973 (Fennoscandia and D e n m a r k ) ; R o z k o s n y
1982-1983 (revision), 1997c (genera), 2000 (species); Stubbs & Drake 2001 (British Isles);
W o o d l e y 1995 (Beridinae); C M P D : R o z k o s n y 1997b; C A T : R o z k o s n y & Nartshuk 1988.

162
Streblidae (key couplet 1 6 9 , 2 0 9 ;fig.1 7 5 , 3 7 2 - 3 7 3 , 4 4 4 )
Systematics: Calyptrate Brachycera; superfamily
H i p p o b o s c o i d e a ; in S o u t h e r n Europe 1 species, Brachytarsina
flavipennis Macquart.
Characters: M i n u t e to small (0.8-5 m m ) , y e l l o w to y e l l o w i s h
b r o w n flies. S o m e n o n - E u r o p e a n species w i t h w i n g s reduced
or absent. Head small, n o t adpressed to the thorax; eyes and
ocelli r u d i m e n t a r y or absent. First tarsal s e g m e n t short; termi
nal tarsal s e g m e n t w i d e n e d .
Biology: T h e adults are ectoparasites o f bats, w i t h a preference
for colonial c a v e - d w e l l i n g M i c r o c h i r o p t e r a , m o s t s p e c i e s
b e i n g confined to a single species or g e n u s o f host. T h e larvae
d e v e l o p inside the female a b d o m e n o f the fly and are deposit
ed o n e b y o n e , generally in the direct v i c i n i t y o f bat roosts,
565 Brachytarsina flavipennis
after w h i c h they i m m e d i a t e l y pupate. A d u l t s o f fully w i n g e d Macquart, male; Hurka 1998b
species, although h i g h l y m o b i l e , remain m o s t closely associat
ed w i t h the host bat. T h e flies generally d o n o t leave the h o s t except for deposition o f the
larvae or w h e n the h o s t is disturbed.
Identification references: Stackelberg i 9 8 g b d (former U S S R ) ; C M P D : Hurka 1998b; C A T :
Hurka ck. Soos 1986b.

Strongylophthalmyiidae (key couplet 168; fig. 332, 368-

369)
Systematics: Acalyptrate Brachycera; superfamily D i o p s o i d e a ;
2 species in Europe: Strongylophthalmyia ustulata
(Zetterstedt) and S.pictipes Frey.
Characters: Small to m e d i u m sized (3.5-5.5 m m ) , slender, large
ly black flies w i t h y e l l o w i s h legs. Arista short pubescent; ocel
li present; O c - b r i s d e s present; P-bristles diverging; 2 pairs o f
F-bristles; interfrontal brisdes absent; vibrissae absent. W i n g
clear w i t h v a g u e markings in the central part or at the w i n g tip;
costa w i t h a subcostal break; v e i n Sc incomplete; crossvein
B M - C u present; cell c u p closed. Tibiae w i t h o u t dorsal preapi-
cal bristle.
Biology: T h e b i o l o g y o f the European species is little k n o w n .
566 Strongylophthalmyia ustulata
T h e larvae live under bark of dying or decaying deciduous trees, (Zetterstedt), female; Papp 1978
apparently w i t h a preference for w e t , dead ash w o o d . T h e adults
occur o n l o w vegetation in w e t deciduous forests near the breeding sites of the larvae.
Identification references: Krivosheina 1981; Shatalkin 1993 (review); Stackelberg 1989V
(former U S S R ) ; C M P D : Iwasa 1998a; C A T : Krivosheina 1984a.

Synneuridae (key couplet 24; fig. 60)

Systematics: Nematocera; superfamily Scatopsoidea; the fami
lies Canthyloscelidae and Synneuridae are often considered
one family; in N o r t h e r n , Central and Eastern Europe i species:
Synneuron annulipes Lundstrom.
Characters: Small (3-4 m m ) , stout, black Nematocera w i t h a
s o m e w h a t laterally c o m p r e s s e d thorax and elongate a b d o m e n .
Ocelli present; e y e s b e l o w the antenna n o t far apart or e v e n
t o u c h i n g ; antenna w i t h 12 short s e g m e n t s , the terminal seg 567 Wing Synneuron decipiens Hutson;
m e n t elongate. W i n g clear or slightly tinged; R - v e i n s stronger Peterson & Cook 1981 (North America)
than the other v e i n s . Legs relatively l o n g and stout.
Biology: T h e b i o l o g y o f the European species o f Synneuron is little k n o w n . T h e larvae live
in moist, decaying w o o d . A d u l t s are rare, at least in collections; t h e y are p r e s u m e d to
occur especially in old, undisturbed forests.
Identification references: A n d e r s s o n 1982 ( S w e d e n ) ; H u t s o n 1977 (revision); C M P D :
Haenni 1997b; C A T : M a m a e v 8t K r i v o s h e i n a 1986a.

163
Syrphidae (key couplet 6 4 ; fig. 135-137)
Systematics: Lower Cyclorrhapha; superfamily
Syrphoidea; s o m e 9 0 genera and about 830 species in
Europe.
Characters: Small to v e r y large (3.5-35 m m ) , slender to
stout flies, usually coloured black w i t h w h i t e or y e l l o w
m a r k i n g s o n head, thorax and especially a b d o m e n ;
s o m e t i m e s b o d y largely b r o w n , metallic y e l l o w , green,
or blue, or w i t h a w i d e range o f colours; in s o m e cases
strongly r e s e m b l i n g bees, b u m b l e b e e s , or w a s p s . Head
large, w i t h o u t any distinct bristles, m o s t l y consisting o f
e y e s w h i c h are u s u a l l y holoptic in the male. T h i r d 568 Scaeva pyrastri (Linnaeus),
antennal s e g m e n t usually the largest b u t s o m e t i m e s female; Oldroyd 1970
first or second s e g m e n t longer; third s e g m e n t carrying
an apical or dorsal arista or an apical style. T h o r a x w i t h v e r y f e w bristles. W i n g usually
clear, in s o m e cases tinged or w i t h markings; v e n a t i o n characteristic, virtually a l w a y s
w i t h a spurious v e i n (vena spuria), less distinct or absent in Eristalinus and Psilota; v e i n
M i n o t reaching the w i n g m a r g i n b u t c u r v i n g strongly forward and e n d i n g in v e i n R4+5
before the w i n g margin; cell d m present, cell b e l o w cell d m o p e n ; cell c u p elongate, end
ing acutely just before the w i n g margin. Legs u s u a l l y slender and simple b u t s o m e t i m e s ,
especially in the male, w i t h coxa, trochanter, femur, tibia, or tarsus modified a n d / o r bear
ing spurs.
Biology: T h e larvae can b e divided into m y c o p h a g o u s and p h y t o p h a g o u s species (includ
ing a n u m b e r o f species b e i n g pests in agriculture, especially bulb cultivation); predators
of other insects (such as aphids, ants, larvae of w a s p s or beetles, etc.), and species feeding
o n v a r i o u s t y p e s o f organic detritus (decaying w o o d , sap e x u d i n g from tree w o u n d s ,
organic detritus u n d e r n e a t h tree bark or in nests of birds and other animals, leaf litter, liq
uid e x c r e m e n t or other organically enriched w a t e r y solutions). In s o m e cases larvae of this
latter g r o u p inhabit semi-aquatic to entirely aquatic habitats. T h e adults are c o m m o n ,
c o n s p i c u o u s flies, to b e found in various o p e n to w o o d l a n d habitats such as gardens,
m e a d o w s , forest eaves, river banks, etc. T h e y are able to h o v e r in o n e place and are appar
ently capable o f m o v i n g n o t o n l y forward, b u t also b a c k w a r d and laterally. T h e adults feed
o n nectar and p o l l e n , and several o f the frequent visitors to flowers play an i m p o r t a n t part
in pollinating certain plant species.
Identification references: Barendregt 2001 (Netherlands); D o l e z i l & R o z k o s n y 1997
( g e n e r a ) ; H i p p a e t al. 2001 (Eristalis); K o r m a n n 2002 ( s p e c i e s ) ; R e e m e r 2 0 0 0
(Netherlands); R o t h e r a y 1993 (larvae); Stackelberg 1989P (former U S S R ) ; Speight 1999
(Xylotini); Stubbs 8t Falk 2002 (British Isles); V a n V e e n 2 0 0 4 ( N o r t h w e s t e r n Europe);
Verlinden 1991 (Belgium); C M P D : T h o m p s o n 8t R o t h e r a y 1998; C A T : Peck 1988.

Tabanidae (key couplet 50; fig. 2a, 6b, 2 0 , 9 2 , 1 1 0 - m )

Systematics: L o w e r Brachycera; superfamily Tabanoidea;
s o m e 13 genera and about 220 species in Europe.
Characters: Stout, m e d i u m sized to v e r y large (6-30 m m )
flies, d e v o i d o f bristles. B o d y generally b r o w n , black or
grey and w i t h a pattern o f lighter markings; i n s o m e
cases b o d y y e l l o w i s h , greenish or metallic blue. Eyes in
living flies often brilliantly reddish or metallic green,
s o m e t i m e s y e l l o w i s h , often w i t h spots or bands; e y e s in
male holoptic, i n female dichoptic; antenna w i t h 6 or
m o r e segments, u s u a l l y w i t h a large third s e g m e n t and a
tapering style; in m o s t females the m o u t h p a r t s consti
tute a p o w e r f u l , often l o n g piercing proboscis. W i n g
clear, or tinged, or w i t h transverse bands, or darkened
w i t h distinct spots; the costa s u r r o u n d i n g the w i n g
entirely; fork o f v e i n s R 4 a n d R 5 u s u a l l y s t r o n g l y
diverging w i t h v e i n R5 e n d i n g distinctly b e y o n d the
w i n g tip; cell IT13 o p e n ; cell c u p o p e n or closed. L o w e r

164
 calypter c o n s p i c u o u s l y large. Legs p o w e r f u l , tibia o f m i d leg and i n s o m e cases h i n d leg
w i t h apical spurs; e m p o d i u m p u l v i l l i f o r m .
Biology: T h e larvae are aquatic, semi-aquatic or terrestrial (living m a i n l y in soil and litter).
In m o s t genera they are predators b u t also able to feed as facultative saprophages. T h e y
prey o n other invertebrates such as w o r m s , snails and the larvae o f other Diptera, includ
ing those o f their o w n species. T h e adults are able fliers and especially active o n w a r m ,
sunny days. In the majority o f species the females feed o n b l o o d o f m a m m a l s i n c l u d i n g
h u m a n s ; b o t h sexes also feed o n nectar and pollen. S o m e species s h o w territorial b e h a v
iour, the males safeguarding their territories b y h o v e r i n g . A s a c o n s e q u e n c e of their b l o o d
feeding habits and associated transmission o f diseases, Tabanidae can be a serious p r o b
lem locally for h u m a n s as w e l l as cattle.
Identification references: Chvala et al. 1972 (revision); C h v a l a & Jezek 1997 (genera);
Kniepert 2000 (species); Majer 1987 (Central Europe); O l d r o y d 1969 (British Isles);
O l s u f y e v 1989 (former U S S R ) ; Portillo R u b i o 2002 (Iberian Peninsula); Stubbs & Drake
2001 (British Isles); T i m m e r 1980 (Belgium, Netherlands and L u x e m b o u r g ) ; Trojan 1979
(Poland); Zeegers 8t Haaren 2000 (Netherlands); C M P D : n o contribution; C A T : C h v a l a
1988b.

Tachinidae (key couplet 1 7 1 - 1 7 2 , 1 8 1 , 1 9 0 , 2 1 9 ; fig. 11,374-375, /f\\

386-389,399,402,405-406) / / / / I
Systematics: Calyptrate Brachycera; superfamily Oestroidea; lllAyi
s o m e 280 genera and about 880 species in Europe. Ml// i
Characters: Small to large (2-20 m m ) flies. M o s t species are grey ML/ , 1
to slate black, in s o m e cases w i t h y e l l o w or red spots o n the sides / LmSS&.i / /
of abdomen; some species are metallic green or y e l l o w i s h grey. WBSA&ib.I&
Usually many bristles are present, some species may, however, WJv*0^^~**
be quite bare or s h o w a bee-like pilosity. Arista usually quite ^ - r ^ ^ f s S K S S S

bare, rarely p l u m o s e . W i n g w i t h v e i n M i usually strongly IT ~"^5>

curved forward, sometimes cell r4+5 is petiolate. L o w e r calypter \
usually broadly rounded-triangular, in s o m e cases small and ^ T =

elliptical. Meral brisdes always present; subscutellum strongly f \

developed (except in Litophasia); lappets o f posterior spiracle \
usually distinctly unequal in size, the posterior lappet subcircu-
lar, in several genera about equal in size. 570 Mintho rufiventris Fallen,
m a l e ; M l h a | 1 9 8 6
Biology: T h e larvae are endoparasitoids o f other insects, s o m e - y'
times o f other arthropods. Eggs are deposited directly o n or i n
the i m m e d i a t e v i c i n i t y o f the host. T h e larvae o f tachinids w h i c h deposit their e g g s
directly o n the host hatch w i t h i n m i n u t e s (ovolarvipary) and penetrate the host. A c o n
spicuous specialisation is the d e v e l o p m e n t o f m i c r o t y p e eggs b y species o f the tribe
G o n i i n i in the subfamily Exoristinae. Here, tens o f t h o u s a n d s o f e g g s are deposited o n
leaves. T h e eggs only d e v e l o p after a caterpillar has eaten t h e m (along w i t h the leaves t h e y
feed on), after w h i c h they hatch inside the caterpillar's digestive tract. Hence, m o s t
G o n i i n i s h o w a w i d e range of lepidopterous hosts. S o m e other groups s h o w h o s t special
isation: the D e x i i n i are parasitoids o f beetles, the Exoristinae and Tachininae are m a i n l y
parasitoids o f caterpillars and the larvae o f the Phasiinae d e v e l o p inside true b u g s
(Hemiptera); others parasitise besides caterpillars also larvae o f Tipulidae (Tachininae:
Siphona). Several species are of e c o n o m i c importance, b e i n g useful in biological pest c o n
trol. T h e adults o f the Tachinidae are largely found o n w a l l s , leaves, branches and similar
spots; only a small n u m b e r o f species regularly visit flowers. S o m e species are territorial.
Identification references: A n d e r s e n 1996 (Siphonini); B e l s h a w 1993 (British Isles);
Mesnil 1 9 4 4 - 1 9 8 0 (Lindner); M i h a l y i 1 9 8 6 , 1 9 9 4 ; Tschorsnig 8c H e r t i n g 1 9 9 4 (Central
Europe); Z i m i n et al. 1989 (former U S S R ) ; C M P D : Tschorsnig & Richter 1998; C A T :
Herting & D e l y - D r a s k o v i t s 1993.

(Taeniapteridae): part o f the Micropezidae.

