8

Identification

Concepts differ from facts by continuing to change over time (Mayr, 1991)

8.1 INTRODUCTION

The principal impediment to the study of the natural history of scuttle flies
has been the difficulty of identifying specimens with confidence. For several
regions the lack of a key to genera means it is difficult to even start. The use of
a generic key for another region can leave the user wondering whether an
error has been made at an earlier stage. The aim of this chapter, therefore, is to
provide a key to the genera of the world currently recognised in the literature
and to indicate which keys or papers are required for identification to the
species level.

8.1.1 SYSTEMATICS
The most recent key to the genera of the world is that by Schmitz (1929g).
This dealt with nearly 100 genera. Section 8.3 lists 229 genera, allowing for
some reinterpretations proposed below. Not all of these genera will prove to
be valid. Some will prove to be the 'missing' sex of another genus currently
only known in one sex. More than 50% of the described genera are still only
known in one sex. It seems likely that some modern molecular techniques
could be used to establish the conspecificity of sexually dimorphic species
whose two sexes are currently assigned to separate genera, each of which is
currently considered to be only known in one sex. Some genera will
disappear as they are amalgamated with others, following the description of
new species which erode the gaps between two genera. Others will be split up
as it is discovered that they are polyphyletic. New genera will continue to be
described, particularly as tropical faunas become better known.
Implicit in these few comments is the recognition that a genus embodies a
set of opinions about the affinities of species and the status of the clusters of
species we have constructed to reflect those opinions. The process of
clustering related species, systematization, is in principle a process of
scientific discovery through the erection of testable hypotheses. The

R. H. L. Disney, Scuttle Flies: The Phoridae

© R H L Disney 1994
 IDENTIFICATION

assignment of a rank (genus, subgenus or whatever) to a cluster of species,

categorization, owes as much to historical accident and prejudice as to
scientific insight. A classification, therefore, is partly a reflection of the
results of scientific discovery and partly the result of arbitrary decisions. The
latter introduce an element of historical constraint, which needs to be
respected in the interests of nomenclatural stability. It is too often forgotten
that a classification is meant to be a system of reference to aid communi-
cation. Continual changes in names, or the concepts signified by them,
impair effective communication.
I conclude that even if we were confident that all species had been correctly
assigned to monophyletic clusters, the status of any such monophylum is to a
large degree a matter of opinion. My own prejudice is in favour of
minimizing unnecessary nomenclatural changes. Thus I will change a name if
I discover evidence that a species has been assigned to the wrong genus. I
object, however, to the practice of raising subgenera to the generic level in a
futile attempt to render different genera 'equivalent' (whatever that might
mean - see Disney, 1983d). Likewise I recognize no obligation to reduce a
genus to the subgeneric level because it is found to be a distinct clade within a
larger monophyletic 'genus'. For example, the genera Gymnophora and
Woodiphora are essentially distinct clades within a monophylum comprising
these two genera plus the genus Megaselia (Disney, 1989f). The purist will
argue that this means the 'genus' Megaselia is, therefore, an incomplete
monophylum (a paraphyletic taxon) and that this is unacceptable. The
solution, it is argued, is to resolve the paraphyly either by reducing the
excluded 'genera' to subgenera or by raising the subgenera of Megaselia to
generic level. If we were in the position to adopt the second option (which we
are not, in our present state of knowledge), the result would be to change the
names of over 1300 species at a stroke. If we pursued the first option, then all
the species of Gymnophora and Woodiphora would have to change to become
species of Megaselia. It is for this reason that I have proposed (Disney, 1989j)
that we accept that we will be stuck with numerous unresolved paraphyletic
genera as far as we can see into the future. If we wish to indicate that we have
identified cases of this, as with Megaselia, we can do so by calling these
paraphyletic genera 'Paragenera'. In these cases the first word of the species
binomial, e.g. Megaselia in the case of Megaselia scalaris, is then a paragenus
rather than a genus in the narrow cladistic sense. In the interests of
nomenclatural stability I make no apologies for this conscious adoption of
non-equivalent notions of genera being applied to different clusters of phorid
species. I repeat that a major function of a formal classification is as a system
for the ordering and retrieval of biological data. The purist's abhorrence of
para genera is a recipe for continuous name changing, to the detriment of
unambiguous communication.
The notion of a genus may be contentious, but the suprageneric
classification is even more so. In section 8.4 I indicate the sort of problems
currently being debated. Because there is, as yet, no consensus I offer no key
to subfamilies and tribes, but go straight into keys to genera, one for males
and one for females. These keys are followed by a guide to the literature for
each genus. A section on fossil Phoridae then follows.

204
 INTRODUCTION

cercus

hypandrium

(b)
Figure 8.1 (a) Frontal view of head of Triphleba nudipalpis male, showing bristles
(0 = ocellars, PL = postero-lateral, PO = pre-ocellar, ML = medio-Iateral, AL =
antero-lateral, A = antial, SA = supra-antennal). (b) Side view of male hypopygium
of a generalized Megaselia-type (T6 = abdominal tergite 6). (Scale bars = 0.1 mm, as in
all other figures.)

205
 IDENTIFICATION

MF
o

I
o
o

(a)

,"" ,?
. ..1:'../
\ . ", ,/

'-:::'~~
/ ~':: ~: .. ,

(c)

Figure 8.2 Details of head. (a) Frons of Megaselia deningi male, showing median
furrow (MF) and bristle sockets, labelled as in Fig. 8.1 (a). (b) and (c) Proboscis of male
(b) and female (c) of M. chlumetiae (L = labellum, U = labrum). (d) Antenna of male
M, microcurtineura (1-3 = segments 1-3).

8.1.2 MORPHOLOGY
The principal details of the morphology of a scuttle fly are indicated in Figs
8.1-8.15. The head is typically as in Fig. 8.1(a). In many genera a median
furrow (MF in Fig. 8.2a) runs from the anterior ocellus to the front margin.
The 'standard' set of bristles on the frons is labelled in Fig. 8.1 (a). Bristles are
liable to be knocked off, but their positions are always indicated by their basal

206
 INTRODUCTION

sockets (Fig. 8.2a). In many genera a pair of the 'standard' bristles may be
absent. Sometimes there are additional bristles, especially more than four
supra-antennals. The proboscis is frequently sexually dimorphic (e.g. Figs
8.2b,c), mainly in terms of the breadths and lengths of the labrum and labella.
The antennae (Fig. 8.2d) typically comprise three basal segments and a
three-segmented arista, but the arista may have a reduced number of
segments or be absent. The first (basal) segment (the scape) (1 in Fig. 8.2d) is
fully exposed; the second (the pedicel, labelled 2) is largely enveloped by the
enlarged third segment (the first flagellar segment, labelled 3). The third
segment may be elongated (e.g. Fig. 8.31a).
The thorax viewed from above mainly comprises the enlarged scutum
(mesonotum) and the scutellum behind. In flightless females the scutellum is
frequently reduced or absent. Details of the sides of the thorax are useful in
both classification and identification. The principal features are labelled in
Fig. 8.3. There is sometimes a gland opening at the front end of the
notopleuron (the edge of the scutum bordering the mesopleuron), which
may be partly or completely concealed in a notopleural cleft (Fig. 8.28b and
8.93b,c).
The legs are typical in having a coxa, trochanter, femur, tibia and tarsus,
the latter normally having five segments (Fig. 8.4a). Sometimes the front
tarsus has only four or two segments. The fifth tarsal segment may be longer
than the fourth, and is sometimes tapered (e.g. Fig. 8.4b). The presence of
isolated bristles on the tibiae (Fig. 8.4c) is a useful character. Likewise the
presence of, usually near-dorsal, longitudinal hair palisades (Figs 8.4d,e), as
opposed to spaced spines (e.g. Fig. 8.4c), is a useful character.
The wings have a surprisingly uniform venation within the family. De-
spite this they exhibit a number of variations useful in identification. The
anterior series of veins, the so-called 'thick veins', comprise the following.
The marginal costal vein (C in Fig. 8.Sb) bears costal cilia (c in Fig. 8.Sa),
usually in two rows. In published descriptions lengths of costal cilia refer to
those on the anterior row of costal section 3. Typically the costa extends
about half the wing length (measured from the level of the basal bristle - B
in Fig. 8.Sb - along the mid line to the wing tip). It may be longer or
shorter. In the traditional phorid literature the Costal Index is much used. It
is the length of the costa (measured from the basal bristle) divided by the
wing length. The subcosta may extend to vein 1 and be fused to it (S in Fig.
8.Sb), be free (e.g. Fig. 8.Sa) or be absent. Vein 1 (Rl) occasionally has hairs
along its upper face. Vein Rs (R in Fig. 8.Sb) may also have hairs along its
upper face, but typically has only one at the base (e.g. Fig. 8.Sh) or none. It
frequently divides into vein 2 (R2+3) and vein 3 (R4+S). When vein 2 is
missing, the single vein Rs/R4+ S is still referred to as vein 3, and is said to
be unforked. In the traditional literature the costal ratios are much used.
These are the ratios of the lengths of the costal sections 1 to 3, or 1 to 2 only
if vein 3 is unforked, to each other (Sl, S2 and S3 in Fig. 8.Sb). They are
best expressed by dividing the lengths of 1 and 2 hy the length of 3, so that
section 3 always scores as 1.
The abdomen typically has tergites on the first six segments, and typically
no sternites. Rarely there is a sternite on segment 6 only. Even more rarely

207
 ,
.'

Figure 8.3 Left sides of thorax of Megaselia ciliata (a) and Conicera dauci (b).
s = spiracle, Pr = propleuron, Me = mesopleuron, f = mesopleural furrow,
hu = humerus, Pt = pteropleuron, St = sternopleuron, Hy = hypopleuron, ha =
halt ere, c = coxa.

208
 INTRODUCTION

(a)

(b)

! "

1/ I1
! II
I III ~
p ,',I, \
1
/ \I \

(c)
(e)

Figure 8.4 Parts oflegs, (a) Hmd leg of Microselza beaveri female, omlttmg coxa and
base of trochanter. (b) M, beaven female, last three tarsal segments of mIddle leg,
(c) Front tibIa and tarsus of Spimphora bergenstammi, showmg an Isolated bnstle and
spaced spmes on tibIa and smgle spmes (s) at tips of first two tarsal segments, (d) and
(e) postenor faces ofhmd tibiae bearmg a near-dorsal haIr palisade (P), of Pseudacteon
brevicauda (d) and Megaselia biarticulata (e),

there is a sternite on segment 4 and vestiges on segment 3. Tergite 1 is

typically shorter than 2 and frequently the tergites are progressively narrower
from 2 onwards (e. g. Figs 8.6a and 8. 7c). In females the tergites of the first six
segments may be reduced in both size and number. The terminal segments of
the abdomen are discussed in section 8.2(A) below.
The terms used for the parts are those most widely used in the literature on
the family. In recent years there has been a tendency to modify the

209
 IDENTIFICATION

81 82 S3
B

(b)
Figure 8.5 DetaIls of WIng (a) RIght WIng of Megaselta stenoterga male (A = aXIllary
bnstles, c = costal CIlIa on costal veIn, 4-7 = tIpS of veInS 4-7). (b) Antenor basal
regIOn of nght WIng of M. subtumlda female (B = basal bnstle of costa, H = humeral
veIn, C = tIp of costal veIn, 1-3 = veInS 1-3, S = subcostal veIn (Sc), Sl-S3 = costal
sectIOns 1-3).

morphologIcal terms m every new major text on the Diptera of a particular

biogeographic region. There IS little sIgn of any stability being achieved in the
lIterature of the second half of the twentieth century. In view of this it is
considered more useful to stick with the tradltionallabels. The passion to
aVOId identical terms in dIfferent orders of ammals, in case these are taken to
Imply homology of the structures so labelled in the two orders, is seen to be
absurd if taken to its logical conclusion. Thus the labelling of a leg bone in
man as the 'femur' certainly has precedence over the labelling of the section of
an msect leg with the same term. The lOgIC of the purist's argument,
therefore, IS that we should com new terms for such parts of the msect leg! I

210
 KEYS TO GENERA

consider it most useful to employ the terms most frequently employed in the
literature required to identify specimens to the species level.

8.2 KEYS TO GENERA

Many specimens can be identified to the genus by examining them in a

watchglass of 70% alcohol under a stereo, dissecting, microscope with a
good light. I use a relatively cheap instrument but with a separate fibre-optic
lighting system. I mainly use this, however, for making slide mounts of the
flies. Details of slide mounting are given in section 9.7. Slides are examined
under a compound microscope using X 10, x20 and x40 objectives. Success
in seeing fine detail tends to be achieved more often by adjustment of the
lighting than by switching to a higher magnification!
In the following keys variation between species in a genus has been dealt
with by keying out some genera at more than one couplet. Also some
couplets can be reached by more than one route. The effect of this device is
that by one route the user may only confront a particular choice once, but by
another route a particular choice may be encountered twice.
A number of genera, particularly several small Neotropical genera, I have
encountered only in the literature. There is, therefore, an increased risk of
misapprehension in my perception of these genera. I would welcome
notification of any errors in these keys. Comparison with other published
keys to phorid genera will quickly reveal that those I offer below represent a
radical new approach. Having been repeatedly frustrated when trying to use
the existing keys, I have adopted an entirely novel arrangement. The most
radical difference is in my total disregard of the supposed subfamilies or
tribes. The result is that quite unrelated genera may end up in the same
couplet, simply because they resemble each other in terms of the recognition
characters I have chosen to emphasize.
The presence or absence of certain bristles are very useful in the recognition
of certain genera. The basal sockets of detached bristles remain intact and can
be readily observed. Traditional keys have given great emphasize to the
inclination of the 'supra-antenna!' bristles as well as of the 'antials'. I have
disregarded this, as it is evident that this is not as significant, taxonomically,
as has been supposed. Furthermore some of the differences given in the
literature are almost certainly based on erroneous homologies of the bristles
in question. Thus some cases of upswept 'supra-antennals' in conjunction
with 'missing antials' could equally plausibly be interpreted as missing
supra-antennals and approximated antials! In view of this sort of possibility
for confusion, it seemed wiser to use other characters.

(A) DETERMINA TION OF THE SEX OF A

SCUTTLE FLY

The first task is to distinguish between the males and the females. In some
females there may be a reduction in the size and number oftergites on the first

211
 IDENTIFICATION

(a)
I (b)

Figure 8.6 Parts of female abdomen. (a) Tergites 1-6 of Megaselia humida.
(b) Tergite 7 and terminal segments of M. humida from above. (c) Terminal segments
of M. termitomyca in side view .

six abdominal segments. The main external differences between the sexes,
however, are in the terminal segments (those beyond segment 6).
InJemales the two cerci are separated (Figs 8.6b and 8. 7a). In some they are
reduced (e.g. Fig. 8.8a), or absent, but in these cases the abdomen ends in an
obvious ovipositor-like structure (e. g. Figs 2.5-2.7). The basic abdominal

212
 KEYS TO GENERA

~---

J
\

~u
T7[]
T8 \J
(b) V
(c) c::'~
Figure 8.7 Female abdominal tergites and sternites of Megaselia subnudipennis.
(a) Tergite to epiproct. (b) Sternite 7 to hypoproct. (c) All tergites and epiproct.
(T6--T10 = tergites 6--10, S7 and S8 = sternites 7 and 8, C = cercus, E = epiproct,
H = hypoproct.)

terminalia are as in Figs 8.6b,c and 8.7. Tergite 9 is normally fused to the
anterior apodemes oftergite 10, thus appearing to be absent (Fig. 8.7a). Only
rarely is it distinct (e.g. Fig. 8.8a). Sternite 8 may be simple (e.g. Fig. 8.8c),
modified (e.g. Fig. 8.8b) or divided into two (e.g. Fig. 8.7b). Sternite 9 is
either absent or internalized. In the latter case it is typically ring-shaped (e. g.
Fig. 8.8a), but is occasionally a more elaborate structure bearing hairs.

213
 IDENTIFICATION

87

(a)
(b)
I

(e)

Figure 8.8 Female abdommal tergltes and stermtes (a) Termmal tergltes of
Chonocephalus blacklthorum (b) Stermtes of C blacktthorum (c) Stermte 8 of
Chaetopleurophora erythronota T7-10 = tergltes 7-10, S7-9 = stermtes 7-9, sp =
spermatheca, H = hypoproct, c = cercus

In males the cerCI are closely approxImated and lIe along the dorso-Iateral
regIOns of the proctIger, frequently bemg completely fused to the latter, and
the three structures form the anal tube (FIg 8 Ib) ThIs anal tube emerges
from the postenor regIOn of the epandrium (= the enlarged TergIte 9) The
latter embraces the pems complex, whIch may be completely wIthdrawn
(FIg 8 9a) or be fully extruded (FIg 8 9b) Beneath the pems complex lIes the
hypandrium (FIg 8 1b), but m some genera an asymmetnc development of
the epandnum may cause the hypandnum to be dIsplaced to one SIde
SometImes the hypandnum and epandnum are fused to form a smgle
structure Adjacent to, and below, the anal tube there may be a paIr of
surstyli(Flg 810) Thesemaybefusedtotheepandnum(e g FIg 811a,b),
to form surstylar lobes A surstylus may be present on one SIde only (e g FIgs
8 llc and 8 12a,b) SometImes the terglte and stermte of segment 10 are stIll
retamed and fused to form a rmg (FIg 8 12c) or cylmder (FIg 8 13b) at the

214
 KEYS TO GENERA

(b)

Figure 8.9 Male hypopygla from left sIdes. (a) Megaselw nepenthina with pems
wIthdrawn. (b) M barbulata with penis extruded.

base of the anal tube. In the latter case any surstyli may be fused to this
cylinder (e.g. Fig. 8.13a). Normally the remnants of the sclerites of segment
10 are lost or fused, at least in part, to the rear of the epandrium. The
hypandrium frequently has one or two posterior lobes (e.g. Fig. 8.14b) and

215
 IDENTIFICATION

Figure 8.10 Male hypopygium of Beyermyia spinulicosta from right (a) and left (b)
sides. (S = surstyli.)

216
 KEYS TO GENERA

(a)

Figure 8.11 Male hypopygia. (a) Maculiphora aenictophila left face and (b) the same,
right face. (c) Megaselia biarticulata left face.

may rarely carry a pair of gonopods (e.g. Fig. 8.1Sa). The latter, however, are
usually absent or represented by a pair of bristles only (Fig. 8.14a). The entire
terminal complex (i.e. segment 9 and all beyond) is called the hypopygium.

217
 IDENTIFICATION

...