165
Tanypezidae (key couplet 158; fig. 342-343)
Systematics: A c a l y p t r a t e B r a c h y c e r a ; s u p e r f a m i l y
D i o p s o i d e a ; i n Europe i w i d e s p r e a d species, Tanypeza
longimana Fallen.
Characters: T h e o n l y European species, Tanypeza longi
mana, is a slender, black lustrous, m e d i u m sized (5-8
m m ) fly w i t h silvery markings o n the head and t h e sides
o f the thorax, and w i t h long, slender, largely y e l l o w legs.
Head rounded; arista pubescent; ocelli present; O c - b r i s -
tles present; P-bristles diverging; 2 pairs o f F-bristles,
c u r v i n g b a c k w a r d ; interfrontal bristles absent; vibrissae
absent. W i n g u n m a r k e d , costa w i t h a w e a k subcostal
break; v e i n Sc reaching the costa, r u n n i n g parallel to v e i n
Ri and approaching or t o u c h i n g it just before reaching the 571 Tanypeza longimana Fallen,
male; Seguy 1934
costa; crossvein B M - C u present; cell c u p closed. Tibiae
w i t h o u t dorsal preapical bristle.
Biology: T h e larvae o f T. longimana feed o n rotting vegetable matter, possibly in decaying
w o o d or other plant detritus. T h e adults prefer l o w vegetation in m o i s t w o o d l a n d , in par
ticular near water.
Identification references: V a n der G o o t 8t V a n V e e n 1 9 9 6 ( N o r t h w e s t e r n Europe);
Stackelberg 1989U (former U S S R ) ; C M P D : Rohacek 1998a; C A T : S o o s 1984b.

(Tendipedidae): n a m e u s e d in the past for C h i r o n o m i d a e .

Tephritidae (key couplet 107,150; fig. 9,239,328-331)

Systematics: A c a l y p t r a t e B r a c h y c e r a ; s u p e r f a m i l y
Tephritoidea; s o m e 7 0 genera and about 270 species in
Europe.
Characters: G e n e r a l l y small to m e d i u m sized (2.5-10 m m ,
e x t r e m e s from 1-20 m m ) , usually colourful flies, w i n g
often w i t h characteristic m a r k i n g s and v e i n Sc abruptly
0
b e n t forward t o w a r d the costa at nearly 9 0 . A r i s t a bare
to short pubescent; ocelli present; Oc-bristles present; P -
bristles parallel to diverging; 2-8 pairs o f F-bristles, at
least 1 b u t usually several l o w e r pairs c u r v i n g i n w a r d , at 572 Ceratitis capitata (Wiedemann),
female; Balachowsky & Mesnil 1935
least 1 o f the u p p e r pairs c u r v i n g b a c k w a r d , in s o m e cases
F-bristles inserted o n a raised tubercle; interfrontal s e t u -
lae m o s t often absent or represented b y 1-2 tiny setulae near the lunula; vibrissae absent
b u t several genera w i t h strong bristles near the vibrissal angle. W i n g w i t h yellow, b r o w n
or black markings or dark coloured w i t h lighter markings; in a f e w species w i n g clear;
costa w i t h a h u m e r a l and a subcostal break; apical part o f v e i n Sc usually less distinct or
e v e n transparent and at about a right angle w i t h respect to the basal part; crossvein B M -
C u present; cell c u p closed, nearly a l w a y s n a r r o w i n g to an acute angle, closed b y a genic
ulate v e i n C u A 2 , v e i n C u A 2 rarely straight or c o n v e x . Tibiae w i t h o u t dorsal preapical bris
tle. Female w i t h oviscape, n o n retractable basal s e g m e n t o f the ovipositor.
Biology: T h e larvae o f nearly all Tephritidae are p h y t o p h a g o u s . T h e female uses her tele
scopic ovipositor to deposit e g g s in l i v i n g , healthy plant tissue. T h e larvae d e v e l o p in
flowers, seeds, fruits, leaves, stems or roots o f their h o s t plant, d e p e n d i n g o n t h e species.
S o m e species induce gall-forming. Several species are serious pests in agriculture and
horticulture, others play a role in the biological control of w e e d s . A n exception to the phy
t o p h a g o u s lifestyle are Euphranta toxoneura (Loew) a n d Chetostoma stackelbergi
R o h d e n d o r f w i t h l a r v a e d e v e l o p i n g i n galls f o r m e d b y s a w f l i e s ( H y m e n o p t e r a :
S y m p h y t a ) . A d u l t Tephritidae are g o o d fliers. T h e y are often found o n the host plant or
w h i l e feeding o n nectar, pollen, plant juices, rotting plant material or h o n e y d e w .
Identification references: Darvas & Papp 2000 (keys to exotic species occurring in Europe);
Kabos & Van Aartsen 1984 (Netherlands); Merz 1994 (Northern and Central Europe); Richter
i 9 8 g f (former U S S R ) ; W h i t e 1988 (British Isles); W h i t e 8t Elson-Harris 1994 (review of eco
nomically important species); C M P D : n o contribution; C A T : Foote 1984.

166
(Tetanoceridae): n a m e used in the past for S c i o m y z i d a e .

Tethitlidae (key couplet 1 1 6 , 1 1 8 , 1 6 8 ; fig. 251-252, 255, 367)

Systematics: Acalyptrate Brachycera; superfamily Carnoidea; 3
genera and about 35 species i n Europe.
Characters: M i n u t e to small (1.5-3.5 m m ) , y e l l o w to black flies
w i t h a grey, y e l l o w i s h , or b r o w n i s h pruinosity. Arista bare to
short pubescent; ocelli present; Oc-bristles present; P-bristles
converging; 1-5 pairs o f F-bristles, the u p p e r o n e s c u r v i n g out
w a r d or b a c k w a r d , the l o w e r pairs, i f present, c u r v i n g out
w a r d , forward a n d / o r inward; interfrontal bristles present or
absent; vibrissa-like bristles present, in s o m e cases small and
inconspicuous; i n the subfamily Tethininae a small, lustrous
bulge in b e t w e e n the antennae and the anterior vibrissa-like
bristle. W i n g unmarked, w i t h a pale y e l l o w to b r o w n i s h tinge; 573 Tethina czernyi (Hendel); male;
Merz 2002
costa w i t h a subcostal break; v e i n Sc first r u n n i n g parallel to
and then m e r g i n g w i t h R i just before the costa; crossvein B M -
C u absent or v e r y faded; cell c u p present b u t small. Tibiae w i t h o u t dorsal preapical bris
tle.
Biology: Tethinidae are almost exclusively h a l o p h i l o u s flies, m a i n l y found along the coast
although they occur i n saline inland habitats as w e l l ; only a f e w species are also k n o w n
from habitats that are apparently w i t h o u t increased salinity (forests, m e a d o w s , deserts).
T h e y are often present in great n u m b e r s . S o m e species are recorded from secondarily
man-created habitats, such as m e a d o w s polluted b y industrial emissions, slaughterhous
es and poultry farms, w h e r e they live synanthropically in alkaline d a m p places.
Identification references: B e s c h o v s k i 1 9 9 4 (Eastern Europe); C o l l i n 1966 (revision);
Munari & Baez 2000 (Atlantic islands); Munari 8t M e r z 2002 (Mediterranean species);
Soos 1981 (Central Europe); Stackelberg i g 8 9 a u (former U S S R ) ; C M P D : Munari 1998;
C A T : Soos 1 9 8 4 m ; Munari 2002.

Thaumaleidae (key couplet 15; fig. 47-48)

Systematics: Nematocera; superfamily C h i r o n o m o i d e a ; 3 g e n
era and about 75 species in Europe.
Characters: Small (3-5 m m ) , delicate, y e l l o w to orange or dark
b r o w n , lustrous Nematocera; u p p e r side o f thorax often w i t h
a U - s h a p e d colour pattern. Eyes holoptic in b o t h sexes; ocelli
absent; antenna short and directed forward, w i t h 12 segments,
tapering to its tip. W i n g clear, w i t h b r o w n tinge and relatively
broad. Legs short and slender, tibiae w i t h o u t apical bristles or
spurs.
Biology: T h e larvae and pupae are found in unpolluted aquatic
environments, usually in shaded, hilly or mountain areas. T h e lar
vae live in the water film of streams and splash zones o f springs,
banks, rocks and other objects protruding from the water, holding
to the substratum w i t h thoracic and abdominal pseudopodia.
They feed o n vegetable debris and diatoms. T h e adults are clumsy 574 Thaumalea testacea Ruthe, male;
fliers and are generally found near the larval habitats. Seguy1951
Identification references: D i s n e y 1 9 9 9 (British Isles);
S t a c k e l b e r g 19896 (former U S S R ) ; W a g n e r lggji, 2002
(species); C M P D : W a g n e r 1997c; C A T : M a r t i n o v s k y &
R o z k o s n y 1988.

167
Therevidae (key couplet 60; fig. 100,127)
Systematics: L o w e r m u s c o m o r p h B r a c h y c e r a ; s u p e r f a m i l y
A s i l o i d e a ; s o m e 17 genera and about 100 species in Europe.
Characters: Slender to m o r e or less stout, small to large (2.5-15
m m ) flies, often strongly p u b e s c e n t w i t h stronger bristles o n
thorax and legs; colour v a r y i n g from pale y e l l o w to black.
Frons o f female i n m a n y species w i t h g l o s s y black markings;
eyes u s u a l l y holoptic in the male; antenna variable, first or
third s e g m e n t often the longest; the latter f o l l o w e d b y n o , or 1-
2 smaller s e g m e n t s and a small apical style. W i n g clear or
tinged, in s o m e cases w i t h markings or bands; cell m 3 strong
ly n a r r o w i n g t o w a r d h i n d margin, or closed; cell c u p closed.
Legs relatively l o n g and slender; tibiae w i t h apical bristles;
e m p o d i u m bristle-like or absent.
575 Thereva nobilitata (Fabricius),
Biology: T h e larvae live in sandy soil, u n d e r n e a t h tree bark and female; Toth 1977
in dead w o o d , w h e r e t h e y prey o n a range o f small inverte
brates, i n particular beetle larvae. T h e adults are found in various habitats b u t prefer o p e n ,
s u n n y places; in v e r y d r y areas t h e y are often found near water. T h e y are able fliers b u t
often sit o n the g r o u n d or o n rocks, leaves, branches, tree trunks, etc. Contrary to m a n y
other l o w e r Brachycera the adults are n o t predatory b u t feed o n nectar, plant or animal
juices and other organic matter.
Identification references: V a n der G 0 0 U 9 8 5 ( N o r t h w e s t e r n Europe); Haarto 8t W i n q v i s t
2006 (Finland); O l d r o y d 1969; S t u b b s & Drake 2001 (British Isles); V F . Z a i t z e v 1989a
(former U S S R ) ; C M P D : Majer 1997b; C A T : Lyneborg 1989.

(Thyreophoridae): part o f the Piophilidae.

Tipulidae 2 2 2
(key couplet 6 , 1 9 5 ; fig. 19, 22-23, 5> 9> 3 3 . 4 5 ~ 4 7 ) 2

Systematics: N e m a t o c e r a ; superfamily T i p u l o i d e a ; traditional

ly this family includes the C y l i n d r o t o m i d a e , Limoniidae and
Pediciidae; 10 genera and about 4 7 0 species in Europe.
Characters: M e d i u m sized to v e r y large (7-35 m m ) N e m a t o c e r a
w i t h l o n g and slender antennae, w i n g s , legs and a b d o m e n .
Tipulidae include s o m e o f the largest Diptera. R o s t r u m w e l l -
d e v e l o p e d and often w i t h nasus; ocelli absent; antenna usual
ly consisting o f 13 s e g m e n t s , in s o m e cases more. T h o r a x w i t h
a V - s h a p e d transverse suture. W i n g w i t h 2 anal v e i n s and in
s o m e species w i t h c o n s p i c u o u s markings; in a f e w species
female w i t h w i n g s short or reduced.
Biology: T h e larvae are rarely aquatic, generally semi-aquatic to
terrestrial; s o m e species live in m o s s e s , l i v e r w o r t s , or i n d y i n g
and d e c a y i n g w o o d o f d e c i d u o u s trees. T h e larvae feed m a i n l y
o n plant remains and detritus b u t in a n u m b e r of species l i v i n g
576 Tipula trifascingulata Theowald;
parts o f m o s s e s , l i v e r w o r t s and seed plants are eaten as w e l l ;
female; Seguy 1951
the larvae o f Tipula, subgenus Tipula and some Nephrotoma
species feed o n pasture grasses, tree saplings, y o u n g cultivated
plants, etc., and can cause serious d a m a g e t o agriculture as t h e y are often present in large
n u m b e r s . Tipulidae larvae are an i m p o r t a n t food source especially for birds. T h e adults
occur in a w i d e range o f habitats v a r y i n g from dark m o i s t forests to d r y semi-deserts or
h i g h alpine regions. T h e y do n o t feed; at m o s t , t h e y rarely drink from d e w - d r o p s or take
s o m e nectar from flowers and similar liquids.
Identification references: H o f s v a n g i g 9 7 (genera); M a n n h e i m s 1951-1968; O o s t e r b r o e k e t
al 2006 (Ctenophorinae); T h e o w a l d 1973-1980 (Lindner); S a v c h e n k o 1989b (former
U S S R ) ; S t u b b s 1972-1996 (British Isles); C M P D : n o contribution; C A T : Oosterbroek &
T h e o w a l d 1992; Oosterbroek 2006.

(Trepidariidae): n a m e u s e d i n the past for Micropezidae: Calobatinae.

168
Trichoceridae (key couplet 4 ; fig. 27-28)
Systematics: Nematocera; superfamily Trichoceroidea; 2 g e n -
era, Diazosma and Trichocera, and about 50 species in Europe.
Characters: Small to m e d i u m sized (3-9 m m ) , generally light
coloured Nematocera, w i t h l o n g slender antennae, w i n g s , legs
and a b d o m e n . Ocelli present; antenna w i t h 18 s e g m e n t s .
W i n g w i t h a short, c u r v e d v e i n A 2 , the l o n g e s t in Diazosma.
Tibiae w i t h terminal bristles or spurs. Female o f Trichocera
w i t h a m o r e or less d o w n w a r d c u r v i n g ovipositor.
Biology: T h e larvae are terrestrial and saprophagous, living in
moist earth, rotting vegetable matter, under bark o f fallen
c o n i f e r o u s trees, m u s h r o o m s a n d bracket f u n g i , carrion
(including h u m a n corpses), excrement, etc. T h e adults o f
Trichocera are active from a u t u m n to spring, t h r o u g h o u t the
w i n t e r ("winter gnats"), w i t h especially the females found
w a l k i n g o n snow. A d u l t s o f Diazosma occur m a i n l y in s u m -
mer and early a u t u m n . T h e males have a characteristic b o u n c -
ing flight and often form s w a r m s , i n particular o n s u n n y days.
Identification references: D a h l 1 9 6 6 , 1 9 6 7 ( S w e d e n ) ; 577 Trichocera maculipennis Meigen,
Stackelberg 1989a (former U S S R ) ; Stary 8t M a r t i n o v s k y 1993 male; Zilahi-Sebess 1960
(Diazosma); C M P D : D a h l 8c K r z e m i n s k a 1997; C A T : D a h l
1992.

(Trichomyiidae): part o f the Psychodidae.

Trixoscelididae (key couplet 93; fig. 192,205-206)

Systematics: Acalyptrate Brachycera; superfamily
Sphaeroceroidea; traditionally part o f the H e l e o m y z i d a e ; in
Europe 1 genus, Trixoscelis, w i t h about 25 species.
Characters: Small (2-4 m m ) , m o s t l y grey flies. Arista w i t h
microscopic hairs to short p l u m o s e ; ocelli present; Oc-bristles
present, situated n e x t to the ocellar triangle at the same level
as the anterior ocellus or l o w e r ; P-bristles converging; 2 pairs
of F-bristles, c u r v i n g b a c k w a r d ; interfrontal setulae absent;
vibrissae present. W i n g usually patterned or partly tinged, in
only a f e w species w i n g clear; costa w i t h a subcostal break;
v e i n Sc complete b u t apical part thin and especially near costa
close to v e i n R i ; crossvein B M - C u present; cell c u p closed.
Tibiae w i t h dorsal preapical bristle
Biology: T h e larvae o f Trixoscelis are u n k n o w n ; s o m e species
have b e e n bred from nests o f birds. A d u l t s are t h e r m o p h i l o u s 578 Trixoscelis frontalis (Fallen), male;
Papp 1988
and prefer o p e n w e l l insolated habitats, like d u n e s , beaches,
heather, sandy steppes and forest steppes.
Identification references: H a c k m a n 1 9 7 0 ( E u r o p e ) ; S o o s 1981 ( C e n t r a l E u r o p e ) ;
Stackelberg 1989am (former U S S R ) ; C M P D : Papp 1998!; C A T : S o o s 1984J.

(Trypetidae): n a m e used in the past for Tephritidae.

(Tylidae): n a m e used in the past for Micropezidae.

169
UUdiidae (key couplet 1 3 9 , 1 4 1 , 1 6 1 , 1 6 3 ; fig. 301, 3 4 9 -
351. 356-357)
\
Systematics: A c a l y p t r a t e B r a c h y c e r a ; s u p e r f a m i l y
T e p h r i t o i d e a ; f o l l o w i n g r e c e n t classifications t h e
Otitinae are treated here as a subfamily; traditional
classifications d i s t i n g u i s h b e t w e e n U l i d i i d a e and
Otitidae as t w o families or include t h e Ulidiinae as a
subfamily o f the Otitidae; s o m e 17 genera and about
105 species in Europe.
Characters: Small to large (2.5-11 m m ) , m o d e r a t e l y
robust flies, colour v a r y i n g from partially y e l l o w and
dark to all black, in s o m e cases grey or metallic. A r i s t a
bare to l o n g pubescent; ocelli present; in a f e w species
all bristles o n t h e head are reduced or absent, b u t the
usual bristle configuration is: Oc-bristles present, P-
bristles parallel to diverging, 1-2 pairs o f F-bristles, 579 Physiphora alceae (Preyssler),
male; Stackelberg 1956
scattered interfrontal setulae present, and vibrissae
absent. W i n g u s u a l l y w i t h a p a t t e r n o f spots or
stripes or w i t h a spot at the w i n g tip, rarely all clear; costa w i t h or w i t h o u t a subcostal
a n d / o r a humeral break; v e i n Sc c o m p l e t e ; crossvein B M - C u present; cell c u p closed.
Tibiae usually w i t h o u t dorsal preapical bristle. Female w i t h oviscape, n o n retractable
basal s e g m e n t o f the ovipositor.
Biology: T h e b i o l o g y o f m o s t species is little k n o w n . Larvae are found in rotting vegetable
matter, leaf litter, fruit, d u n g , sap e x u d i n g from tree w o u n d s and u n d e r n e a t h tree bark;
s o m e species are fully p h y t o p h a g o u s , feeding o n sugar beet, o n i o n s , m a i z e , etc. A d u l t s are
found in various habitats ranging from saline b i o t o p e s to sandy areas or m o i s t and marshy
g r o u n d s , o n flowers, tree trunks or o n excrement.
Identification references: C l e m e n t s 1 9 9 0 (British Isles); Kabos & V a n A a r t s e n 1 9 8 4
(Netherlands); M e r z 1996a, 1996c (Switzerland); S o o s 1980 (Central Europe); Richter
i 9 8 g d , 19896 (former U S S R ) ; C M P D : G r e v e 1998 (Otitinae o n l y ) ; C A T : V.F. Z a i t z e v 1984
(Ulidiinae); S o o s i g 8 4 f (Otitinae).

(Usiidae): part o f the B o m b y l i i d a e .

Vermileonidae (key c o u p l e t 4 i , 52; fig.

87,113-114)
Systematics: L o w e r Brachycera; super-
family Tabanoidea; in Europe 9 species
in 2 genera, Lampromyia (5) a n d
Vermileo (4).
Characters: M e d i u m sized to large (5-18
m m ) , slender, pale to dark b r o w n flies,
d e v o i d o f bristles. Eyes dichoptic; api
cal s e g m e n t s o f the antenna tapering,
m a k i n g u p a style; m o u t h p a r t s c o n
s p i c u o u s l y elongate in Lampromyia,
short in Vermileo. W i n g l o n g and slen
der, w i t h o u t alula; clear, tinged or w i t h 580 Vermileonld larva digging a funnel-trap in dry sand;
markings along t h e v e i n s or across the Wheeler 1931
w i n g ; cell m 3 and cell c u p b o t h either
o p e n or closed. Legs l o n g , tibiae w i t h apical bristles or spurs; e m p o d i u m pulvilliform, in
Lampromyia pulvilli a n d / o r e m p o d i u m s o m e t i m e s extremely reduced.
Biology: T h e larvae o f the V e r m i l e o n i d a e prey o n small insects and other invertebrates,
u s i n g a funnel trap they d i g o u t in d r y and sandy areas, similar to the traps o f ant lions
(Neuroptera: M y r m e l e o n t i d a e ) . T h e adults visit flowers.
Identification references: N a g a t o m i e t al. 1 9 9 9 ( W o r l d genera); S t u c k e n b e r g 1965
(Vermileo), 1998 (Lampromyia); C M P D : N a g a t o m i 1997b; C A T : Majer I988f.

170
Xenasteiidae (key couplet 101; fig. 226-227)
Systematics: Acalyptrate Brachycera; superfamily
O p o m y z o i d e a ; in Southern Europe 2 species, Tunisimyia con-
vergens V e n t u r a & C a r l e s - T o l r a (Balearic Islands) a n d
Xenasteia excellens (Papp) (Italy, Malta).
Characters: M i n u t e (1.3-1.8 m m ) , b r o w n t o black lustrous flies;
legs and parts o f the head y e l l o w i s h . A r i s t a m i c r o p u b e s c e n t ;
ocelli present; Oc-bristles present; small, thin p s e u d o p o s t o -
cellar bristles present, c u r v i n g forward or c o n v e r g i n g ; 3 pairs
of F-bristles, t h e l o w e r pair c u r v i n g i n w a r d , the others back
w a r d or the upper pair obliquely b a c k w a r d ; interfrontal bris
tles present; vibrissae present. W i n g u n m a r k e d ; alula w i t h
long hair-like bristles; v e n a t i o n reduced; costa w i t h h u m e r a l
and subcostal breaks; v e i n Sc c o m p l e t e u p to the costa, b u t api
cal part pallid; v e i n M i largely pallid, n o t reaching the w i n g
margin; crossveins B M - C u and D M - C u b o t h absent; cell c u p
closed; anal v e i n absent. Tibiae w i t h o u t dorsal preapical bris- 581 Xenasteia excellans (Papp),
male; Papp 1980
de.
Biology: T h e larvae o f this small family are u n k n o w n . T h e
adults are found in saline habitats, b o t h coastal and inland and in deserts. T h e y are found
on f l o w e r i n g plants and trees.
Identification references: V a n i n 2003 (Italy); Ventura 8t Carles-Tolra 2002 (Balearic
Islands); C M P D : Papp 1 9 9 8 ^ C A T : Papp 1984I.

Xylomyidae (key c o u p l e t 4 i , 53; fig. 86,115)

Systematics: L o w e r Brachycera; superfamily Stratiomyoidea;
in Europe 8 species in 2 genera, Solva (5) and Xylomya (3).
Characters: M e d i u m sized to large (6-20 m m ) , slender flies,
w i t h o u t bristles o n thorax and a b d o m e n . B o d y dark, w i t h l i m
ited to extensive lighter markings. Eyes dichoptic; antenna
w i t h acute tip, w i t h 10 segments. W i n g clear or tinged; b o t h
the cell m3 and cell c u p closed. Tibia o f m i d and h i n d leg w i t h
apical bristles or spurs; e m p o d i u m p u l v i l l i f o r m .
Biology: Larvae, often several together, l i v i n g u n d e r n e a t h the
bark of d e c i d u o u s trees, feeding o n injured and dead insect lar
vae and other small invertebrates. T h e adults are m a i n l y found
near the larval habitats, often sitting o n trees.
Identification references: B r u g g e 2002 ( N e t h e r l a n d s ) ;
Krivosheina 1999a, 1999b; Nartshuk 19896 (former U S S R ) ;
O l d r o y d 1969; Stubbs 8t Drake 2001 (British Isles); R o z k o s n y 582 Xylomya maculata (Meigen),
1973 ( F e n n o s c a n d i a a n d D e n m a r k ) ; C M P D : N a g a t o m i & male; Sguy 1926
R o z k o s n y 1997a; C A T : K r i v o s h e i n a 1988b.

171
Xylophagidae (key couplet 4 3 , 5 2 ; fig. 90,112)
Systematics: L o w e r Brachycera; superfamily X y l o p h a g o i d e a ; in
Europe 1 genus, Xylophagus, w i t h 5 species.
Characters: M e d i u m sized t o large (5-11 m m ) flies, largely
devoid o f bristles, w i t h long, slender w i n g s , legs and
a b d o m e n . B o d y usually dark w i t h y e l l o w markings, in s o m e
cases largely y e l l o w . Eyes dichoptic; antenna simple, w i t h 12,
rarely 11 segments. W i n g w i t h o u t alula and w i t h markings
along t h e v e i n s or across the w i n g ; cell m3 o p e n ; cell c u p o p e n
or closed. Legs in s o m e cases banded; tibiae w i t h apical bristles
or spurs; e m p o d i u m p u l v i l l i f o r m .
Biology: T h e larvae live u n d e r n e a t h tree bark, in rot holes o f
trees and in decaying d e c i d u o u s and coniferous trees w h e r e
they are t h o u g h t to prey o n other invertebrates, i n c l u d i n g
their o w n species. A d u l t s are m a i n l y found near t h e larval
habitats, especially i n m o r e rich and h u m i d forests, often sit
583 Xylophagus ater Melgen,
ting o n tree trunks.
male; Verrall 1909
Identification references: V a n der G o o t 1985 ( N o r t h w e s t e r n
Europe); Haenni lggzd (Switzerland); K r i v o s h e i n a 8t M a m a e v
1972 (species); Nartshuk i g 8 9 d (former U S S R ) ; O l d r o y d 1969; Stubbs 8t Drake 2001
(British Isles); C M P D : N a g a t o m i 8t R o z k o s n y 1997b; C A T : Krivosheina & M a m a e v
i988d.