Figure 8.12 Male hypopygia. (a) and (b) left and right faces of Phora penicillata.
(c) left face of Dohmiphora comuta (T9 = tergite 9, C = cercus, T10 + S10 = fused
tergite 10 and sternite 10 to form a ring).

218
 KEYS TO GENERA

T10

(b)

Figure 8.13 Left faces of male hypopygJa of Diplonevra. (a) D. predicta. (b)
D. nitidula (T9 = tergJte 9 (epandnum), TI0+SI0 = fused terglte and stermte 10 to
form cylmder, C = cercus).

219
 IDENTIFICATION

,~
:::"

(a)

(b)
Figure 8.14 Male hypopygia. (a) Gymnophora healeyae hypandrium from below.
(b) Megaselia badia, left face ofhypopygium.

220
 (b)

Figure 8.15 (a) Ventral view of male hypopygium ofChonocephalus blackithorum (S9
= sternite 9 (hypandrium), RG and LG = right and left gonopods). (b) Acontistoptera
brasiliensis, right wing offemale.

221
 IDENTIFICA TION

(B) KEY TO GENERA OF MALES

1. Wings absent 2
Wings present 3
2. Fewer than 15 ommatidia in each eye Arrenaptenus
More than 40 ommatidia in each eye Aptinandria
3. Vein Rs (i.e vein 3 before any apical fork) with several hairs along the
dorsal face (e.g. Fig. 8.21a). These hairs may be short and fine (or at
least shorter and finer than a more bristle-like hair at base of vein) 4
Apart from a few hairs (usually only one) in the basal quarter, the dorsal
face of vein Rs is bare (apart from microsetae) 13
4. Vein 1 with hairs along its dorsal face Phlebothrix
Vein 1 bare on its dorsal face 5
5. Hind tibia with at least one dorsal, or near-dorsal, longitudinal hair
palisade (e.g. Figs 8.4d,e) 6
Hind tibia without dorsal hair palisades, but there may be spaced hairs
(as in Fig. 8.4c) 10
6. A longitudinal hair palisade present on front tibia 7
Front tibia without a dorsal, longitudinal hair palisade, but there may
be a row of spaced bristles or spines 8
7. Distance between anterior ocellus and front of frons much greater than
distance between two posterior ocelli. Vein 3 forked or unforked (Fig.
8.21a) Aenigmatias
Distance between the anterior ocellus and front of frons less than
distance between posterior ocelli. Vein 3 unforked Misotermes
8. Height offrons much greater than width Trineurocephala (part)
Frons width about equal to height (from anterior ocellus to front
margin) or greater 9
9. Posterior ocelli close to eye margin and ocellar triangle strongly
demarcated at front by a sinuous furrow (Fig. 8.94d) Stichillus
Posterior ocelli well removed from eye margin and ocellar triangle not
demarcated in this way at front Borophaga
10. Third antennal segment apically elongated and tapered, with apical
arista Burmophora
Third antennal segment more globose; arista frequently pre-apical and
dorsal in position 11
11. Costa less than two fifths of wing length. Medio-Iateral and pre-ocellar
bristles absent Euryophora
Costa extends about half wing length, or more. Medio-Iateral and
pre-ocellar bristles present 12
12. More than 10 hairs on vein Rs Anevrina
Fewer than 10 hairs on vein Rs 13
13. Apex of third antennal segment drawn out into long, narrow,
long-haired process; with longer, short-haired, arista originating near
base of this apical process Diplostylocera
Antenna not of this form 14
14. Apex of third antennal segment drawn out into long, tapered extension
bearing apical arista (e.g. Figs 8.16b and 8.31a) 15

222
 KEYS TO GENERA

- Apex of third antennal segment not drawn out in this way, except in
some cases where arista is pre-apical in position or is absent. Sometimes
arista replaced by apical pseudo-arista (which is an extension of the
third antenna I segment itself, rather than articulating with it) 39
15. Arista with only one or two segments and mesopleuron bare 16
- Without this combination (i.e. arista normally with two short basal
segments and long apical segment, but if with only one or two
segments, then mesopleuron has hairs) 17
16. Arista with single long segment. Veins 1 and 3 well separated apically
(Fig. 8.36b). Only fine hairs below basal half of hind femur Dahliphora
(part)
- Arista short and with two segments (Fig. 8.49c). Veins 1 and 3 closely
approximated (as Fig. 8.s0d). Base of hind femur with spinules (Fig.
8.49b). (Hypopygium as Fig.4ge) Iridophora
17. Mid tibia with one or two differentiated dorsal or near-dorsal bristles in
basal half 18
- Mid tibia without such bristles, although a row of postero-dorsal
differentiated hairs may be present along the length of tibia 25
18. Only one bristle in basal half of mid tibia. Vein 3 forked or un-
forked 19
Two bristles in basal half of mid tibia. Vein 3 unforked 20
19. Vein3unforked 83
- Vein 3 forked 84
20. Hind tibia with two longitudinal hair palisades Antipodiphora
- Hind tibia without any longitudinal hair palisades 21
21. Hind tibia with isolated bristles including at least one pair (one dorsal
and one antero-dorsal) in basal half(e.g. Fig. 8.32a) 22
- Hind tibia without any isolated bristles situated close enough to another
to form such a pair in basal half 24
22. Hind tibia with at least eight bristles before apical quarter Contopteryx
- Hind tibia with only four bristles in upper three-quarters 23
23. Hind tibia with lower dorsal bristle beyond middle (Fig. 8.32a). Vein Sc
does not reach vein 1 (Fig. 8.31c). (Antenna and claspers (surstyli) as
Fig.8.31b) Conicera (part)
Hind tibia with lower dorsal bristle just before middle. Tip of vein Sc
reaches vein 1 and fuses with it Darwiniphora (part)
24. At least one strong spine at postero-lateral extremity of left side of
epandrium (Fig. 8.47c) Gymnoptera (part)
No such spines on epandrium (see also couplet 71) Coniceromyia (part)
25. Hind tibia with a longitudinal dorsal hair palisade 26
- Hind tibia without a dorsal hair palisade, but there may be a row of
spaced hairs 34
26. Vein 3 forked 27
- Vein 3 unforked 31
27. Frons breadth equal to or clearly greater than height 28
- Frons clearly with height greater than breadth 39
28. Anal tube not straight, narrow and long (not exceeding mid-line length
ofepandrium) 29

223
 IDENTIFICATION

Anal tube strikingly long (exceeding mid-line length of epandrium),

straight and narrow 39
29. Mesopleuron with hairs and a bristle 30
Mesopleuron bare 39
30. Antero-Iateral bristles absent from frons Acanthophorides
Antero-Iaterals present Macrocerides (part)
31. Mesopleuron with hairs. (Arista very short.) Tubicera (part)
Mesopleuron bare 32
32. Tip of vein 3 does not reach margin of wing but runs to tip of the costa,
which is deflected rearwards through a right angle bend Stenoneurellys
Tip of costa not deflected in this way 33
33. Anal tube long and slender, being clearly longer than length of
epandrium 39
Anal tube only about as long as epandrium and not obviously
slender Ceratoconus
34. Vein 3 forked Rhopica
- Vein 3 unforked 35
35. Costa extends less than one-third of wing length. Tapered extremity of
third antennal segment bears hairs which are clearly much longer than
those on segment 3 of arista Leptilla
Costa extends for at least half wing length. Hairs on tapered extremity
of third antennal segment little, if any, longer than those on segment 3
ofarista 36
Note: If neither applies try couplet 196.
36. Vein six deflected anteriorly in middle section, so that space embraced
by it and vein 5 has characteristic 'waist' (Fig. 8.56a) Metopina (part)
Vein six not deflected in this manner 37
37. Frons clearly broaderthan high 38
Frons with height at least as great as breadth or greater. Aristocerina
38. Frons bearing conspicuous, strongly developed, bristles (Fig. 8.16b).
Mesopleuron bare (see also Figs 8.16a, 8.17a) Adelopteromyia
Frons devoid of bristles. Mesopleuron with a few hairs dorsally
Cyphocephalus
39. Front part of frons at a right angle to the top face of frons, the transition
being marked by an arch-shaped ridge (Fig. 1.4) Aenigmatistes (most)
Frons not of this form 40
40. At least two differentiated dorsal or near-dorsal bristles in basal
two-thirds of mid tibia 41
Only one bristle or none in basal two thirds of mid tibia, but there may
be a longitudinal row of dorsal, differentiated hairs 78
41. Hind tibia with dorsal, longitudinal hair palisade (Fig. 8.4d,e).
Sometimes with two or three such palisades (Fig. 8.39f) 42
Hind tibia without longitudinal hair palisade, but there may be a row of
spaced hairs or a series of overlapping transverse combs 53
42. Vein 3 unforked 43
- Vein 3 forked 49
43. Arista absent Abaristophora
Arista present 44