172
CHECKLISTS OF EUROPEAN
Andorra: Carles-Tolr Hjorth-Andersen, M.
(ed.), 2002. Catalogo de los Dptera de
Espaa, Portugal y Andorra. - Monografas
Sociedad Entomolgica Aragonesa 8:1-323.
Belgium: Grootaert, P., Bruyn, L. de & Meyer,
M. de (eds.), 1992. Catalogue of the Dptera
of Belgium. - Studiedocumenten van het
K.B I.N. 70:1-338.
Bulgaria: Lavciev, V., 2003. Dptera: Fanniidae,
Muscidae, Stomoxydidae. - Catalogus
Faunae Bulgaricae 5:1-77 (the other volumes
in this series are not about Dptera).
Czech Republic: Chvla, M. (ed.), 1997.
Checklist of Dptera (Insecta) of the Czech
and Slovak Republics. Karolinum-Charles
University Press, Prague: 1-130.
Denmark: Petersen, F.T. 8t Meier, R. (eds.),
2001. A preliminary list of the Dptera of
Denmark. - Steenstrupia 26:119-276.
Finland: Hackmann, W., 1980. A check list of
the Finnish Dptera. - Notulae
Entomologicae 60:17-48 (Part I.
Nematocera and Brachycera (s. str.)); 117-162
(Part II. Cyclorrhapha).
Germany: Schumann, H., Bhrmann, R. 8t
Stark, A . (eds.), 1999. Checkliste der
Dipteren Deutschlands. - Studia
Dipterologica, Supplement 2: - 3 5 4 .
(Additions in Studia Dipterologica 9 (2003):
437-445, and 11 (2005): 619-630.)
Great Britain: Chandler, P.J., (ed.), 1998.
Checklists of insects of the British Isles
(New Series) part 1: Diptera. - Handbooks
for the Identification of British Insects 12:
i-vx, 1-234. (Additions and corrections in
subsequent issues of Dipterists Digest.)
COUNTRIES
Hungary: Papp, L. (ed.), 2001. Checklist of the
Diptera of Hungary. Hungarian Natural
History Museum, Budapest: 1-550.
Ireland: see Great Britain.
Italy: Minelli, A., Ruffo, S. 8t Posta, S. la (eds.),
1995. Checklist delle specie della fauna
Italiana, vols. 63-78 (each volume w i t h sep-
arate page numbers).
Lithuania: Pakalniskis, S., Rimsaite, J.,
Spragnauskaite-Bernotiene, R., Butautaite,
R. 8t Podenas, S., 2000. Checklist of
Lithuanian Diptera. Acta Zoologica
Lituanicaio(i): 3-57. (Additions: 2000:
Acta Zoologica Lituanica 10 (3): 20-26).
Netherlands: Beuk, P.L.T. (ed.), 2002.
Checklist of the Diptera of the Netherlands.
K N N V Uitgeverij, Utrecht: 1-448
Poland: Razowski, J. (ed.), 1991. Checklist of
animals of Poland, Vol. 2, Polska Akademia
Nauk: 1 - 342 (Diptera 77-269).
Portugal: see Andorra
Slovakia: see Czech Republic
Spain: see Andorra.
Switzerland: Merz, B., Bachli, G, Haenni, J.- P.
8t Gonseth, Y. (eds.), 1998. Diptera -
Checklist Fauna Helvetica 1:1-369.
(Additions: 2002: Mitteilungen der
Entomologische Gesellschaft Basel 51:110-
140.)
For a systematic survey of all European
animals at family, genus, (sub)species
and/or country level, including distribu-
tion maps for the (sub) species, see the
Fauna European Web Service website at
http://www.faunaeur.org
173
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Krivosheina, N.P., 1986c. Family Anisopodidae. - Same as
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Krivosheina, N.P., 1988a. Family Diadociidae. - In: Sos, A.
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Krivosheina, N.P, i997d. Family Mycetobiidae. - Same as
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Majer, J , i988d. Family Hilarimorphidae. - Same as 1988a:
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Majer, J , 1988e. Family Coenomyiidae. - Same as 1988a: 31-
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Mamaev, B.M, 1989c. Family Axymyiidae. - Same as 1989a:
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Sos, A, 1984b. Family Tanypezidae. - Same as 1984a: 26-
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Sos, , 1984c. Family Psilidae. - Same as 1984a: 28-35.
Sos, , 1984a. Family Pyrgotidae. - Same as 1984a: 36-38.
Sos, A , 1984e. Family Platystomatidae. - Same as 1984a:
38-45.
Sos, A , 1984! Family Otitidae. - Same as 1984a: 45-59.
Sos, , 1984g. Family Dryomyzidae. - Same as 1984a: 152-
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Sos, A , 1984b. Family Neottiophilidae. - Same as 1984a:
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Sos, , 1984L Family Clusiidae. - In: Sos, . 8c Papp, L.
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Sos, A, 1984J. Family Trixoscelididae. - Same as 1984!: 45-
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Sos, , 1984k. Family Opomyzidae. - Same as 1984!: 53-
6
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Sos, , 1984I. Family Chyromyidae. - Same as 1984!: 56-
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Sos, , 1984m. Family Tethinidae. - Same as 1984!: 107-
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Sos, . 8c Hrka, K, 1986. Family Hippoboscidae. - In:
Sos, . 8c Papp, L. (eds), Catalogue of Palaearctic
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Sos, . 8c Minr, J , 1986a. Family Gasterophilidae. - In:
Sos, A. 8c Papp, L. (eds), Catalogue of Palaearctic
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Sos, A. 8c Minf, J , 1986b. Family Oestridae. - Same as
1986a: 240-244.
Sos, . 8c Minr, J , 1986c. Family Hypodermatidae. -
Same as 1986a: 244-251.
Sos, A. 8c Oosterbroek, P., 1992. Family Cylindrotomidae.
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Spencer, K.A, 1976. The Agromyzidae (Diptera) of
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Spencer, K.A, 1987. Agromyzidae. - In: McAlpine, J.F. et al.
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the USSR. - Opredeliteli po fauna SSSR 60:1-164. (In
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Stackelberg, A.A, 1989a. Family Trichoceridae. - In: Bei-
Bienko, G.Y. 8c Steyskal, G.C. (eds), Keys to the insects
of the European Part of the USSR. Volume V. Diptera
and Siphonaptera, Part 1. Brill, Leiden, etc.: 68-71 (origi-
nally published in Russian in 1969).
Stackelberg, A.A, 1989b. Family Ptychopteridae. - Same as
1989a: 164-166.
Stackelberg, A.A, 1989c. Family Dixidae. - Same as 1989a:
209-215.
Stackelberg, A.A, I98gd. Family Chaoboridae. - Same as
1989a: 216-217.
Stackelberg, A.A, 1989e. Family Thaumaleidae. - Same as
189
 1989a:370-372-
Stackelberg, A.A., 1989^ Family Pachyneuridae. - Same as
1989a: 373
Stackelberg, A.A., i989g. Family Bolitophilidae. - Same as
1989a: 374-385-
Stackelberg, A.A., 1989b. Family Ditomyiidae. - Same as
1989a: 386-388.
Stackelberg, A.A., 19891. Family Ceroplatidae
(Keroplatidae). - Same as 1989a: 389-394.
Stackelberg, A.A., 1989). Family Diadocidiidae. - Same as
1989a: 395.
Stackelberg, A.A., 1989k. Family Macroceridae. - Same as
1989a: 396-400.
Stackelberg, A.A., 1989I. Family Manotidae. - Same as
19893:401.
Stackelberg, A.A., 1989m. Family Hesperinidae. - Same as
1989a: 666.
Stackelberg, A.A., ig8gn. Family Anisopodidae. - Same as
1989a: 681-682.
Stackelberg, A.A., 19890. Family Lonchopteridae. - Same as
1989a: 1153-1155.
Stackelberg, A.A., 1989P. Family Syrphidae. - In: Bei-
Bienko, G.Y. 8c Steyskal, G.C. (eds), Keys to the insects
of the European Part of the USSR. Volume V. Diptera
and Siphonaptera, Part 2. Brill, Leiden, etc.: 10-148
(originally published in Russian in 1970).
Stackelberg, A.A., 19891. Family Calobatidae
(Trepidariidae). - Same as 1989P: 179.
Stackelberg, A.A., 1989s. Family Micropezidae (Tylidae, in
part). - Same as 1989P: 176-178.
Stackelberg, A.A., 19891. Family Megamerinidae. - Same as
1989P: 180-181.
Stackelberg, A.A., 1989U. Family Tanypezidae. - Same as
1989P: 182.
Stackelberg, A.A., 1989V. Family Strongylophthalmyiidae. -
Same as 1989P: 183.
Stackelberg, A.A., 1989W. Family Psilidae. - Same as 1989P:
184-191.
Stackelberg, A.A., 1989X. Family Pyrgotidae. - Same as
1989P'. 192-193.
Stackelberg, A.A., 1989^ Family Helcomyzidae. - Same as
1989P: 277.
Stackelberg, A.A., 1989Z. Family Coelopidae. - Same as
1989P: 278-279.
Stackelberg, A.A., 198933. Family Dryomyzidae. - Same as
1989P: 280-281.
Stackelberg, A.A., 19893b. Fsmily Sepsidae. - Same as
1989P: 282-289.
Stackelberg, A.A., 19893c. Family Sciomyzidae. - Same as
ig8gp: 290-317.
Stackelberg, A.A., 19893d. Family Lauxaniidae. - Same as
1989P: 318-330.
Stackelberg, A.A., 198936. Family Periscelididae. - Same as
1989P: 346.
Stackelberg, A.A., ig89af. Family Piophilidae. - Same as
8 :
9 9P 347-35-
Stackelberg, A.A., I9893g. Family Neottiophilidae. - Same
as 1989P: 352-353-
Stackelberg, A.A., i98gah. Family Pallopteridae. - Same as
1989P: 354-357-
Stsckelberg, A.A., ig8gai. Family Lonchaeidae. - Same as
i98gp: 358-373.
Stackelberg, A.A., 1989a). Family Odiniidae. - Same as
1989P: 374-37-
Stsckelberg, A.A., ig8gak. Family Clusiidae. - Same as
1989P: 505-508.
Stackelberg, A.A., ig8gal. Family Acartophthalmidae. -
Same as ig8gp: 5og.
190
Stackelberg, A.A., ig8gam. Fsmily Trixoscelididae. - Same
as 1989P: 538.
Stackelberg, A.A., ig8gan. Family Pseudopomyzidae. -
Sameasi989p: 539.
Stackelberg, A.A., ig8gao. Family Anthomyzidae. - Same as
1989P: 540-545-
Stsckelberg, A.A., ig8gap. Family Opomyzidae. - Same 3S
198gp: 546-550.
Stackelberg, A.A., I98gaq. Family Chyromyidae. - Same as
I98gp: 551-552.
Stackelberg, A.A., I98gar. Family Aulacigasttidae. - Same as
I98gp: 553-554-
Stackelberg, A.A., 19893s. Family Asteiidae. - Same as
199P: 555-556-
Stackelberg, A.A., 198931. Family Cryptochaetidae. - Same
as ig8gp: 557-556.
Stackelberg, A. A., 19893U. Family Tethinidae. - Same as
I98gp: 591-592-
Stackelberg, A. A., 19893V. Family Milichiidae. - Same as
I98gp: 593-601.
Stackelberg, A. A., ig8gaw. Family Canaceidae. - Same as
1989P: 602-603.
Stackelberg, A.A., ig8gax. Family Braulidae. - Same as
ig8gp: 604.
Stackelberg, A.A., I98gay. Family Camillidae. - Same as
ig8gp: 647.
Stackelberg, A.A., ig8gaz. Family Diastatidae. - Same as
i98gp: 648-650.
Stackelberg, A.A., i98gba. Family Drosophilidae. - Same as
1989P: 651-665.
Stackelberg, A.A., ig8gbb. Family Curtonotidae. - Same as
ig8gp: 666.
Stackelberg, A.A., ig8gbc. Family Eginiidae. - Same as
1989P: 975.
Stackelberg, A.A., ig8gbd. Family Streblidae. - Same as
1989P: 987-988.
Stackelberg, A.A., ig8gbe. Family Nycteribiidae. - Same as
ig8gp: g8g-994.
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193
FIGURE DETAILS
2a habitus Tabanus americanus (Tabanidae);
McAlpine 1981a
2b habitus Brachystoma vesiculosum
(Brachystomatidae), male; Niesiolowski 1992
3 habitus brachycerous fly; Lane 8c Crosskey 1993
4 head calyptrate fly; Lane 8c Crosskey 1993
5 head calyptrate fly; Lane 8c Crosskey 1993
6a antenna Aedes alternans (Culicidae), female;
Colless 8c McAlpine 1991
6b antenna Tabanidae; Merz 8c Haenni 2000 (draw
ing by B. Merz)
6c antenna calyptrate fly; Lane 8c Crosskey 1993
7 head calyptrate fly; Lane 8c Crosskey 1993
8 head calyptrate fly; Colless 8c McAlpine 1991
9 head Paracantha gentilis (Tephritidae); Hennig 1973
10 head Anorostoma currani (Heleomyzidae);
Hennig 1973
11 thorax Tachinidae; Colless 8c McAlpine 1991
12 thorax Calliphoridae; Lane 8c Crosskey 1993
13 hypothetical ground plan of the Diptera wing;
Lane 8c Crosskey 1993
14 wing Calliphora vicina (Calliphoridae); Lane 8c
Crosskey 1993
15 elements of the Diptera leg; Oosterbroek 1981
16 bristle (macrotrichium) with socket (alveolus);
Merz 8c Haenni 2000 (drawing by B. Merz)
17 antenna Aedes alternans (Culicidae), female;
Colless 8c McAlpine 1991
18 antenna Aedes alternans (Culicidae), male; Colless
8c McAlpine 1991
19 antenna Dictenidia bimaculata (Tipulidae), male;
Mannheims 1951
20 antenna Tabanidae; Merz 8c Haenni 2000 (draw
ing by B. Merz)
21 head calyptrate fly; Lane 8c Crosskey 1993
22 head Tipula trivittata (Tipulidae); McAlpine 1981a
23 habitus Tipula luna (Tipulidae); Lindner 1948
24 wing Mycetobia pallipes (Mycetobiidae);
Krivosheina i997d (drawing by N.P. Krivosheina)
25 thorax Tipulidae; Hendel 1928
26 thorax Cylindrotoma distinctissima
(Cylindrotomidae) ; Peus 1952
27 wing Trichocera saltator (Trichoceridae); Dahl 8c
Krzeminska 1997
28 wing Diazosma tibeticum (Trichoceridae); Dahl
1957
29 head Prionocera turcica (Tipulidae); Tjeder 1948
30 wing Tricyphona protea (Pediciidae); Alexander 8c
Byers 1981
31 habitus Elephantomyia edwardsi (Limoniidae);
Tjeder 1953
32 wing Elliptera tennessa (Limoniidae); Alexander 8c
Byers 1981
33 wing Tipula tricolor (Tipulidae); Alexander 8c
Byers 1981
34 habitus Cylindrotoma distinctissima
(Cylindrotomidae); Lindner 1948
35 wing Phalacrocera replicata (Cylindrotomidae);
Alexander 8c Byers 1981
36 hypopygium Triogma trisulcata
(Cylindrotomidae); Peus 1952
37 ovipositor Triogma trisulcata (Cylindrotomidae);
Peus 1952
194
38 wing Sylvicola fenestralis (Anisopodidae);
Peterson 1981
39 wing Mochlonyx velutinus (Chaoboridae); Saether
2002
4 0 wing Ptychoptera lacustris (Ptychopteridae); Peus
1958
41 haltere Ptychoptera contaminata (Ptychopteridae);
Oosterbroek 1981
42 habitus Tinearia alternata (Psychodidae); Wagner
1997a (drawing by A. Szappanos)
42a wing Trichomyia urbica (Psychodidae); Wagner
1997a
43 habitus Dixa maculata (Dixidae), female; Wagner
1997c (drawing by A. Szappanos)
4 4 antenna Dixella nova (Dixidae); Peters 1981
45 habitus Aedes vexans (Culicidae), female; Minar
2000a (drawing by A. Szappanos)
46 head Chaoborus plumicornis (Chaoboridae),
male; Oldroyd 1964
47 wing Thaumalea tricuspis (Thaumaleidae); Tjeder
1949
48 head Thaumalea brevidens (Thaumaleidae) ;
Wagner 2002
49 habitus Cecidomyia pini (Cecidomyiidae), male;
Skuhrav 1997 (drawing by M. Skuhrav)
50 wing Greniera spec. (Simuliidae); Peterson 1981
51 head Twinnia spec. (Simuliidae); Peterson 1981
52 wing Chironomidae; Oosterbroek 1998
53 wing Culicoides antennalis (Ceratopogonidae);
Colless 8c McAlpine 1991
54 wing Liponeura bilobata (Blephariceridae) ;
Hennig 1973
55 head Blepharicera tetrophthalma
(Blephariceridae), female; Hennig 1973
56 wing Protaxymyia melanoptera (Axymyiidae);
Krivosheina 2000 (drawing by N.P. Krivosheina)
57 wing Groveriella carpathica (Cecidomyiidae);
Skuhrav 1997 (drawing by M. Skuhrav)
58 head Swammerdamella acuta (Scatopsidae);
Haenni 1997a (drawing by A. Szappanos)
59 habitus Scatopse notata (Scatopsidae), male;
Colless 8c McAlpine 1991
60 wing Synneuron decipiens (Synneuridae);
Peterson 8c Cook 1981
61 wing Hyperoscelis eximia (Canthyloscelidae);
Haenni 1997b (drawing by S. Podenas)
62 hind leg Hyperoscelis eximia (Canthyloscelidae);
Haenni 1997b (drawing by S. Podenas)
63 habitus Bibio marci (Bibionidae), male; Skartveit
1997 (drawing by A. Szappanos)
64 tibia fore leg Dilophus femoratus (Bibionidae);
Zilahi-Sebess i960
65 head Hesperinus brevifrons (Hesperinidae), male;
Hardy 1981
66 wing Hesperinus brevifrons (Hesperinidae), male;
Hardy 1981
67 wing Plecia americana (Pleciidae), male; Hardy
1981
68 habitus Penthetria funebris (Pleciidae), male;
Sguy1940
69 head Pnyxia scabiei (Sciaridae); Steffan 1981
70 wing Pnyxia scabiei (Sciaridae); Menzel 8c Mohrig
1997 (drawing by F. Menzel)
71 wing Pachynema fasciata (Pachyneuridae);
Krivosheina 1997a (drawing by N.P. Krivosheina)
72 wing Mycetobia pallipes (Mycetobiidae);
Krivosheina I997d (drawing by N.P. Krivosheina)
73 head Sciara spec. (Sciaridae); Colless & McAlpine
1991
74 wing Zygoneura sciarina (Sciaridae); Menzel &
Mohrig 1997 (drawing by F. Menzel)
75 habitus Mycetophila propria (Mycetophilidae),
female; Colless 8c McAlpine 1991
76 wing Ditomyia fasciata (Ditomyiidae); Hendel
1928
77 wing Keroplatus spec. (Keroplatidae); Hendel 1928
78 habitus Keroplatus tipuloides (Keroplatidae),
male; Sguy 1940
79 wing Macrocera spec. (Keroplatidae); Hendel 1928
80 habitus Macrocera centralis (Keroplatidae),
female; Sguy 1940
81 wing Bolitophila spec (Bolitophilidae); Hendel
1928
82 wing Diadocidia spec. (Diadocidiidae); Hendel
1928
83 wing Sciarosoma borealis (Sciaroidea); Chandler
2002
84 wing Sylvicola fenestralis (Anisopodidae);
Peterson 1981
85 head Rachicerus obscuripennis (Rachiceridae),
female; James 1981
86 habitus Xylomya maculata (Xylomyidae), female;
Lindner 1938
87 wing Vermitigris orientalis (Vermileonidae);
Edwards 1932
88 habitus Beris clavipes (Stratiomyidae), female;
Rozkosny 1973
89 habitus Coenomyia ferruginea (Coenomyiidae),
male; Rozkosny 8c Nagatomi 1997a (drawing by J.
Pal)
89a spined scutellum Coenomyia ferruginea
(Coenomyiidae), male; Oosterbroek 1981
90 habitus Xylophagus ater (Xylophagidae), female;
Sguy 1926
91 head calyptrate fly; Sguy 1923
92 head Haematopota bigoti (Tabanidae), female;
Sguy 1926
93 head Stichopogon trifasciatus (Asilidae), female;
W o o d 1981b
9 4 head Arthroceras leptis (Rhagionidae), female;
James & Turner 1981
95 head Atherix variegata (Athericidae), female;
Webb 1981a
96 head Platypalpus infectus (Hybotidae); Collin
1961
97 head Hercostomus nobilitatus (Dolichopodidae);
Oldroyd 1970
98 habitus Ogcodes gibbosus (Acroceridae), male;
Sack 1936
99 habitus Cyrtus pusillus (Acroceridae), female;
Sguy 1926
100 wingThereva nobilitata (Therevidae), female;
Goot 1985
101 wing Empidideicus hungaricus
(Mythicomyiidae); Greathead 8c Evenhuis 1997
(drawing by L. Papp)
102 wing Oestrus ovis (Oestridae); Minar 1980
103 fifth tarsal segment Inopus rubriceps
(Stratiomyidae); Colless 8c McAlpine 1991
104 fifth tarsal segment Musca domestica (Muscidae);
Colless 8t McAlpine 1991
105 wing Ibisia marginata (Athericidae); Thomas
1997
106 antenna Atherix marginata (Athericidae); Verrall
1909
107 wing Fallenia fasciata (Nemestrinidae); Richter
1997a (drawing by V.A. Richter)
108 habitus Nemestrinus obscuripennis
(Nemestrinidae), female; Richter 1997a (drawing
by Florinskaya)
109 wing Trichopsidea costata (Nemestrinidae),
female; Sack 1933
n o a - d antenna: a: Heptatoma pellucens; b:
Haematopota pluvialis; c: H. italica; d: Tabanus
autumnalis (Tabanidae); Sguy 1926
111 habitus Tabanus autumnalis (Tabanidae), female;
Oldroyd 1970
112 habitus Xylophagus ater (Xylophagidae), female;
Sguy 1926
112a antenna Xylophagus ater (Xylophagidae), male;
Sguy 1926
113 habitus Vermileo degeeri (Vermileonidae), male;
Sguy 1926
114 habitus Lampromyia cylindrica (Vermileonidae),
male; Sguy 1926
115 habitus Xylomya maculata (Xylomyidae), female;
Lindner 1938
116 habitus Oxycera rara (Stratiomyidae), female;
Sguy 1950
117 habitus Coenomyia ferruginea (Coenomyiidae),
male; Rozkosny 8c Nagatomi 1997a (drawing by J.
Pal)
117a spined scutellum Coenomyia ferruginea
(Coenomyiidae), male; Oosterbroek 1981
118 habitus Rhagio scolopaceus (Rhagionidae),
female; Verrall 1909
119 habitus Chrysopilus splendidus (Rhagionidae),
male; Sguy 1926
120 wing Hispanomydas hispicus (Mydidae); Sack
1933
121 head Leptomydas padischach (Mydidae), male;
Richter 1997b (drawing by V.A. Richter)
122 wing Hilarimorpha ditissa (Hilarimorphidae);
Webb 1981b
123 head Dioctria engeli (Asilidae), male; Goot 1985
124 head Proagonistes athletes (Asilidae); Oldroyd
1964
125 head Scenopinus fenestralis (Scenopinidae),
female; Kelsey 1981
126 habitus Scenopinus fenestralis (Scenopinidae),
female; Tth 1977
127habitusThereva nobilitata (Therevidae), female;
Goot 1985
128 habitus Bombylella atra (Bombyliidae), male;
Tth 1977
129 wing Amphicosmus elegans (Bombyliidae); Hall
1981
130 habitus Apolysis andalusiaca (Bombyliidae),
female; Engel 1932
131 habitus Brachystoma vesiculosum
(Brachystomatidae), male; Niesiolowski 1992
132 wing Scenopinus unifasciatus (Scenopinidae);
Engel 1932
133 habitus Chelifera diversicauda (Empididae), male;
Niesiolowski 1992
134 habitus Clinocera dimidiata (Empididae), male;
Niesiolowski 1992
135 habitus Episyrphus balteatus (Syrphidae), male;
Thompson 8c Rotheray 1998 (drawing by J. Pal)
195
136 wing Erstalinus sepulchralis (Syrphidae); Verrall
1909
137 wing Psilota anthracina (Syrphidae); Verrall 1909
138 wing Conops flavipes (Conopidae); Smith 1969b
139 habitus Pipunculus campestris (Pipunculidae),
female; Kozanek et al. 1998 (drawing by J. Pal)
140 habitus Chalarus spurius (Pipunculidae), female;
Bankowska 1979
141 wing Empidideicus hungaricus
(Mythicomyiidae); Greathead 8c Evenhuis 1997
(drawing by L. Papp)
142 habitus Cyrtisiopsis melleus (Mythicomyiidae),
female; Engel 1932
143 habitus Myopa stigma (Conopidae), male;
Bankowska 1979
1 4 4 habitus Leopoldius calceatus (Conopidae), male;
Bankowska 1979
145 habitus Dalmannia punctata (Conopidae), male;
Bankowska 1979
146 wing Lonchoptera uniseta (Lonchopteridae),
male; Peterson 1987
147 wing Lonchoptera bifurcata (Lonchopteridae),
female; Peterson 1987
148 habitus Phoridae s p e c ; Hendel 1928
149 head Megaselia meconicera (Phoridae); Oldroyd
1964
150 wing Oplodontha viridula (Stratiomyidae);
Rozkosny 1982
151 head calyptrate fly; Seguy 1923
152 antenna calyptrate fly; Lane 8c Crosskey 1993
153 wing Oestrus ovis (Oestridae); Minaf 1980
154 head Platypalpus infectus (Hybotidae); Collin
1961
155 head Hercostomus nobilitatus (Dolichopodidae);
Oldroyd 1970
156 antenna Empis pennipes (Empididae); Chvala
1983
157 wing Opetia nigra (Opetiidae), male; Chandler
1998a (drawing by P. Chandler)
158 wing Opetia nigra (Opetiidae), female; Chandler
1998a (drawing by P. Chandler)
159 habitus Agathomyia falleni (Platypezidae), male;
Chandler 8c Shatalkin 1998 (drawing by P.
Chandler)
160 habitus Polyporivora picta (Platypezidae),
female; Chandler 2001 (drawing by P. Chandler)
161 habitus Microphor holosericeus (Microphoridae),
male; Chvala 1983
162 wing Sympycnus allectorius (Dolichopodidae);
Colless 8c McAlpine 1991
163 habitus Dolichopus popularis (Dolichopodidae),
male; Oldroyd 1970
164 habitus Sciapus basilicus (Dolichopodidae),
male; Meuffels & Grootaert 1990
165 wing Nemedina alamirabilis (Atelestidae);
Chandler 1981
166a habitus Meghyperus sudeticus (Atelestidae),
female; Chvala 1983
166b wing Atelestus pulicarius (Atelestidae); Chvala
1983
167 habitus Anthalia schoenherri (Hybotidae), male;
Chvala 1983
168 habitus Hybos culiciformis (Hybotidae), male;
Chvala 1983
169 habitus Drapetis assimilis (Hybotidae), male;
Collin 1961
170 front view thorax Empis pennipes (Empididae);
Chvala 1983
196
171 front view thorax Hybos grossipes (Hybotidae);
Chvla 1983
172 antenna Empis pennipes (Empididae); Chvla
1983
173 antenna Leptopeza borealis (Hybotidae); Chvla
1983
174a ventral view thorax Melophagus rupricaprinus
(Hippoboscidae); Theodor & Oldroyd 1964
174b habitus Ornithomya rupes (Hippoboscidae),
female; Biittiker 1994 (drawing by W . Buttiker)
175 head and fore legs Brachytarsina flavipennis
(Streblidae), male; Hrka 1998b (drawing by A.
Szappanos)
176 thorax calyptrate fly; Lane 8c Crosskey 1993
177 antenna calyptrate fly; Lane & Crosskey 1993
178 head Lispe tentaculata (Muscidae); Gregor et al.
2002
179 wing Calliphora vicina (Calliphoridae); Lane 8c
Crosskey 1993
180 upper side of thorax calyptrate fly; Oldroyd 1970
181 upper side of thorax acalyptrate fly; Oldroyd 1970
182 head Dasiops ensifer (Lonchaeidae), female;
Morge 1963
183 habitus Setisquamalonchaea fumosa
(Lonchaeidae); Sos 1980
184 habitus Limosina silvtica (Sphaeroceridae);
Mihlyi 1972
185 habitus Copromyza equina (Sphaeroceridae);
Rohcek et al. 2002
186 head Prosopomyia pallida (Lauxaniidae); Papp 1978
187 head Lauxania cylindricornis (Lauxaniidae);
Hendel 1928
188 head Coremacera marginata (Sciomyzidae);
Hendel 1928
189 head Clusiodes albimana (Clusiidae); Mihlyi
1972
190 habitus Clusiodes spec. (Clusiidae), male;
Sakawa 1998 (drawing by A. Szappanos)
191 wing base Camilla glabra (Camillidae); Hennig
1958
192 wing Trixoscelis marginella (Trixoscelididae);
Sos 1981
193 habitus Pachycerina seticornis (Lauxaniidae),
male; Papp 1978
194 head Orygma luctuosum (Sepsidae); Steyskal
1987
195 head Saltella sphondylii (Sepsidae); Steyskal 1987
196 posterior spiracle Orygma luctuosum (Sepsidae);
Oosterbroek 1981
197 posterior spiracle Nemopoda nitidula (Sepsidae);
Oosterbroek 1981
198 habitus Sepsis punctum (Sepsidae), female; Sos
1959
199 head Orygma luctuosum (Sepsidae); Pont 8c
Meier 2002
200 head Prosopomyia pallida (Lauxaniidae); Papp
1978
201 habitus Orbellia myopiformis (Heleomyzidae),
male; Sos 1981
202a head Coelopa frigida (Coelopidae); Hendel 1928
202b tarsus Coelopa frigida (Coelopidae);
Oosterbroek 1981
203 wing Malacomyia sciomyzina (Coelopidae);
McAlpine 1998b (drawing by A. Szappanos)
204 habitus Curtonotum helvum (Curtonotidae);
McAlpine I987d
205 head Trixoscelis marginella (Trixoscelididae);
Hennig 1958
206 wing Trixoscelis marginella (Trixoscelididae);
Sos 1981
207 wing Camilla glabra (Camillidae); H e n n i g i 9 5 8
208 wing Euthychaeta spectabilis (Campichoetidae);
Chandler 1987
209 head Camilla nigrifrons (Camillidae); Mihlyi
1972
210 femur fore leg Camilla flavicauda (Camillidae);
Beuk & de Jong 1994
211 habitus Coenia palustris (Ephydridae), male; Papp
1975
212 head Limnellia quadrata (Ephydridae);
Zatwarnicki 1997
213 head Thinoscatella spec. (Ephydridae);
Zatwarnicki 1997
214 head Odinia boletina (Odiniidae); Oldroyd 1970
215a habitus Prosopantrum flavifrons
(Cnemospathidae); Ismay 8c Smith 1994
215b detail wing Prosopantrum flavifrons
(Cnemospathidae); Ismay 8c Smith 1994
215c thorax Prosopantrum flavifrons
(Cnemospathidae); Ismay 8c Smith 1994
216a head Chiropteromyza wegelii
(Chiropteromyzidae); original, drawing by D.
Langerak
216b habitus Chiropteromyza wegelii
(Chiropteromyzidae); original, drawing by D.
Langerak
217 habitus Drosophila spec. (Drosophilidae);
Oldroyd 1970
218 head Chymomyza amoena (Drosophilidae);
Wheeler 1987
219 wing Drosophila deflexa (Drosophilidae);
Shorrocks 1972
220 frons Drosophilidae a: Leucophenga maculata, b;
Microdrosophila congesta; Bchli 1998 (drawings
by G. Bchli)
221 head Diastata adusta (Diastatidae); Oldroyd 1970
222 habitus Diastata fuscula (Diastatidae); Chandler
1986
223 head Euthychaeta spectabilis (Campichoetidae);
Chandler 1987
224 head Campichoeta zernyi (Campichoetidae);
Chandler 1987
225 wing Euthychaeta spectabilis (Campichoetidae);
Chandler 1987
226 antenna Tunisimyia shalam (Xenasteiidae);
Freidberg 1994
227 wing Tunisimyia shalam (Xenasteiidae);
Freidberg 1994
228 wing Asteia amoena (Asteiidae); Oldroyd 1970
229 head Asteia elegantula (Asteiidae); Papp 1973
230 wing Stenomicra cogani (Stenomicridae); Irwin
1982
231 head Stenomicra delicata (Stenomicridae); Collin
1944
232 head Santhomyza inermis (Anthomyzidae);
Rohcek 1998b (drawing by J. Rohcek)
233 femur fore leg Anthomyza gracilis
(Anthomyzidae); Mihlyi 1972
234 wing Paranthomyza nitida (Anthomyzidae);
Hennigi958
235 wing Cercagnota collini (Anthomyzidae);
Rohcek 1998b (drawing by J. Rohcek)
236 wing base Geomyza combinata (Opomyzidae);
Hennig 1958
237 habitus Geomyza combinata (Opomyzidae),
male; Sos 1981
238a head Canace salonitana (Canacidae); Mathis
1998 (drawing by W.N. Mathis)
238b wing Xanthocanace ranula (Canacidae); Hendel
1928
239a wing Chaetostomella cylindrica (Tephritidae);
Mihlyi i960
239b head Chaetostomella cylindrica (Tephritidae);
Merz 1994
240 wing Rhodesiella plumiger (Chloropidae);
Hennig 1958
241 head Ophiomyia maura (Agromyzidae); Spencer
1987
242 wing Melanagromyza laetifica (Agromyzidae);
Spencer 1987
243 habitus Acartophthalmus nigrinus
(Acartophthalmidae), female; McAlpine 1987a
2 4 4 habitus Themira putris (Sepsidae), male; Sguy
1934
245 head Heteromeringia nigrimana (Clusiidae);
Sos1959
246 habitus Heteromeringia nigrimana (Clusiidae),
female; Sos 1981 (drawing by J. Pal)
247 head Thyreophora cynophila (Piophilidae);
Hendel 1928
248 habitus Piophila casei (Piophilidae), male; Sos
1959
249 head Aulacigaster leucopeza (Aulacigastridae);
Oldroyd 1970
250 wing Aulacigaster leucopeza (Aulacigastridae);
Hennig 1958
251 wing Pelomyiella hungarica (Tethinidae); Munari
1998
252 head Pelomyia steyskali (Tethinidae); Munari
1998 (drawing by A. Szappanos)
253 head and thorax Borboropsis puberula
(Heleomyzidae); Gill 8c Peterson 1987
254 habitus Nidomyia cana (Heleomyzidae), male;
Papp 1998I
255 head Tethina strobliana (Tethinidae); Sos 1981
256 wing Chyromya flava (Chyromyidae); Hendel
1928
257a wing Parochthiphila coronata (Chamaemyiidae);
Hendel 1928
257b head Leucopis psyllidiphaga (Chamaemyiidae);
McLean 1998b
258 wing Leiomyza laevigata (Asteiidae); Hennig
1958
259 head Periscelis annulata (Periscelididae);
McAlpine 1987b
260 wing Periscelis annulata (Periscelididae);
McAlpine 1987b
261 wing Odinia maculata (Odiniidae); Papp 1998c
262 wing Turanodinia tisciae (Odiniidae); Papp 1998c
263 wing Liriomyza alpicola (Agromyzidae); Hendel
1931
264 wing Phytomyza affrnis (Agromyzidae); Hendel
1928
265 head Napomyza elegans (Agromyzidae); Oldroyd
1970
266 habitus Agromyza albipennis (Agromyzidae),
female; Spencer 1987
267 wing Paranthomyza nitida (Anthomyzidae);
Hennig 1958
268 femur fore leg Anthomyza gracilis
(Anthomyzidae); Mihlyi 1972
269 head Amygdalops thomasset (Anthomyzidae);
Rohcek 1998b (drawing by J. Rohcek)
270 head Aphaniosoma quadrivittatum
197
 (Chyromyidae); McAlpine 1987c
271 head Cacoxenus indagato! Loew (Drosophilidae);
Papp 1973
272a head Hemeromyia spec. (Carnidae); Papp 1998b
(drawing by A. Szappanos)
272b wing Hemeromyia spec. (Carnidae); Sabrosky
1987
273 head Desmometopa m - n i g T u m (Milichiidae);
Hennigi958
274 head Phyllomyza securicornis (Milichiidae);
Hennig 1937b
275 head Phyllomyza longipalpis (Milichiidae);
Hennig 1937b
276 head Madiza britannica (Milichiidae); Hennig
1937b
277 wing Madiza glabra (Milichiidae) ; Hennig 1937b
278 wing Milichia speciosa (Milichiidae); Hennig
I937t
279 wing Leiomyza laevigata (Asteiidae); Hennig
1958
280 wing Asteia amoena (Asteiidae); Oldroyd 1970
281 head Asteia elegantula (Asteiidae); Papp 1973
282 head Chloropidae: a: Trachysiphonella scuteilata;
b: Gampsocera numerata; c: Eurina calva; d;
Chlorops scutellaris; Dely-Draskovits 1978
283 thorax Chlorops certimus (Chloropidae);
McAlpine 1981b
284 habitus Meromyza pratorum (Chloropidae),
f e m a l e ; Balachowski 8c Mesnil 1935
285 wing Rhodesiella plumiger (Chloropidae);
Hennig 1958
286 head Carnus hemapterus (Carnidae); Sabrosky
1987
287 head Meoneura obscurella (Carnidae); Hennig
1937b
288 wing Carnus hemapterus (Carnidae); Sabrosky
1987
289 wing Meoneura obscurella (Carnidae); Hennig
1937b
290 habitus Coenia palustris (Ephydridae), male;
Papp 1975
291 wing Paralimna cinerella (Ephydridae); Hennig
1958
292 head Limnellia quadrata (Ephydridae);
Zacwarnicki 1997