224
 KEYS TO GENERA

44. Frons narrow, the height being clearly greater than width.
Trineurocephala (part)
- Frons with width equal to or greater than height 45
45. Mid tibia with one or two dorsal or near-dorsal longitudinal hair
palisades. Veins 1 and 3 divergent apically 46
- Mid tibia without longitudinal hair palisades. Veins 1 and 3 subparallel
throughout 47
46. Mid tibia with two longitudinal hair palisades. Supra-antennal bristles
absent. Anal tube relatively short Aenigmatistes (part)
- Mid tibia with only one longitudinal hair palisade. A well-developed
pair of supra-antennal bristles. Anal tube relatively long (Fig.
8.45a) Epicnemis
Note: The unknown male of one Diplonevra species may key out here,
but will be distinguished by its cerci being well separated from the epandrium
(see couplet 52), and by having both supra-antennals and two palisades.
47. Frontal bristles strongly developed. Form of hypopygium as Fig.
8.48e. (Typically palp as Fig. 8.48b and tip of vein 1 as Fig.
8.49a.) Hypocerides
- Frontal bristles greatly reduced (Figs 8.67a and 8.89a). Hypopygium
relatively short but high (e. g. Fig. 8.66c, d) 48
48. Palp with several bristles (Fig. 8.66b) and supra-antennal bristles
distinct (Fig. 8.67a). (Hypopygium as Fig. 8.66c,d, mid tibia as Fig.
8. 66a.) Palpiclavina
- Palp with only one bristle (Fig.8.89b) and supra-antennal bristles
vestigial or absent (Fig. 8.89a) Thaumatoxena
49. Frons lacks supra-antennals and other bristles at front. Base of broad
anal tube projects above level of dorsal face of epandrium. Scutellum
with at least ten bristly hairs Cyrtophorina (part)
- Supra-antennals and other bristles present at front offrons. Base of anal
tube not projecting above dorsal face of epandrium. Scutellum usually
with fewer than ten hairs, but when ten present then at least two clearly
differentiated as strong bristles 50
50. Costal cilia very reduced and coloured as costa, so that at low
magnifications latter appears to be bare Rhynchomicropteron (part)
- Black costal cilia clearly evident at low magnifications 51
51. Mesopleuron with hairs in upper part. Axillary ridge with single,
generally pale, bristle which is broadest in middle (Fig. 8.40b).
Hypopygium characterized by short but tall epandrium and hairless
sclerotized ring (the fused tergite and sternite 10) at base of anal tube
(Fig. 8.12c). (Hind femur typically with patch of spinules at base of
posterior face - Fig. 6.12, and sometimes with sensory pit as well- Fig.
8.40a, or other modifications) Dohrniphora
- Mesopleuron bare. Axillary ridge with at least two bristles, which are
generally dark brown or black and evenly tapered. Epandrium
generally longer than high and sclerites of segment 10 either form hairy
cylinder, with or without paired surstyli fused to the sides (Fig. 8.13),
or entirely absent 52
52. Sclerites of segment 10 form cylinder bearing hairs only (Fig. 8.13b) or

225
 IDENTIFICA TION

hairs and surstyli (Fig. 8. 13a). In either case bases of cerci are weB
separated from epandrium. (Bases of hind femora frequently with
spinules and/or processes - Fig. 8.39.) Diplonevra
- Sclerites of segment 10 not evident, so that bases of elongated cerci lie
immediately behind rear margin of epandrium Psyllomyia
53. Vein 3 unforked 54
- Vein 3 forked 68
54. Axillary ridge with single bristle which is very long plumose (Fig.
8.29a). (Tip of costa swollen - Fig. 8.2%. Hypopygium as Fig.
8.10.) Beyermyia
- Axillary ridge without such a long-plumose bristle, and frequently
with more than one bristle 55
55. No isolated bristles on upper two-thirds of hind tibia 56
- At least one bristle on upper two-thirds of hind tibia 57
56. Arista apical Billotia
- Arista in dorsal, pre-apical position 57
57. Mesopleuron with hairs 58
- Mesopleuron bare 61
58. Third antenna I segment globose 59
Third antenna I segment more elongated, being more pear-shaped 68
59. Hind tibia with series of overlapping transverse combs along length of
dorsal face (Fig. 8.46c). (Hypopygium as Fig. 8.48a.) Hypocera
- No such combs 60
60. Arista in dorsal, pre-apical position. Costa at least two-fifths of wing
length Plethysmochaeta
- Arista apical. Costa less than two-fifths of wing length Mannheimsia
61. Width of frons clearly less than height. Mid tibia with one or more true
dorsal bristles below one in basal pair of bristles Phora
- Width of frons subequal to or greater than height. Only true dorsal
bristle in upper two-thirds of mid tibia is one in basal pair of bristles 62
62. Hind tibia with two or more isolated bristles in upper two-thirds (e.g.
Fig. 8.32a) 63
- Hind tibia at most with only one isolated bristle in upper two-thirds 68
63. Veins 1 and 3 apically divergent (Fig. 8.31c). Arista apical in pos-
h~n M
Veins 1 and 3 subparallel (Fig. 8.65c). Arista dorsal in position.
(Hypopygium as Fig. 8.65d,e; hind leg as Fig. 8.65b.) Obscuriphora
64. Palps longer than breadth offrons, strongly tapered, devoid of bristles,
and directed upwards in front of broad hairless band running from
anterior ocellus to front offrons Bothroprosopa
Neither palps nor frons of this form 65
65. The most basal anterior bristle of hind tibia is paired with a dorsal
~~ ~
- Without dorsal bristle adjacent to most basal anterior bristle of hind
tibia 67
66. One pair of supra-antennal and eight other bristles on frons in front of
anterior ocellus. (Antenna as Fig. 8.32b, claspers (surstyli) of hypo-
pygium as Fig. 8.31b.) Conicera (part)

226
 KEYS TO GENERA

Both supra-antennals and antero-Iaterals absent, leaving only six

bristles in front of anterior ocellus Darwiniphora (part)
67. Apical ventral region of front metatarsus developed into an elongated
process 68
Front metatarsus simple Gymnoptera (part)
68. Costal cilia very reduced and coloured as costa, so that at low
magnifications latter appears to be bare Rhynchomicropteron (part)
Dark costal cilia clearly evident at low magnifications 69
69. Third antennal segment conspicuously elongated apically and tapered
from swollen base 70
Third antennal segment globose, pear-shaped or oval, but without
strongly tapered apical portion 71
70. Arista clearly pre-apical in position Cyphometopis (part)
Arista clearly apical 71
71. Front metatarsus with apical-ventral region developed into elongated
process. (Front tibia sometimes with strong spine, which may be
twisted, and/or notched) Coniceromyia (part)
Front metatarsus without such a process 72
72. Palps greatly elongated, being longer than breadth offrons, and devoid
of bristles Palpocrates
Palps shorter, or if enlarged there are small bristles at tip at least 73
73. Mesopleuron with hairs, and usually with strong bristle at rear as
well Chaetopleurophora
Mesopleuron bare 74
74. Medio-Iateral bristles absent Tarsocrates
Medio-Iateral bristles present (but antero-Iaterals sometimes reduced or
absent) 75
75. Mid tibia with anterior bristle in third quarter clearly longer than width
of tibia at point ofinsertion Spiniphora
Mid tibia with anterior bristle in third quarter greatly reduced or
absent 76
76. Fork of vein 3 complete Triphleba (part)
Base ofinner branch (R2+ 3) of'fork' of vein 3 obliterated, so there is no
actual forking of vein 3 as such 77
77. Restricted to New Zealand Kierania
Restricted to Holarctic Region Triphleba (part)
Note: See comment under review of Kierania below.
78. One differentiated bristle in basal half of mid tibia 79
No differentiated bristle in basal half of mid tibia, but there may be
longitudinal row of near-dorsal or dorsal hairs 85
79. Hind tibia with one or more longitudinal hair palisades 80
Hind tibia without longitudinal hair palisades 81
80. Hind tibia with only one, dorsal, hair palisade. Scutellum with at least
ten hairs Cyrtophorina (part)
Hind tibia with several longitudinal hair palisades (Fig. 8.60a).
Scutellum with only four hairs or bristles. (Frons and hypopygium as
Fig. 8.61, wing as Fig. 8.60d.) Multinevra (part)
81. Third antennal segment conspicuously elongated apically and tapered

227
 IDENTIFICA TION

from swollen base. Arista clearly pre-apical in position

Cyphometopis (part)
Third antennal segment otherwise. If tapered apically then arista is
apical; otherwise arista is dorsal 82
82. Vein 3 forked Triphleba (part)
- Vein 3 unforked 83
83. Arista dorsal, in pre-apical position. Subcosta well developed
Triphleba (part)
- Arista apical. Sub costa weakly developed or absent Xanionotum
84. Frons with antial and antero-Iateral bristles absent. A weak anterior
bristle in basal half of hind tibia Ceratoplatus
- Antial bristles present above the upper pair of supra- antennal bristles in
a pre-ocellar position. No isolated bristle in basal half of hind
tibia Ceratusa
85. Hind tibia with at least one dorsal, or near-dorsal, longitudinal hair
palisade (e.g. Fig. 8.4d,e) 86
- Hind tibia without longitudinal hair palisades, but there may be row of
spaced hairs 175
86. Front tibia with longitudinal dorsal, or near-dorsal, hair palisade 87
- Front tibia at most with spaced hairs or spinules along dorsal face 89
87. Hind tibia with several longitudinal hair palisades (Fig. 8.60a). (Frons
and hypopygium as Fig. 8.61; wing as Fig. 8.60d.) Multinevra (part)
Hind tibia with a single, dorsal, hair palisade 88
88. Vein 3 forked. Costa at most only halflength of wing, and ending level
with tip of vein 3 89
- Vein 3 unforked. Costa extends well past middle of wing and
overreaching tip of vein 3 Eremophora
89. Mesopleuron with hairs, and sometimes with one or more bristles also.
Rarely with a single strong bristle only 90
- Mesopleuron bare 109
90. Vein 3 unforked 91
- Vein 3 forked 95
91. Scutellum with at least six bristles. With six or more supra-antennal
bristles, but otherwise no bristles on frons between these bristles and
anterior ocellus Aenigmatopoeus
- Scutellum with only four bristles or two bristles and two hairs. With
only four supra-antennal bristles, but with several strong bristles
between these and anterior ocellus 92
92. Abdominal venter with paired pigmented patches on segment 5 (Fig.
8.71). (Front tarsus as Fig. 8. 72b.) Plastophorides
- Abdominal venter without these pigmented patches 93
93. Veins 1 and 3 subparallel (as Fig. 8.65c) 94
- Veins 1 and 3 apically divergent (as Fig. 8.31c) Megaselia (part)
94. Anal tube very short, with tip of proctiger only extending as far as rear
margin of epandrium Syneura (part)
- Anal tube long, extending well beyond rear of epandrium Lepta
Note: Since completion of this key, Brown (1993a) has published a
description of the hitherto unknown male of Co lye ria. It runs to this lead. The