293 head Thinoscatella spec. (Ephydridae);
Zatwarnicki 1997
294 head Pseudopomyza atrimana
(Pseudopomyzidae); Chandler 1983
295 head Pseudopomyza atrimana
(Pseudopomyzidae); Hennig 1958
296 wing Pseudopomyza atrimana
(Pseudopomyzidae); Chandler 1983
297 head Chymomyza amoena (Drosophilidae);
Wheeler 1987
298 wing Drosophila deflexa (Drosophilidae);
Shorrocks 1972
299 habitus Myopa stigma (Conopidae), male;
Bankowska 1979
300 habitus Leopoldius calceatus (Conopidae), male;
Bankowska 1979
301 wing Ulidia albidipennis (Ulidiidae); Hennig 1940
302 wing Colobaea bifasciella (Sciomyzidae);
Stackelberg 19893c
303 wing Eusapromyza poeciloptera (Lauxaniidae);
Papp 1978
304 head Sciosapromyza advena (Lauxaniidae);
Hennig 1958
198
305 head Saltella sphondylii (Sepsidae); Hennig 1958
306 wing Platystoma seminationis (Platystomstidae);
Oldroyd 1970
307 wing Salticella fasciata (Sciomyzidae); Rozkosny
1998a
308 head Coremacers marginata (Sciomyzidae);
Hendel 1928
309 habitus Psacadina verbekei (Sciomyzidae);
Knutson 8c Lyneborg 1965
310 habitus Pelidnoptera nigripennis (Phaeomyiidae),
male; Rozkosny 1998b (drawing by J. Pal)
311 head Eurygnathomyia bicolor (Pallopteridae);
Merz 1998b (drawing by B. Merz)
312 wing Eurygnathomyia bicolor (Pallopteridae);
Merz 1998b (drawing by B. Merz)
313 katepisternum Eurygnathomyia bicolor
(Pallopteridae); Merz 1998b (drawingby B. Merz)
314 head Neuroctena anilis (Dryomyzidae); Czerny
1930
315 habitus Dryomyza flaveola (Dryomyzidae);
Lindner 1948
316 ventral view thorax Dryomyza flaveola
(Dryomyzidae); Hennig 1958
317 ventral view thorax Helcomyza ustulata
(Helcomyzidse); Hennig 1958
318 ventral view thorax Helcomyza mirabilis
(Helcomyzidae); Steyskal 1987
319 head Helcomyza ustuhts (Helcomyzid3e);
McAlpine 1998c (drswing by A. Szsppanos)
320 wing Helcomyza ustulata (Helcomyzidae);
Czerny 1930
321 head Heterocheila buccata (Heterocheilidae);
Czerny 1930
322 habitus Cryptochetum grandicorne
(Cryptochetidae), female; Papp 1978
323 head Adapsilia coarctata (Pyrgotidae); Hendel 1928
324 wing Adapsilia coarctata (Pyrgotidae); Hendel
1928
325 head Loxocera aristata (Psilidae); Hendel 1928
326 wing base Psilidae; Oldroyd 1970
327 habitus Chyliza annulipes (Psilidae): Lyneborg
1964
328 head Orellia falcata (Tephritidae); W h i t e 1988
329 wing base Tephritidae; Oldroyd 1970
330 wing Stemonocera cornuta (Tephritidae); Merz
1994
331 habitus Urophora cardui (Tephritidae); Lindner
1948
332 wing base Strongylophthalmyia ustulata
(Strongylophthalmyiidae); Hennig 1958
333 habitus Dalmannia punctata (Conopidae), male;
Bankowska 1979
334 habitus Sepedon spinipes (Sciomyzidae), male;
Sguy 1934
335a femur hind leg Megamerina dolium
(Megamerinidae); Oldroyd 1970
335b head Megamerina dolium (Megamerinidae);
Hennig 1942
336 habitus Megamerina dolium (Megamerinidae);
Sos1959
337 habitus Sphyracephala europaes (Diopsidae),
male; Simova-Tosic & Stojanovic 1999
338 habitus Themira putris (Sepsidae), male; Sguy
1934
339 wing Micropeza corrigiolata (Micropezidae);
Hendel 1928
340 head Micropeza corrigiolata (Micropezidae);
Hendel 1928
341 head Calobata petronella (Micropezidae);
Oosterbroek 1981
342 wing Tanypeza longimana (Tanypezidae);
Hennig 1937a
343 habitus Tanypeza longimana (Tanypezidae);
Chandler 1975
344 wing Lipoleucopis praecox (Chamaemyiidae);
McLean 1998a (drawing by I. McLean)
345 wing base Parochthiphila spectabilis
(Chamaemyiidae); Hennig 1958
346 head Parochthiphila spectabilis
(Chamaemyiidae); McLean 1998a (drawing by I.
McLean)
347 wing Cremifania nigrocellulata (Cremifaniidae);
Czerny 1936
348 head Cremifania lanceolata (Cremifaniidae);
Papp 1994
349 wing Ulidia albidipennis (Ulidiidae); Hennig
1940
350 habitus Seioptera vibrans (Ulidiidae), female;
Lyneborg 1964
351 habitus Myennis octopunctata (Ulidiidae),
female; Lindner 1948
352 wing Pallopteraustulata (Pallopteridae); Morgeig74
353 habitus Palloptera modesta (Pallopteridae),
female; Morge 1974
354 wing Platystoma seminationis (Platystomatidae);
Oldroyd 1970
355 head Platystoma lugubre (Platystomatidae);
Mihlyi 1972
356 head Ulidia erythrophthalma (Ulidiidae);
Mihlyi 1972
357 habitus Dorycera graminum (Ulidiidae), female;
Mihlyi 1972
358 wing Geomyza hackmani (Opomyzidae); Drake
1993
359 wing Opomyza punctata (Opomyzidae); Drake
1993
360 head Anomalochaeta guttipennis (Opomyzidae);
Trojan 1957
361 head Calamoncosis sasae (Chloropidae);
Nartshuk 1987
362 wing Rhodesiella plumiger (Chloropidae);
Hennig 1958
363 habitus Azorastia mediterranea
(Nannodastiidae), male; Papp 1980
364 wing Azorastia mediterranea (Nannodastiidae);
Papp 8c Mathis 2001
365 wing Paralimna cinerella (Ephydridae); Hennig
1958
366 habitus Ochthera mantis (Ephydridae); Seguy
1926
367 habitus Pelomyia occidentalis (Tethinidae); Merz
et al. 2001
368 wing base Strongylophthalmyia ustulata
(Strongylophthalmyiidae); Hennig 1958
369 habitus Strongylophthalmyia ustulata
(Strongylophthalmyiidae); Papp 1978
370 tarsal claw Ornithomya (Hippoboscidae); Falcoz
1926
371 habitus Ornithomya rupes (Hippoboscidae),
female; Buttiker 1994 (drawing by W . Buttiker)
372 wing Brachytarsina diversa (Streblidae); Theodor
1968
373 head and fore legs Brachytarsina flavipennis
(Streblidae), male; Hrka 1998b (drawing by A.
Szappanos)
374 thorax Tachinidae; Colless 8c McAlpine 1991
375 head Trixa caerulescens (Tachinidae); Tschorsnig
8c Richter 1998 (drawing by H.-P. Tschorsnig)
376 habitus Gasterophilus intestinalis
(Gasterophilidae), female; Grunin 1969
377 habitus Oedemagena tarandi (Hypodermatidae),
female; Grunin 1964-1969
378 head Crivellia corinnae (Hypodermatidae),
female; Grunin 1964-1969
379 head Oestrus ovis (Oestridae); Colless 8c
McAlpine 1991
380 habitus Pharyngomyia picta (Oestridae), female;
Grunin 1966-1969
381 habitus Rhinoestrus purpureus (Oestridae),
female; Grunin 1966-1969
382 head Cephalopina titillator (Oestridae); Grunin
1966-1969
383 habitus Cephalopina titillator (Oestridae);
Grunin 1966-1969
384 thorax Pegomyia quadrivittata (Anthomyiidae);
Suwa 8c Darvas 1998 (drawing by M. Suwa)
385 habitus Graphomya maculata (Muscidae),
female; Sguy 1923
386 meron Chrysotachina alcedo (Tachinidae); W o o d
1987b
387 meron Winthemia fumiferanae (Tachinidae);
W o o d 1987b
388 wing Phyllomya volvulus (Tachinidae);
Tschorsnig 8c Richter 1998 (drawing by H.-P.
Tschorsnig)
389 wing Steleoneura czernyi (Tachinidae);
Tschorsnig 8c Richter 1998 (drawing by H.-P.
Tschorsnig)
390 scutellum Hylemya alcathoe (Anthomyiidae);
McAlpine 1981b
391 habitus Anthomyia spec. (Anthomyiidae), male;
Papp 8c Schumann 2000
392 head Craspedochoeta pullata (Anthomyiidae);
Hennig 1976
393 head Scathophaga stercoraria (Scathophagidae);
Sguy 1934
394 habitus Spaziphora hydromyzina
(Scathophagidae); Sguy 1934
395 wing Fannia canicularis (Fanniidae); Huckett 8c
Vockeroth 1987
396 habitus Musca domestica (Muscidae), female;
Lane 8c Crosskey 1993
397 wing Muscina levida (Muscidae); Huckett 8c
Vockeroth 1987
398 head Azelia cilipes (Muscidae), female; Hennig
1964
399 thorax Exorista larvarum (Tachinidae); McAlpine
1981a
400 scutellum and subscutellum Melanophora
roralis (Rhinophoridae); W o o d 1987a
401 scutellum and subscutellum Calliphoridae;
Mihlyi 1979
402 posterior spiracle Peribaea orbata (Tachinidae);
Draber-Monko 1989 after Crosskey 1977
403 posterior spiracle Phyto discrepans
(Rhinophoridae); Draber-Monko 1989 after
Crosskey 1977
4 0 4 posterior spiracle Melanophora roralis
(Rhinophoridae); Draber-Monko 1989 after
Crosskey 1977
405 habitus Ectophasia crassipennis (Tachinidae);
Mihlyi 1986
406 habitus Cylindromyia brassicaria (Tachinidae);
Mihlyi 1986
199
407 habitus Eginia ocypterata (Muscidae), male;
Sguy 1923
408 meron bristles Eginia ocypterata (Muscidae);
Gregor et al. 2002
409 wing Lucilia caesar (Calliphoridae); Rognes 1991
410 wing Stevenia deceptoria (Rhinophoridae); Pape
1998b (drawing by P. Lidmark)
411 habitus Melanomya nana (Calliphoridae); Mihlyi
1986
412 posterior spiracle Melanophora roralis
(Rhinophoridae); Draber-Monko 1989 after
Crosskey 1977
413 posterior spiracle Phumosia lutescens
(Calliphoridae); Draber-Monko 1989 after
Crosskey 1977
4 1 4 lower calypter and scutellum Rhinophoridae;
Mihlyi 1986
415 scutellum and subscutellum Melanophora roralis
(Rhinophoridae); W o o d 1987a
416 head Rhinomorinia sarcophagina
(Rhinophoridae); Mihlyi 1986
417 scutellum and subscutellum Calliphoridae;
Mihlyi 1979
418 wing Angioneura cyrtoneurina (Calliphoridae);
Rognes 1991
419 thorax Sarcophagidae (detail); Mihlyi 1979
420a habitus Sarcophaga carnaria (Sarcophagidae);
Pape 1987
420b wing Sphenometopa fastuosa (Sarcophagidae);
Venturi 1947
421 thorax Calliphoridae; Mihlyi 1979
422 thorax Protophormia terraenovae (detail)
(Calliphoridae); Rognes 1991
423 thorax Protophormia atriceps (detail)
(Calliphoridae); Rognes 1991
4 2 4 leg Frirenia tenella (Cecidomyiidae); Skuhrav
1997 (drawing by M. Skuhrav)
425 thorax Tipulidae; Hendel 1928
426 head Tipula carinifrons (Tipulidae); Tjeder 1963
427 habitus Tipula carinifrons (Tipulidae), female;
Lantsov 8c Chernov 1987
428 head Tricyphona hannai antennata (Pediciidae);
Tjeder 1963
429 habitus Tricyphona hannai antennata
(Pediciidae); Tjeder 1963
200
430 habitus Dicranomyia lindrothi (Limoniidae);
Tjeder 1963
431 copula Chionea scita (Limoniidae); Byers 1983
432 thorax Tricyphona hannai antennata (Pediciidae) ;
Tjeder 1963
433 copula Clunio pacificus (Chironomidae); Oka
1930
434 wing Telmatogeton pectinata (Chironomidae);
Goetghebuer & Lenz 1950
435 habitus Epidapus gracilis (Sciaridae), female;
Menzel 8c Mohrig 1997 (drawing by F. Menzel)
436 head Swammerdamella acuta (Scatopsidae);
Haenni 1997a
437 habitus Aprionus miki (Cecidomyiidae);
Skuhrav 1997 (drawing by M. Skuhrav)
438 underside Melophagus rupricaprinus
(Hippoboscidae); Theodor 8c Oldroyd 1964
439 tarsal claw Braula coeca (Braulidae); Hennig 1938
4 4 0 habitus Braula coeca (Braulidae); Hendel 1928
441 habitus Penicillidia dufouri (Nycteribiidae); Sos
1955
4 4 2 habitus Crataerina pallida (Hippoboscidae),
female; Buttiker 1994 (drawing by R. Heinertz
after original by Schneider-Orelli)
443 habitus Melophagus ovinus (Hippoboscidae),
female; Buttiker 1994 (drawing by R. Heinertz)
4 4 4 head and fore legs Brachytarsina flavipennis
(Streblidae), male; Hurka 1998b (drawing by A.
Szappanos)
4 4 5 habitus Aenigmatias lubbocki (Phoridae),
female; Oldroyd 1970
4 4 6 habitus Ariasella pandellei (Hybotidae), male;
Sguy 1926
447 habitus Aptilotus paradoxus (Sphaeroceridae),
female; Sguy 1926
4 4 8 habitus Tricimba brachyptera (Chloropidae),
female; Sos 1980
4 4 9 habitus Camus hemapterus (Carnidae), female;
De Meijere 1913
450 head Rhynchopsilopa nitidissima (Ephydridae);
Sguy1934
451 habitus Stiphrosoma sabulosum
(Anthomyzidae); Rohcek 2006
FIGURE CREDITS
Permission to use figures was gratefully
received from:
Minister of Public Works and Government Services,
Canada, 2005. Agriculture and Agri-Food Canada
(from McAlpine et al. (eds) 1981,1987, Manual of
Nearctic Diptera: 2a, 22,30,32,33, 35, 3 8 , 4 4 , 50,
51, 60, 65, 6 6 , 6 7 , 6 9 , 84, 85, 9 3 , 9 4 , 95,122,125,
1 2 9 , 1 4 6 , 1 4 7 , 1 9 4 , 1 9 5 , 204, 218, 241,242, 243, 253,
259, 260,266, 270,272b, 283,286, 288, 297, 318,
386, 387, 390, 395, 397, 3 9 9 . 4 0 0 , 4 1 5 , 4 6 3 , 567)
Apollo Books, Denmark (from Nilsson (ed.), 1997.
Aquatic Insects of North Europe, Vol. 2:105,212,
213,292,293,498, 503)
Brill Academic Publishers, the Netherlands (from
Fauna Entomologica Scandinavica: 88,156,159,
160,161,166a, 166b, 167,168,170,171,172,173,199,
409,418,420,422,423, 512)
British Entomological Society, United Kingdom
(from British Journal of Entomology and Natural
History: 358, 359)
Peter Chandler, United Kingdom (83,157,158,159,
160,165, 208, 222, 223, 224, 225,294,296, 343, 533,
546)
Chapman & Hall, United Kingdom (From Lane 8c
Crosskey, 1993, Medical Insects and Arachnids: 3,
4, 5, 6c, 7 , 1 2 , 1 3 , 1 4 , 21,153,176,177,179, 396)
Milan Chvla, Czech Republic (156,161,166a, 166b,
167,168,170,171,172,173, 512)
CSIRO Australia, Australia (from Colless 8c
McAlpine, 1991, Insects of Australia: 6a, 8,11,17,
18, 53. 59,73,75, r 3 . 1 0 4 , 1 6 2 , 374. 379)
Deutsches Entomologisches Institut des Leibniz -
Zentrums fr Agrarlandschafts- und
Landnutzungsforschung, Germany (from Beitrge
zur Entomologie): 191, 205, 207, 234, 236,240,
250, 258, 267, 273,279, 285, 291, 295,304,305, 316,
317.332. 345. 362,365, 368)
Entomological Society of Israel and Amnon
Freidberg, Israel (226,227)
Yves Gonseth, Switzerland (174b, 3 7 1 , 4 4 2 , 4 4 3 )
Frantisec Gregor, Czech Republic (178,408)
Jean-Paul Haenni, Switzerland (58, 6 1 , 6 2 , 4 3 6 , 556)
John Ismay, United Kingdom (215, 216,485)
Leif Lyneborg, Denmark 1964 (327, 350)
I.F.G. McLean, United Kingdom (257b)
Bernhard Merz, Switzerland (6b, 16, 20, 239b, 311,
312, 313,330,367, 573)
Henk Meuffels, the Netherlands (164)
NEST Foundation, Hungary (from Papp 8c Darvas
(eds), 1997 - 2000: Contributions to a Manual of
Palaearctic Diptera: 6b, 16, 20, 24, 27,42,42a, 43,
4 5 , 4 9 , 56, 57. 58, 61, 62, 6 3 , 7 0 , 7 1 , 7 2 , 7 4 , 89,101,
107,108,117,121,135,139,141,158,159,160,175,
190, 203,220, 232, 235,238a, 251, 252, 261,269,
272a, 307,310,311, 312,313, 319, 344, 346, 373,375,
384, 3 8 8 , 3 8 9 , 3 9 1 , 4 1 0 , 4 2 4 , 4 3 5 , 4 3 6 , 4 3 7 , 4 4 4 ,
4 7 4 , 4 7 9 , 4 8 9 , 4 9 3 , 4 9 4 , 506, 509, 516, 524, 533,
535, 537, 565, 578)
Natural History Museum, Basel, Switzer-land (from
Mitteilungen der Entomol-ogischen Gesellschaft
Basel: 367)
Stefan Niesiolowski, Poland (2b, 131,133,134)
Thomas Pape, Denmark (410,420)
Laszlo Papp, Hungary (183,186,192,193,198, 200,
201, 206, 211, 229,237, 239, 2 4 4 , 2 4 5 , 246, 248,
254, 255, 261, 262, 271, 281, 290, 303,322,336, 338,
348, 363, 364, 3 6 9 , 4 4 1 , 4 4 8 , 4 5 2 , 4 5 9 , 4 6 2 ,
4 6 7 , 4 7 1 , 4 7 2 , 4 7 8 , 4 8 3 , 4 8 4 , 4 9 2 , 4 9 5 , 515, 520,
525, 528, 531, 538, 549, 553, 557, 562, 566, 570, 575,
577, S81)
Adrian Pont, United Kingdom (199)
Knut Rognes, Norway (409,418,422,423)
Jindrich Rohcek, Czech Republic (185,451,457)
Royal Entomological Society, United Kingdom (from
Handbooks for the Identification of British
Insects: 97,111,138,155,163,180,181, 214, 217, 221,
228, 249,265, 280, 306, 326,329, 335a, 3 5 4 , 4 4 5 ,
4 6 4 , 4 6 6 , 4 9 7 , 511, 540, 552, 554, 558, 560, 568)
Rudolf Rozkosny, Czech Republic (88, 89, 93,117,
150, 307. 310, 564)
E. Schweizerbart'sche Verlagsbuchhandlung,
Germany (from Lindner (ed.), 1930-1974, Die
Fliegen der Palaarktischen Region: 19, 23,26, 34,
36, 37,40, 86, 98,109,115,120,130,132,142,174a,
263,274, 275, 276, 277, 278,287, 289, 301, 314, 315,
320, 321, 331, 335b, 342, 347, 349, 351, 376, 377,378,
380,381,382, 383,392,398,434,438,439,453,
4 9 0 , 505, 513, 532)
Bryan Shorroclcs, United Kingdom (219)
Duska Simova-Tosic, Serbia (337,496)
The author has taken much care to ask permission
from the copyright owners for the figures in
this book. If you think that your copyright is affected,
please contact the publisher.
201
ACKNOWLEDGEMENTS

The author would like to thank the following
persons and institutions: My good friend Nico
Schonewille for putting me on this project. The
co-authors of the Dutch precursor, Herman de
Jong and Liekele Sijstermans, w h o because of
lack of time could not participate in preparing
the present book. W i l l e m Hurkmans for trans-
lating the Dutch version. The publisher (KNNV
Publishing, Utrecht) for the generous financial
support with the translation and with the dis-
tribution of the test copy, and especially W i l m a
Seijbel and Rijnvis van W i r d u m , for advice and
guidance during the development stage of the
book.
Special thanks are due to all the people w h o
tested the key and/or critically reviewed the
text. The generous help received w i t h the
Dutch version is acknowledged therein. A test
copy of the English version was mailed to some
40 Dipterists throughout Europe and even fur-
ther. In return, many valuable corrections,
comments and additions were received. This
substantially improved the book and it is there-
fore a great pleasure to say thanks to Daniele
Avesani (Italy), Rudolf Bahrmann (Germany),
Miroslav Bartak (Czech Republic), Miguel
Carlos-Tolra (Spain), Peter Chandler (UK),
Pierfilippo Cerretti (Italy), Graham Collins
(UK), A m n o n Freidberg (Israel), Paul Gatt
(Malta), David Gibbs (UK), David Greathead
(UK), Lita Grve Jensen (Norway), Jean-Paul
Haenni (Switzerland), W i l l e m Hurkmans
(Netherlands), Ladislav Jedlicka (Slovakia),
Paul Kramer (UK), Nina Krivosheina (Russia),
Franco Mason (Italy), Bernard Merz
(Switzerland), Lorenzo Munari (Italy), Emilia
Nartshuk (Russia), Andrey Przhiboro (Russia),
Vera Richter (Russia), Jindrich Rohacek (Czech
Republic), Rudolf Rozkosny (Czech Republic),
Jukka Salmela (Finland), Wolfgang Schacht
(Germany), Vasily Sidorenko (Russia), Bradley
Sinclair (Germany), Daniel Whitmore (Italy),
Kaj Winqvist (Finland) and Joachim Ziegler
(Germany).
Finally, with respect to Chiropteromyza
wegelii Frey, I w o u l d like to thank Pekka
Vilkamaa (Finland) for checking the t y p e -
material, Jean-Paul Haenni (Switzerland) and
Laszlo Papp (Hungary) for the loan of speci-
mens, and Dick Langerak (Netherlands) for the
drawings he made so that illustrations for the
family Chiropteromyzidae could be included as
well.