228
 KEYS TO GENERA

anal tube extends beyond the rear margin of epandrium but is clearly shorter
than it. The mesopleuron I?as a single strong bristle only. The body and
hypopygium are somewhat flattened dorso-ventrally.
95. Eight bristles on scutellum 96
At most six bristles on scutellum, usually with four bristles or with two
bristles and two hairs 97
96. Epandrium with a pair of long surstylar lobes, extending from rear
margins below anal tube. Terminal hairs of proctiger clearly longer and
stronger than those on cerci Egregiophora
Epandrium without surstylar lobes. Hairs at tip of proctiger not longer
or stronger than those on cerci Johowia (part)
97. Hind tibia with antero-dorsal row of differentiated hairs, as well as
postero-dorsal row (the two rows being separated by hair palisade) 98
Hind tibia without an antero-dorsal row of differentiated hairs 100
98. Hind tibia with antero-ventral row of differentiated hairs as well
Physoptera (part)
Without this antero-ventral row of hairs 99
99. Palp bristles and supra-antennal bristles only about size of upper
occipitals, but the former sometimes a little longer
Phalacrotophora (part)
Either palp bristles or supra-antennals, or both, clearly longer than
upper occipitals Megaselia (part)
100. Frons narrow, with height clearly much greater than breadth 101
Frons not obviously narrowed or with breadth greater than height 102
101. Costa only extending about one-third of wing length. Third antennal
segment globose, but with pointed apex. Stenodicrania
Costa extends about half wing length. Third antennal segment about
twice as long as broad Ceratophoromyia
102. Frons about three times as broad as long. Third antenna I segment with
apex drawn out so that it is an elongated pear shape. Anal tube very long
and thin. Costa about halflength of wing Brachycephaloptera
Frons at most twice as broad as long. Without other characters in
combination 103
103. Anal tube clearly longer than length of dorsal face of epandrium.
(Mesopleuron with a long bristle at rear in addition to hairs.) 104
Anal tube little, if any, longer than dorsal face of epandrium 106
104. Section 1 of costa with a swelling. Only two minute supra-antennal
bristles. (Anal tube slender.) Pleurophorina
Costal section 1 without a swelling. With two or four supra-antennal
bristles, of which at least two are not obviously reduced in size 105
105. Anal tube slender and very long relative to length of epandrium (Fig.
8.S4a) Menozziola
Anal tube not so long and slender Megaselia (part)
and Rhyncophoromyia (part)
Note: These two genera can only be distinguished in the female sex.
Rhyncophoromyia is only known from the Nearctic and Neotropical Regions.
106. Third antennal segment elongated Macrocerides (part)
- Third antennal segment globose 107

229
 IDENTIFICATION

107. More than four supra-antennal bristles Johowia (part)

Only two or four supra-antennal bristles 108
108. Anal tube very short (both cerci and proctiger being only about as long
as high). Epandrium with well-developed paired surstylar lobes (at
least twice as long as width at base) which bear more conspicuous hairs
than rest ofepandrium.
Megaselia (part, including subgenus Kerophora)
Anal tube usually longer. Surstylar lobes absent or if developed they are
shorter and the hairs of rest of epandrium are usually more conspicu-
ous Diocophora and Megaselia (part)
Note: It is not clear how to separate these two genera in the male sex. The
hypopygium of a Diocophora is shown in Fig. 8.38(c). This genus is only
known from the Nearctic and Neotropical Regions.
109. Arista absent, but third antennal segment drawn out into a long,
tapered, apical extension 110
Arista present 112
110. Vein 3 forked. (The narrowed extension of third antennal segment at
least two thirds of total length. ) Maeroeerides (part)
Vein 3 unforked 111
111. Anal tube with inflated cerci (third antennal segment short-haired)
Tubieera (part)
Cerci not inflated (third antennal segment with at least some long hairs
in apical half) 218
112. Vein3unforked 113
Vein 3 forked 136
113. Two longitudinal hair palisades on hind tibia Distiehophora (part)
Only one palisade on hind tibia 114
114. More than four supra-antennal bristles 115
At most with four supra-antennal bristles 116
115. Part offrons bearing supra-antennals is extended forwards so that front
margin is almost V -shaped. Anal tube shorter than mid-line length of
dorsal face of epandrium Cataclinusa
Front margin of frons only imperceptibly produced in middle. Anal
tube longer than epandrium Antieojimbria
116. Anal tube at most as long as mid-line length of dorsal face of
epandrium 117
Anal tube longer than mid-line length of epandrium 120
117. Notopleural cleft present (Fig. 8.93b,c) Woodiphora (part)
- No notopleural cleft 118
118. Veins 1 and 3 almost subparallel, only slightly divergent apically 119
Veins 1 and 3 clearly divergent in apical third or more 120
119. Frons clearly broader than high. Third antenna I segment globose
Syneurina
Frons higher than broad. Third antenna! segment more oval
Syneura (part)
120. Third antennal segment globose 121
Third antennal segment more oval, conical, or at least always longer
than greatest breadth 124

230
 KEYS TO GENERA

121. Anal tube relatively short 122

- Anal tube relatively long (e.g. Fig. 8.25) 123
122. Last third of costal section 1 distinctly narrower than first two-
thirds Pheidolomyia
- Costal section 1 of even thickness throughout or gradl:lally thickening
from base Megaselia (part)
123. Anal tube with narrow cerci (Fig. 8.91b) (Costa about one-third of
wing length) Trucidiphora
Anal tube with greatly inflated cerci Dinocercus
Note: Since I completed this key, Brown (1993a) has published a
description of the hitherto unknown male of Borgmeieriphora. It runs to this
couplet. Its anal tube is similar to that of Trucidiphora, but its costa is at least
half the wing length and the tip of vein 4 is deflected rearwards close to wing
margin.
124. Entire third antennal segment long-haired, these hairs being much
longer than those on arista Dichocerina
Third antennal segment with any hairs longer than those on arista
restricted to apical region 125
125. Without supra-antennal bristles in a median position at front of
frons 126
At least two supra-antennal bristles in a median position at front of
frons 131
126. Antero-lateral and/or medio-lateral bristles absent 127
Both antero-lateral and medio-Iateral bristles present 128
127. Pre-ocellar and antial bristles also absent Melaloncha (part)
- Pre-ocellar and antial bristles present 130
128. A strong pair of supra-antennal bristles present: but each standing in a
triad with an antial and antero-lateral bristle and thus much further
apart than pre-ocellars (e.g. Fig. 8.27b) Auxanommatidia (part)
- Supra-antennal bristles entirely absent 129
129. Costa only about one-third of wing length. Hairs at tip of proctiger
much longer than those on cerci Cremersia (part)
- Costa about half wing length. Hairs at end of proctiger only about as
long as those on cerci. (Antennae close together.) Melaloncha (part)
Note: Since I completed this key, Brown (1993a) has published a
description of the hitherto unknown male of Melittophora. It runs to this
couplet. The costa is about half the wing length. The widely-separated
antennae and somewhat dorso-ventrally flattened body will distinguish it
from Melaloncha.
130. Antero-lateral bristles offrons absent Apocephalus (part)
Antero-lateral bristles present Melaloncha (part)
131. Antero-lateral and/or medio-lateral bristles offrons absent 132
- Both antero-lateral and medio-Iateral bristles present 133
132. Epandrium symmetrical and with broad, hairy, postero-Iateral
lobes Styletta
Epandrium tends to be asymmetrical and at least one postero-Iateral
lobe tends to be narrow and tapered (e. g. Fig. 8.25a) Apocephalus (part)
133. Anal tube relatively short and thick 134