ERWIN LINDNER
Die Fliegen der palaearktischen Region
Edited by Prof. Dr. Erwin Lindner

This large series was founded in 1924 and is now nearing

completion. It provides dipterists with basic information on
species, genera and families with the intention to describe all
DIE FLIEGEN
DER species known so far.
PALAEARKTISCHEN
REGION A l s o see the website: www.schweizerbart.de/joumals
to find information on all numbers published until today.

Contents of Vol. 1 (Handbook)

Geschichte der Dipterologie und Literatur, Morphologie und Terminologie, Anatomie,
Sinnesorgane, Metamorphose (Ei, Larve, Puppe, Pathenogenese), Biologie (Parasitismus,
Pflanzenschdlinge, Blutsauger, Krankheitsbertrger), Stammbaum, systematische Einteilung,
Fang, Zucht, Prparation, Sammlung.
Vol. II to XII

Descriptions of all families of the palearctic region with many figures, literature etc.

Please send orders or inquiries to

E. Schweizerbart'sche Verlagsbuchhandlung (Ngele u. Obermiller), Johannesstr. 3A
70176 Stuttgart / Germany. Tel./-Fax +49 (0)711 351 456-0 / -99 mail@schweizerbart.de

2 0 2
REGISTER OF FAMILY NAMES
(INCLUDING SYNONYMY)

Acartophthalmidae 67,110 Drosophilidae 63,73,77,109,132

Acroceridae 39,111 Dryomyzidae 81,132
Aenigmatidae 111 Eginiidae 133
Agromyzidae 67,71,111 Empididae 47, 53,123
Anisopodidae 37. " 2 Ephydridae 61,75,91,109,133
Anthomyiidae 97.99.ii2 Erinnidae 134
Anthomyzidae 65,73,109,112 Eurygnathomyiidae 134
Asilidae 45. " 3 Fanniidae 99,134
Asteiidae 65,73.3 Fungivoridae 134
Atelestidae 53,"4 Gasterophilidae 93,134
Athericidae 4i,"4 Helcomyzidae 81,135
Aulacigastridae 69,115 Heleidae 135
Axymyiidae 31,115 Heleomyzidae 59,69,109,135
Bibionidae 33,ii6 Helomyzidae 136
Blephariceridae 31,116 Hesperinidae 33,107,136
Bolitophilidae 37,116 Heterocheilidae 81,136
Bombyliidae 45, " 7 Hilarimorphidae 45,136
Borboridae 117 Hippoboscidae 93,109,137
Borboropsidae "7 Hybotidae 53,109,137
Brachystomatidae 45,"7 Hyperoscelididae 138
Braulidae 107,118 Hypodermatidae 95,138
Calliphoridae 103,118 Itonididae 138
Calobatidae 119 Keroplatidae 35,138
Camillidae 61,119 Larvaevoridae 139
Campichoetidae 63,"9 Lauxaniidae 57,77,139
Canacidae 65,120 Leptidae 139
Canthyloscelidae 33,120 Leptoconopidae 139
Carnidae 73.75,109,120 Leptogastridae 139
Cecidomyiidae 29,31,105,107,121 Lestreminiidae 139
Ceratopogonidae 31,121 Limnobiidae 134
Chamaemyiidae 69,87,122 Limoniidae 27,105,139
Chaoboridae 29,123 Liriopeidae 140
Chironomidae 31,105,123 Lonchaeidae 55,i4o
Chiropteromyzidae 61,123 Lonchopteridae 49,140
Chloropidae 75,91,109,124 Lycoriidae 141
Chyromyidae 69,73,124 Macroceridae 141
Clusiidae 57,67,125 Manotidae 141
Clythiidae 125 Megamerinidae 85,141
Cnemospathidae 61,125 Melusinidae 141
Coelopidae 59,126 Micropezidae 87,141
Coenomyiidae 39,43,126 Microphoridae 51,142
Conopidae 49,77,83,126 Milichiidae 73,142
Cordiluridae 127 Muscidae 99,101,143
Corynoscelidae 127 Musidoridae 143
Cremifaniidae 87,127 Mycetobiidae 35,H3
Cryptochetidae 81,127 Mycetophilidae 35,107,144
Culicidae 29,128 Mydidae 43,144
Curtonotidae 59,128 Mythicomyiidae 47, H S
Cylindrotomidae 27,129 Nannodastiidae 9i,i45
Cypselidae 129 Nemestrinidae 4L 145
Cyrtidae 129 Nemopalpidae 146
Dacidae 129 Neottiophilidae 146
Diadocidiidae 37,129 Nycteribiidae 107,146
Diastatidae 63,130 Ochthiphilidae 146
Diopsidae 85,130 Odiniidae 61,71,146
Ditomyiidae 35,130 Oestridae 95,147
Dixidae 29,131 Olbiogastridae 147
Dolichopodidae 51,109,131 Omphralidae 147
Dorylaidae 132 Oncodidae 147

203
 Opetiidae 5i,H7 Sepsidae 57, 67,77, 85,160
Opomyzidae 65,89,109,148 Simuliidae 31,160
Ortalidae 148
Otitidae Siphonellopsidae 161
148 Solvidae
Pachyneuridae 161
33,148 Spaniidae
Pallopteridae 161
79,89,148 Sphaeroceridae
Pediciidae 55,109,161
27,305,149 Stenomicridae
Periscelididae 65,162
7\ H 9 Stomoxydidae 162
Petauristidae 150 Stratiomyidae 37,43,49,162
Phaeomyiidae 79.150 Streblidae 93,109,163
Phlebotomidae 150
Phoridae Strongylophthalmyiidae 91,163
49,109,150 Synneuridae
Phryneidae 150 33,163
Syrphidae 47,164
Phthiriidae 150 Tabanidae 41,164
Piophilidae 67.151 Tachinidae 93,99,103,109,165
Pipunculidae 47,151 Taeniapteridae 165
Platypezidae 5h 152 Tanypezidae 87.166
Platystomatidae 67, 89,152 Tendipedidae 166
Pleciidae
33,107,153 Tephritidae 65, 83,166
Pseudopomyzidae 77,153 Tetanoceridae 167
Psilidae
83.153 Tethinidae
Psychodidae 69, 91,167
29.154 Thaumaleidae
Ptychopteridae 2
29.167
9.155 Therevidae
Pyrgotidae 45,i68
81,155 Thyreophoridae
Rachiceridae 168
37.155 Tipulidae 27,105,168
Rhagionidae
43.156 Trepidariidae
Rhinophoridae 101,156 168
Trichoceridae 25,169
Rhyphidae 157 Trichomyiidae 169
Risidae 157 Trixoscelididae 59,169
Sapromyzidae 157 Trypetidae 169
Sarcophagidae 103,157 Tylidae 169
Scathophagidae 97,99,157 Ulidiidae 79, 89,170
Scatomyzidae 158 Usiidae 170
Scatopsidae 31,107,158 Vermileonidae 37,43,170
Scenopinidae 45,47,158 Xenasteiidae 63,171
Sciaridae 33, 35,107,159 Xylomyidae
Sciomyzidae 77,79,85,159 37,43,171
Xylophagidae
Scopeumatidae 160 39,41,172

204
PUBLISHING DATA:

AUTHOR:
Pjotr Oosterbroek
Dept of Entomology, Zoological Museum
University of Amsterdam

TRANSLATION:
W i l l e m Hurkmans,
Bureau Hurkmans, Zwolle

GRAPHIC LAYOUT A N D DESIGN:

Erik de Bruin, Varwig Design, Hengelo

PRINTED BY:
D Z S d.d.

COVER ILLUSTRATIONS:

Front cover illustration:

Habitus Anthrax anthrax
(Bombyliidae), male
After Toth lgjj
Used by courtesy ofLdzlo Papp

Back cover illustration:

Brachycerousfly
After Lane & Crosskey, 1993
Reproduced with permission
from Chapman & Hall.

K N N V Publishing, Utrecht, 2 0 0 6 .
ISBN 9 0 - 5 0 1 1 - 2 4 5 - 5 / 9 7 8 - 9 0 - 5 0 1 1 - 2 4 5 - 1
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OTHER AVAILABLE TITLES ON INSECTS BY KNNV PUBLISHING

HOVERFLIES OF N O R T H W E S T EUROPE
Identification k e y s to t h e Syrphidae
M.P. v a n V e e n

T h i s b o o k identifies the hoverflies o f

N o r t h w e s t Europe and incorporates the k n o w
l e d g e t h a t h a s b e e n p u b l i s h e d i n m a n y articles
over past decades. T h e b o o k covers 500 species
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s y r p h i d fauna. T h e b o o k is a r i c h s o u r c e o f refe
r e n c e for e n t o m o l o g i s t s a n d specialists w o r
k i n g o n n a t u r e c o n s e r v a t i o n , as w e l l as for
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Hardcover \ 2005 I >,5 x 24 cm \ 256 pp \
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WATERBEETLES OF THE NETHERLANDS

M . B . P . D r o s t , H.P.J.J. C u p p e n ,
1 r !>l l w
l
E.J. v a n N i e u k e r k e n
DE WATERKEVERS
T h e p r e s e n t b o o k is an e x c e l l e n t a n d still u p - t o -
VAN NEBERLAND
date k e y t o all t h e 350 s p e c i e s o f w a t e r b e e t l e s
occurring in T h e Netherlands. W i t h 824
d e t a i l e d i l l u s t r a t i o n s , c o n t r i b u t i n g t o a reliable
identification.

Paperback \ Second edition 12006 | 17,5x25,5 cm

I 280pp I 34,95 I Dutch I 978-90-5011-053-2

THE DUTCH S P E C I E S OF C A D D I S LARVAE

Identification and e c o l o g y
Bert H i g l e r

The m a i n b o d y o f the b o o k consists o f k e y s to

all D u t c h s p e c i e s ( a b o u t 150) o f C a d d i s l a r v a e .
T h e a u t h o r d i d r e s e a r c h o n t h e s u b j e c t for
a b o u t 4 0 y e a r s , a n d is t h e D u t c h e x p e r t o n t h e
s u b j e c t . H e m a d e all o f t h e d r a w i n g s h i m s e l f .
To complete the b o o k it contains 2 4 w o n d e r f u l
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BSTRACT KEY

P u r p o s e o f this abstract is to facilitate finding a particular

s e c t i o n o f the m a i n key.

i Body and wing normal. 2

Body strongly flattened (fig. 441-443) and/or wing reduced or absent. 191
2(1) Antenna filiform or resembling a series of beads, with more than 6 seg
ments (fig. 17-19) 3
Antenna with less than 10 segments; third segment nearly always larger
than the subsequent segments (fig. 20-21). 44
3(2) 2 anal veins reaching the wing margin (fig. 23-35). 4"7
Only 1 (or no) anal veinreaching the wing margin (fig. 24). 8
8(3) Discal cell or cell dm absent (fig. 39-83). 9
Discal cell or cell dm present (fig. 38). 38-43
9(8) Ocelli absent. 10-18
Ocelli present (fig. 58,73). i9"37
44(2) Ptilinal suture absent (fig. 92-97). 45
Ptilinal suture present (fig. 91). 81
45(44) Thorax humpbacked and abdomen usually globose (fig. 98-99). Acroceridae
- Combination of characters not as given above. 46
46(45) Vein R4+5 forked into R4 and R5 and cell cup very elongate(fig. 105,113),
or wing venation aberrant as shown in fig. 107-109. 47
Not at the same time vein R4+5 forked into R4 and R5 and cell cup very
elongate (e.g. fig. 101-102,133). 62
47(46) Empodium pulvilliform (last tarsal segments with 3 cushions; fig. 103). 48-55
- Empodium bristle-like or absent (fig. 104). 56-61
62(46) Vein R4+5 forked (fig. 132-134). 63
Vein R4+5 not forked (all wing figures beyond 134). 64-80
81(44) Coxae of mid legs not far apart. Tarsal claws normal, not strongly curved.
Fly not ectoparasitic on birds or mammals. 82
Coxae of mid legs, and usually of the other legs as well, far apart (fig. 174a). Tarsal
claws strongly curved (fig. 174b) or last tarsal segment widened and eyes reduced
or absent (fig. 175). Fly living as an ectoparasite on birds or mammals. 169
82(81) Greater ampulla present (fig. 176). Usually 1 or more strong vibrissae pres
ent, as well as incurved lower F-bristles (fig. 178). 170
- Greater ampulla inconspicuous; if well developed, then vibrissae absent. 83
83(82) Haltere dark brown to black. Head large, wide and high (hemispherical);
only 1 pair of F-bristles, situated at the level of the oceilar triangle and
curving backward (fig. 182). Hind margin of anepisternum with a row of
bristles (fig. 183). Lonchaeidae
Haltere not dark brown to black or other characters different. 84
84(83) Vibrissae (fig. 184-189) or vibrissa-like bristles (fig. 186) present. 85
N o vibrissae or vibrissa-like bristles (fig. 187-188). 134
85(84) First tarsal segment of hind leg much swollen, and usually shorter than the
second segment (fig. 184,185 Sphaeroceridae
- First tarsal segment of hind leg not swollen, usually long and slender. 86
86(85) Dorsal preapical bristle at least present on tibia of mid leg, but usually on
all tibiae (fig. 190-204). 87
- All tibiae without dorsal preapical bristle. 101
87(86) Vein Sc complete, reaching the costa separate from vein Ri (fig. 190-193). 88-93
- Vein Sc incomplete, does not reach the costa separately: Sc reduced or
shortened (fie. 191), or merging with vein Ri before the costa (fig. 215b).
94-100
101(86) Vein R2+3 ends in the costa about halfway up the wing; vein Mi apically
pallid, not reaching the wing margin (fig. 227). Xenasteiidae
- Vein R2+3 is strikingly short (fig. 228) or ends in the costa distinctly beyond
the middle (fig. 230-235), or if vein R2+3 ends in the costa about halfway up the
wing, then vein Mi reaching the wing margin (fig. 240). 102
102(101) Wing base narrow, alula and anal vein small or absent (fig. 228-237). 103-105
W i n g base not narrow, alula well developed. 106
106 (102) Gena with 1 or more strong upcurved bristle(s) (fig. 238a). Canacidae
- Gena without strong upcurved bristles. 107
107(106) Vein Sc abruptly bent forward toward the costa at nearly 9 0 ; cell cup
closed by a geniculate vein CuA2 and with an acute apical end (fig. 239a).
Tephritidae
Vein Sc and cell cup not as given above. 108
108(107) Vein Sc complete, usually reaching the costa separate from vein Ri (fig
243-257)- 109-119
Vein Sc absent, or short and not reaching the costa, or merging with vein
Ri distinctly before the costa (fig. 240, fig. 258-262,267-298), or reduced to
a fold that may or may not reach the costa (fig. 263-264). 120
120(108) Crossvein BM-Cu present (cells bm and dm separate) (fig. 260-272^. 121-127
Crossvein B M - C u absent (cells b m and dm fused) (fig. 258). 128-133
134(84) A t the same time cell cup elongate (approaching the hind margin of the
w i n g ) and vein C11A2 (i.e., the vein closing cell cup) more or less straight;
cell r4+5 narrowing toward the wing tip, or closed (fig. 299- 300).
Conopidae
- Cell cup short or absent, its apex not near the hind margin of the wing (fig.
302-347), or if long, then closed by a geniculate vein CuA2 (fig. 301). 135
1 1
35( 34) Dorsal preapical bristle present at least on tibia of mid leg hut usually on
tibiae of all legs (fig. 309). 136-146
Tibiae without dorsal preapical bristle. 147
147(135) Arista absent; third antennal segment very large (fig. 322). Cryptochetidae
Arista present. 148
148(147) Ocelli absent. Pyrgotidae
Ocelli present. 149
0
149(148) Vein Sc abruptly bent forward toward the costa at nearly 9 0 , going
toward the costa as a transparent line or trace (fig. 326-330). 150
- Vein Sc not abruptly bent forward. 151
151(149) Vein Sc complete, its apical part not pallid, reaching the costa separately
from vein Ri (fig. 333-357). 152-163
Vein Sc absent, shortened, merging with Ri before the costa, or its apical
part pallid and vague (fig. 332, 358-369). 164-168
170(82) Mouth opening small, its diameter up to 1 / 4 to 1/8 of the width of the
head; mouthparts minute, rudimentary (fig. 375-382). 171-174
Mouth opening large, its diameter at least 1 / 4 of the width of the head;
mouthparts well developed, well visible (fig. 4,7-8,393-394). 175
175(170) Meron without one or more rows of bristles near its hind margin (fig. 384).
176-180
Meron with bristles near the hind margin, arranged in one (fig. 386) or
several (fig. 387) rows, or in a small cluster (fig. 408). 181-190
191(1) Antenna usually long, filiform or resembling a string of beads, with 6 or
(usually) more segments; usually all segments or those beyond the second
more or less similar (fig. 426,437). 192-205
Antenna not filiform or resembling a string of beads, with less than 6 seg
ments; usually the first 2 segments short, the third segment large and the
subsequent segments making up an arista or style. 206-220
 The European
Families of the Diptera
Identification, diagnosis, biology

T
ihe European Families of the Diptera
presents an identification key and family
descriptions of all 132 families of Diptera
(midges, mosquitoes, gnats, true flies) occurring
in Europe. It is written by a well-known Dutch
specialist, in collaboration w i t h over 30
European dipterists.
For the extensive identification key a new
combination of important characters is employ
ed, enabling relatively easy identification of
families which are aberrant or traditionally
considered difficult to identify. Over 600 line
drawings are included to illustrate characters
and families. Apart from the key, the book
includes an extensive chapter on terminology.
Information on systematics and the number of
genera and species in Europe, a survey of the
main characters, a summary of the biology and
of the pertaining identification literature is
included for all families.
Due to its design and content, the book will
be of interest to the lay public and the serious www.knnvuitgeverij.nl
amateur alike.