231
 IDENTIFICA TION

Anal tube long and slender (e.g. Fig. 8.74a) 135

134. More than four bristles on axillary ridge of wing Apodicrania
Fewer than three axillary bristles 173
135. Frons more than 1.5 times as broad as high Veruanus
Frons less than 1.5 times as broad as high Pseudacteon
136. With at least eight bristles on scutellum Chaetaspidia
With at most only six bristles on scutellum 137
137. Hind tibia with more than one longitudinal hair palisade, even if some
are irregular (e.g. Fig. 8. 92c,d and 8. 73a) 138
- Hind tibia with a single, dorsal, hair palisade 141
138. Mesopleuron with hairs Beckerina (part)
Mesopleuron bare 139
139. At anterior end of no top leuron the orifice of the duct from notopleural
gland is fully exposed and its diameter is less than diameter of anterior
thoracic spiracle (Fig. 8.28b) Beckerina (part)
Orifice of duct from notopleural gland at least partly concealed within a
notopleural cleft (Fig. 8. 93b, c). The length of this cleft is greater than
diameter of anterior thoracic spiracle 140
140 Hind tibia as Fig. 8.73a. (Hypopygium as Fig. 8.70b) Plectanocnema
Hind tibia with fewer and less complex hair palisades (as Fig.
8.92c,d) Woodiphora (part)
141. Postero-dorsal hairs of hind tibia set in enlarged asymmetric pits and
with broadened tips. Wings with a distinct elongated dark mark
between bases of veins 5 and 7. Costa extends well beyond apex of vein
3. Pelidnophora
Without this combination 142
142. At least six supra-antennal bnstles Johowia (part)
At most only four supra-antennal bristles 143
143. Six short bristles on scutellum. With four subequal supra-antennal
bristles and eight other bristles on frons below pre-ocellars. Costa
extending almost two-thirds of wing length Clinochaeta
Without this combination 144
144. Frons breadth more than five times mid-line height, the anterior ocellus
being at the front margin. Third antennal segment concave on external
face and with greatest breadth about the same as frons width
Lecanocerus
Frons not so broad. Third antennal segment otherwise 145
145. Outer two-thirds of hind femur postero-ventrally hollowed to accom-
modate tibia, and this region with a distinctive oblique dark mark at its
proximal end (Fig.8. 74d). A pre-apical tooth present on all tarsal claws
(Fig. 8.74b). Anal tube very short. More than eight bristles on axillary
ridge of wing Pseudohypocera
Without this combination 146
146. Basal half of hind margin of wing fringed with fine hairs a little longer
than costal cilia on costal section 3. Costa more than half wing length.
Third antennal segment globose. Anal tube clearly much shorter than
epandrium Woodiphorides
Not with this combination 147

232
 KEYS TO GENERA

147. Hind femur with triangular projection at base (Fig. 8.78c). Anal tube
shorter than epandrium (Fig. 8.78d). Third antenna I segment clearly
longer than broad (Fig. 8.78e) Razorjemora
- With a different combination 148
148. Eye with lower facets (ommatidia) at least twice diameter of those
uppermost (e.g. Fig. 8.27b). Anal tube long relative to epandrium (e.g.
Fig. 8.27a). Third antennal segment longer than greatest breadth
Auxanommatidia (part)
- Without these features combined 149
149. Supra-antennal bristles absent, but antials may be in middle of front
margin offrons 150
- At least two small supra-antennal bristles are differentiated from
adjacent hairs on frons 152
150. Anal tube clearly longer than epandrium and slender 151
- Anal tube either shorter than mid-line length of epandrium or if a little
longer it is not slender. (Third antenna I segment oval or pear-
shaped.) Neodohrniphora and Cremersia (part)
Note: The males of both genera are poorly known. The known
Neodohrniphora have the costa at least two-fifths of wing length, whereas it is
only about one-third oflength in the Cremersia species keying out here.
151. Pre-ocellar bristles absent. (Costa about half of wing length.)
Melaloncha (part)
- Pre-ocellar bristles present 152
152. The last quarter of vein 6 (third thin vein) absent. Frons more than 2.5
times as broad as high. Antial bristles close to antero-Iaterals. Third
antennal segment globose, but with small apical point Exochaeta
- Without this combination 153
153. Hind tibia with a clearly differentiated antero-dorsal row, as well as a
postero-dorsal row, of hairs (the two rows separated by hair pali-
sade) 154
- At most with a postero-dorsal row of differentiated hairs on dorsal face
of hind tibia 155
154. Palp bristles and supra-antennals only about size of upper occipitals, but
former sometimes slightly longer Phalacrotophora (part)
- Palp bristles and at least upper pair of supra-antennals clearly longer
than upper occipitals 155
155. Third antennal segment globose 156
- Length of third antennal segment clearly greater than greatest
breadth 162
156. At front end of notopleuron there is the opening of duct from
notopleural gland, either fully exposed (Fig. 8.28b) or concealed (in
part at least) within a notopleural cleft (Fig. 8. 93b, c). (Anal tube at most
only about as long as epandrium.) 157
- No such opening, or cleft, at front of no top leur on 158
157. Opening of duct from notopleural gland at least partly concealed within
a notopleural cleft (Fig. 8.93b,c). Length of this cleft is greater than
diameter of anterior thoracic spiracle Woodiphora (part)
- Opening of duct from notopleuron gland fully exposed to view and its

233
 IDENTIFICA TION

diameter is less than that of anterior spiracle (Fig. 8. 28b)

Beckerina (part)
158. Costa at least half wing length 159
- Costa less than half wing length Megaselia (part)
159. Anal tube at most as long as mid-line length ofepandrium 160
- Anal tube clearly longer than epandrium 161
160. Costal at most only half wing length Megaselia (part)
Costa more than half wing length Megaselia (part)
and Trophithauma (part)
Note: These two genera can only be separated in the female sex; indeed
the latter may only be aberrant species of Megaselia.
161. With at most only three bristles on axillary ridge of wing
Megaselia (part)
With at least four axillary bristles Megaselia (part),
Physoptera (part)
and Rhyncophoromyia (part)
Note: The males of these three genera are difficult to separate at the
generic level. The last two are restricted to the Nearctic and Neotropical
Regions, and both have obviously elongated anal tubes.
162. Anal tube at most as long as mid-line length ofepandrium 163
Anal tube clearly longer than epandrium (e. g. Fig. 8.25) 166
163. Costa extends less than half wing length 164
Costa extends more than half wing length 165
164. Inner branch of fork of vein 3 either disconnected at base or clearly
much thinner than outer branch (R4+5). Third antennal segment
oval Syneura (part)
- Inner branch of fork of vein 3 intact and strong, even if a little thinner
than outer branch. Third antennal segment oval or pear-shaped
Myrmosicarius (part)
165. Third antennal segment bean-shaped. Only two supra-antennal
bristles Diaclinella
- Third antennal segment more nearly globose. With four supra-
antennals Megaselia (part)
166. Costa extends to middle of wing or beyond 167
Costa less than half wing length 171
167. With at least three bristles on axillary ridge of wing 168
With fewer than three axillary bristles 170
168. Third antennal segment conical (tapering towards apex) 169
- Third antennal segment more oval Megaselia (part)
169. Conical third antennal segment with long, tapered, apical portion.
With four supra-antennal bristles. With at least four axillary
bristles Macrocerides (part)
- Without this combination Apocephalus (part)
170. Vein Sc obscure. Third antennal segment pear-shaped. Only two
supra-antennal bristles. Antial and antero-Iateral bristles
present Anaclinusa
- Without these characters in combination Apocephalus (part)
171. Anal tube relatively long and slender (e.g. Fig. 8.25) 172

234
 KEYS TO GENERA

Anal tube relatively short and broad (e. g. Fig. 8. 57b) 173
172. Supra-antennal bristles as strong as antero-Iaterals Neophora
Supra-antennal bristles either absent or clearly shorter than antero-
laterals or, iflatter are absent, than antials Apocephalus (part) and
Dacnophora
Note: It is not clear how to distinguish these two genera in the male
sex.
173. Epandrium longer than high and postero-Iaterallobes well developed
below anal tube (e. g. Fig. 8. 57b) Microselia (part)
Epandrium as high as high as long, or a little higher, without obvious
postero-Iaterallobes 174
174. Vein 3 forked Synclinusa
Vein 3 unforked Pradea
175. Arista absent, but apex of third antennal segment drawn out into a long
pseudo-arista (e. g. Fig. 8.36a) 176
Arista present and articulated with third antennal segment 177
176. Both hairs and bristles on frons, in front of anterior ocellus Rhopica
(part)
Only hairs on frons, in front of anterior ocellus Dahliphora (part)
177 . Vein 3 forked 178
Vein 3 unforked 187
178. Mesopleuron with fine hairs in postero-dorsal region and notopleuron
with a cleft (Fig. 8.47a,b). Frons with only small hairs and no bristles in
front of ocelli. Costal cilia only about as long as breadth of costal section
3 and costal section 1 clearly longer than sections 2+ 3 Gymnophora
Without this combination and mesopleuron always bare 179
179. Third antennal segment with elongated, tapered, apical part 180
Third antennal segment essentially globose, even if a little pointed
apically 181
180. Frons much wider than high Corynusa
Frons exceptionally narrow, with height very much greater than
breadth Cyphometopis (part)
181. The bend in basal third of vein 5 is opposed to forward deflection in
middle of vein 6, so that space between the two veins is character-
istically narrowed just before middle (as in Fig. 8.56a) Metopina (part)
Without this feature 182
182. Apart from supra-antennals, there are fewer than eight bristles in front
of pair on ocellar triangle. (Frons clearly broader than high.) 183
The normal eight bristles are present on frons in front of ocellars (as in
Fig. 8.1a) 184
183. Medio-Iateral bristles absent Macroselia
Medio-Iateral bristles present, even if somewhat higher on frons than
usual Aphiura
184. Costa clearly extends a little way beyond tip of vein 3 Allochaeta
Costa ends at tip of vein 3 185
185. Vein Sc obscure or absent Phymatopterella (part)
Vein Sc strongly developed and its tip merging with vein 1 186
186. Opening of duct from notopleural gland fully exposed to view and its

235
 IDENTIFICATION

diameter less than diameter of the anterior thoracic spiracle (Fig.

8. 28b) Beckerina (part)
- Opening of duct from notopleural gland at least partly concealed in a
notopleural cleft (Fig. 8.93b,c), the length of latter being greater than
diameter of anterior thoracic spiracle Woodiphora (part)
187. Eyes greatly enlarged and almost meeting behind ocelli, and frons
narrow (Figs 8.70c and 8. 73b). Wing with very short costa and weakly
developed vein 1 (Fig. 8.70d) Postoptica
- Without this combination 188
188. Front tibia with apical ventral process and/or pre-apical triangular
notch in ventral margin Apterophora
- Front tibia without apical ventral process or ventral notch 189
189. Front metatarsus with notch in middle of ventral margin. Scutellum
with two bristles and four hairs (two between the bristles and one either
side) Eurycnemis
- Without these features 190
190. Axillary ridge of wing with at least one bristle 191
- Axillary ridge of wing without bristles 195
191. Sub costa strongly developed. Anal tube at most subequal to length of
epandrium, usually clearly shorter 192
- Sub costa obscure or absent. Anal tube clearly longer than
epandrium Phymatopterella (part)
192. With the posterior pair of bristles on scutellum shorter than the anterior
pair. (Frons about twice as broad as long.) Paraphiura
The four scutellar bristles either sub equal or the anterior pair shorter,
and occasionally reduced to fine hairs 193
193. Front tarsi modified as in Fig. 8. 92a, b Woodiphora (part)
Front tarsi not modified in this way 194
194. Third antennal segment oval. Right side of epandrium with tapered
lobe from hind margin extending almost as far as tip of anal tube, which
is about as long as epandrium. With four bristles on ~cutellum
Distichophora (part)
- Third antennal segment globose. Epandrium without tapered lobe
from hind margin. If anal tube is almost as long as epandrium then
scutellum with posterior pair of bristles and anterior pair of minute
hairs Woodiphora (part)
195. Wings somewhat reduced (Fig. 8. 59a, b). Posterior ocelli absent. No
frontal bristles apart from one reduced pair of supra-antennals and
o~~~ ~~~~~
- Without this combination 196
196. Front femur with at least six long, subventral, hairs in basal half (these
hairs at least as long as greatest breadth offront tibia) Homalophora
- Without these long hairs in basal half offront femur 197
197. Front femur somewhat swollen and with a darkened patch just beyond
middle (Fig. 8.53a, b). Costa clearly less than half wing length and veins
5 and 6 much weaker than veins 4 and 7 (Fig. 8.53c,d). (Hypopygium as
Fig. 8.11a, b.) Maculiphora
- Without these features in combination 198

236
 KEYS TO GENERA

198. Anal lobe of wing large, so that inner margin ofleft wing viewed from
above is strongly S-shaped. Vein 1 with at least distal half missing.
Costa only about two-fifths of wing length. Third antennal segment
conical, with an apical arista. Both antero-lateral and medio-lateral
bristles offrons absent Travassophora
- Not with this combination 199
199. Costa at most only one-third of wing length. Anal tube clearly shorter
than epandrium 200
- Costa at least two-fifths of wing length. Anal tube shorter or longer
than epandrium 208
200. Vein 1 distinct. Vein 3 straight and ending at tip of costa. Vein 4
originates well beyond tip of vein 3. Veins 4-6 end very close to wing
margin Brachycosta
- Without this combination 201
201. Arista long-haired (Figs 8. 76b and 8. 84b) 202
- Arista short-haired (e.g. Fig. 8.63a) or a little longer (Fig. 8.84c) 203
202. Second segment of arista longer than broad (Fig. 8.84b). Veins 4-6 end
very close to wing margin (Fig. 8.85b) Termitophilomyia
- Second segment of arista broader than long (Fig. 8. 76b). Veins 4-6 end
well clear of wing margin (Fig. 8. 76a). (Front leg as Fig. 8. 76e.)
Ptochomyia
203. Vein 4 originates beyond apex of vein 3 (e.g. Fig. 8.82a,d) 204
- Vein 4 originates at or before apex of of vein 3 (e.g. Figs 8.63c and
8.88a) 205
204. A clearly differentiated pair of supra-antennal bristles present. Vein 1
distinct. Veins 4-6 almost reach wing margin. A faint vein 7
present Brachyphlebina
- Any hairs in position of supra-antennals are at most only as strong as
other small hairs on frons. Vein 1 obscure, veins 4-6 ending somewhat
to considerably short of wing margin, and no indication at all of vein 7
(Fig.8.82a,d) Syntermitoxenia
205. Left surstylus withjagged antero-ventral margin (Fig. 8.33a) Perissa
- Left surstylus with smooth, or only slightly irregular, antero-ventral
margin (Figs 8.87c and 8.88b) 206
206. Vein 5 directed towards wing tip (Fig. 8. 88a). Second segment of arista
with breadth greater than or equal to length (Fig. 8.88c). (Hypopygium
as Fig. 8.88b.) Javanoxenia
Vein 5 directed to point behind wing tip (Fig. 8.87d). Second segment
of arista longer than broad (Figs 8. 63a and 8. 84c) 207
207. Fore tarsal segments relatively long (Fig. 8.63b). With two fine bristles
in lower half of propleuron. (Antenna as Fig. 8.63a, wing as Fig. 8.63c,
hypopygium as Fig. 8.69a.) Necperissa
- Fore tarsal segments relatively short (Fig. 8.84d). Propleuron bare.
(Hypopygium as Fig. 8.87e.) Termitoxenia
208. Outer half of costal section 1 with sudden swelling Pachyneurella
- Costal section 1 either of uniform thickness throughout or only
gradually increasing in breadth 209
209. Ocelli absent. (Vein Sc strongly developed and its tip Jommg

237
 IDENTIFICA TION

R1.) Wandolleckia
Ocelli present 210
210. Anal tube clearly shorter than epandrium and cerci little, if any, longer
than high (e.g. Fig. 8.30.) 211
Anal tube subequal or longer than epandrium. When a little shorter,
then cerci are clearly longer than high 214
211. Vein 6 (third thin vein) deflected forwards in middle third, or just
beyond, so that space between veins 5 and 6 is a little to very markedly
(Fig. 8.56a) constricted. The microsetae fringing the rear margin of
wing only about as long as the distance between adjacent
microsetae 212
Veins 5 and 6 almost straight or only slightly sinuous, so there is no
obvious constriction of space between these two veins (e. g. Fig. 8. 29c).
The microsetae fringing rear margin of wing about three times as long
as distance between adjacent microsetae 213
212. Vein 6 strongly deflected and anal region (i.e. part of wing bearing anal
vein - vein 7) with inner border at a steep angle to axillary ridge (Fig.
8.56a) Metopina (part)
Vein 6less strongly deflected and anal/ axillary angle shallow
Penthaplus
213. Wing as Fig. 8.29(c), the base ofRsjoining base of vein 1 at an oblique
angle. (Hypopygium compact and often complex - Figs 8.15a and
8.30.) Chonocephalus
Wing with base of Rs joining base of vein 1 at an angle of about
90 Epichonocephalus
214. Vein Sc strongly developed and its tip fusing with vein 1 (as in Fig.
8.5b) 215
Vein Sc less strongly developed, and its tip not reaching vein 1, or
absent 217
215. Anal tube a little shorter than mid-line length of epandrium. Lower
margin ofleft side of epandrium concavely emarginate in middle. Palps
large (breadth being greater than diameter of third antennal segment)
with only short fine hairs. At most only pre-ocellar bristles on frons
between supra-antennals and ocelli Trophodeinus
Without this combination 216
216. Both antial and antero-Iateral bristles absent Pre-ocellars and medio-
laterals present. (Hypopygium as Fig. 8.54b,c, wing as Fig. 8.59d,
front leg as Fig. 8. 59d.) Mesopathusa
Antero-Iateral and/ or medio-Iateral bristles absent. Antials
present Commoptera and Puliciphora
Note: It is not clear how to distinguish these two genera in the male sex.
Borgmeier (1926a) erroneously stated that the Sc vein is absent in Commop-
tera, despite it being illustrated as being present. The error was then
perpetuated in his Nearctic monograph (Borgmeier, 1963d).
217. Costa just under half wing length. Vein Sc evident, and sometimes with
1-2 small bristles at base Acontistoptera
Costa extending just over half wing length. Vein Sc without bristles
and frequently absent altogether Ecitomyia,

238
 KEYS TO GENERA

Ecitophora,
Ecitoptera
and Ecituncula
Note: These genera are only reliably separated in the female sex, in our
present state of knowledge.
218. Vein 4 runs straight to wing margin and vein 3 has a slight angle near
origin of vein 4 Pseudacteon (part)
Vein 4 almost reaches wing margin but then turns sharply rearwards
and gradually approaches wing margin at a narrow angle. Vein 3
straight in vicinity of origin of vein 4 Trichocerina

(C) KEY TO GENERA OF FEMALES

1. Wings and halteres entirely absent (e.g. Figs 6.7, 8.20b, 8.22a).
Sometimes wing rudiments fused to thorax and not articulated with it
(e. g. Fig. 8.44a) 2
Wings present, even if reduced to rudiments or stumps (e.g. Figs
6.4-6.6, 8.26b, 8.SSa, 8.77a, 8.81d and 8.90). These rudiments
articulate with thorax. Halteres usually present 47
Note: In some cases the wing rudiments are partly fused to the thorax.
Sometimes developments of the anterior segments of the abdomen may tend
to conceal wing stumps (e.g. Fig. 8.46a). These cases are keyed both ways.
2. End segments of abdomen directed rearwards 3
End segments of abdomen deflected ventrally, even to the extent of the
anus being directed forwards (e.g. Fig. 5.1) 87
3. Dorsal-basal region of abdomen extended forwards over top of thorax
(Fig. 8.46a). First antennal segment as long as tapered third segment
(Fig.8.46b) Eutermiphora (part)
Without this combination 4
4. Proboscis more than four times length of head, with distal section
elongated. (Abdominal tergites 1-6 present, but 6 reduced, and 5 with
antero-median gland complex. Only two ocelli present, the anterior
one being absent.) Apterophora
Proboscis little, if any, longer than head and usually shorter 5
5. Abdominal tergites entirely absent 6
At least tergite 2 is present, even if weakly pigmented 9
Note: Variable genera are keyed both ways.
6. A pair of bristles in basal half of mid tibia. Three ocelli present
Diplonevra (part)
Mid tibia without pair of bristles in basal half, but there may be some
long hairs along dorsal face. Ocelli absent 7
7. Diameter of eye clearly less than diameter of third antennal segment 8
Diameter of eye sub equal to diameter of third antennal segment 9
8. All tibiae bearing dorsal, or near-dorsal, hairs which are longer than
greatest breadth of tibia. Similar hairs on femora and most tarsal
segments. Dorsal faces of abdominal segments 2-5 also with conspicu-
ous hairs Oligophora

239
 IDENTIFICATION

Hairing oflegs and abdomen much shorter 9

9. Abdominal tergites greatly reduced in number, with only 2 (which is
sometimes very pale), and sometimes 5 as well, present 10
Abdominal tergites 2-5 present, and sometimes 1 and 6 also 19
10. Ocelli present 11
Ocelli absent 12
11. Thorax (viewed from above) more than twice as wide as long.
Abdomen with no trace of tergite or associated gland opening on
segment 5 Apterella
Thorax at most only twice as wide as long. Abdominal segment 5 with
a reduced tergite associated with a gland opening 12
12. Diameter of eye clearly less than diameter of third antennal segment 13
Diameter of eye subequal to, or greater than, diameter of third antennal
segment 16
13. Abdominal tergite 2 absent Trophodeinus (part)
Abdominal tergite 2 present 14
14. Abdominal tergite 2 with front margin broader than hind margin
Trophodeinus (part)
Abdominal tergite 2 with hind margin broader than front margin 15
15. Hairs on dorsal faces of abdominal segments 1-5 short, being clearly
shorter than bristles on thorax. Sometimes with long hairs on flanks of
segments 3 and 4 Ecituncula (part)
Hairs of dorsal faces of abdominal segments 1-5 long, some being
almost as long as bristles on thorax, contrasting with short hairs on
tergite 2 (apart from longer ones on hind margin) Rhabdomyia (part)
16. Abdominal tergite 2 extends full width of segment, but sometimes
largely represented by patch of longer hairs contrasting with shorter
hairs on rest of abdominal dorsal face 17
Abdominal tergite 2 reduced, and clearly not extending full width of
segment 18
17. Abdominal tergite 2 mainly represented by longer hairs contrasting
with rest of hairs on dorsal face of abdomen. No gland complex evident
on segment 5 Morahania
Abdominal tergite 2 distinctly demarcated and bearing short hairs (no
longer than hairs, as opposed to bristles, on thorax). A distinct gland
opening on segment 5 Zikania
18. Abdominal tergite 2 at most as broad as long, and usually longer than
broad. General hairing of dorsal face of abdomen shorter than hairs on
tarsi Ecituncula (part)
Abdominal tergite 2 more than twice as wide as long. General hairing of
dorsal face of abdomen clearly longer than hairs of tarsi Wandolleckia
19. Not strongly dorso-ventrally flattened; the anterior thoracic spiracles in
normal, lateral, position; head usually not very much wider than long;
antennae usually closely approximated; the postero-Iateral corners of
thoracic notum not forming backwardly-directed tapered processes 20
Strongly dorso-ventrally flattened; head very much wider than long;
antennae widely separated; anterior thoracic spiracle in dorsal position;
postero-Iateral extremities of thoracic notum may form tapered

240
 KEYS TO GENERA

processes (representing wing rudiments fused to notum) (e.g. Fig.

8.44) 54
20. Hind femur with dorsal, pre-apical, feathered bristle and all tibiae with
similar dorsal or near-dorsal bristles (Fig. 8.19). Antennae widely
separated and frons with long hairs (Fig. 8.20a). Thorax and abdominal
tergites also long-haired (Fig. 8. 20b) Aenictacantha
Without these features in combination 21
21. Both eyes and ocelli absent. Third antennal segment pear-shaped.
Abdominal tergites 2 and 3 short but wide, with long bristles on their
concave hind margins. Long bristles along dorsal faces of mid and hind
tibiae Cryptophora
Without this combination 22
22. No supra-antennal bristles in middle at front of frons, as they are
displaced laterally in association with the wide separation of antennae
(e.g. Fig. 6.7). Apart from bristles on vertex, which are either reduced
or absent, there are no other bristles on frons 23
Supra-antennal bristles, usually two pairs, situated in middle at front of
frons. Antennae usually more closely approximated. There may be
other bristles on frons, apart from supra-antennals and those on
vertex 25
23. Abdomen with six tergites extending full widths of segments (Fig. 6.7).
(Terminal abdominal segments as Fig. 8. 8a, b) Chonocephalus
Abdomen with at most only five tergites extending full widths of
segments 24
24. Only three abdominal tergites extending full widths of segments.
(Apart from antero-Iaterally situated supra-antennals, with no bristles
on frons. Ocelli absent. Eyes much reduced. Long bristles on flanks of
abdominal segments 3-6.) Paurophora
Abdomen with five tergites extending full widths of segments 25
25. Dorsal face of thorax at least four times as wide as long (e.g. Fig.
8.55d) 26
Dorsal face of thorax less than four times as wide as long (but if almost
four times as wide as long, then arista is shorter than palp) 28
26. At least four abdominal tergites extending full widths of segments (e.g.
tergites 4-6 as Figs 6.14 and 8. 55b, c) 27
Abdominal tergites 3-5 progressively much narrower than widths of
segments Stethopathusa
27. At rear of abdominal tergites 2-5 at least some of the bristles are as
strong as those on thorax (e. g. Fig. 8.55c) Mesopathusa (part)
Bristles at rear of abdominal tergites 2-5 not as strong as those on
thorax Puliciphora (part)
28. Abdominal tergites 1 and 2 tapered laterally and extending well down
flanks (Fig. 8.4la) Dundophora
Abdominal tergites 1 and 2 not tapered in this way 29
29. A pair of ocellar bristles present (i.e. a pair of closely approximated
bristles in middle of vertex) even though ocelli may be absent. If these
bristles are somewhat small, then diameter of eye at least equal to that of
third antennal segment 30

241
 IDENTIFICA TION

Ocellar bristles absent. (Ocelli absent; antennae widely separated;

dorsal face of thorax more than three times as wide as long; four
abdominal tergites as wide as segments; diameter of eye less than that of
third antennal segment; arista shorter than palp.) Tranopeltoxenos
30. Ocelli absent 31
Ocelli present 40
Note: Variable genera are keyed both ways.
31. At least four bristles on frons between supra-antennals and those on
vertex 32
At most only two bristles on frons between supra-antennals and those
on vertex 33
32. Abdominal tergites 2-4 each with two transverse rows of strong
bristles (as strong as those on thorax), one row in anterior half and one
at rear margin. Tergite 5 with a large semicircular flap occupying a wide
emargination of the front margin (similar to Fig. 6.14 right)
Hexacantherophora
Abdominal tergites 2-4 each with at most a single transverse row of,
weaker, bristles at rear margin only. Tergite 5 with only a small
emargination of middle of front margin (less than one third of tergite
width) and without a semicircular flap Aptinandria
Note: Since I completed this key, Brown (1993a) has described new
species of Borgmeieriphora whose females will run to this lead (rather than
couplet 69), as they lack wing rudiments. Their down-curved, pointed
ovipositors, with cerci vestigial or absent, will distinguish them from
Aptinandria. If none of these options apply, proceed to couplet 40.
33. Eyes greatly reduced, their diameters being at most only one-third of
that of third antennal segment 34
Diameter of an eye at least two-thirds, and usually greater than, that of
third antennal segment 36
34. Frons without bristles between supra-antennals and those on vertex.
Without any indication of a gland opening in middle of front margin of
abdominal tergite 5 Syntermophora
Frons with pair of bristles between supra-antennals and those on
vertex. A small semicircular flap occupying indentation in middle of
front margin of abdominal tergite 5 (e. g. Fig. 8. 78a, b) 35
35. Abdominal tergite 4 much wider than abdominal tergite 5 (e.g. Fig.
8.78a) Puliciphora (part)
Abdominal tergite 4 scarcely any wider than tergite 5. 36
36. Abdominal tergite 4 only as wide as tergite 5. Frons with small pair of
antial bristles, four supra-antennals, and pair of ocellars on vertex
Hylophora
Abdominal tergite 4 wider than tergite 5. With no bristles on frons
between supra-antennals and those on vertex 37
37. Abdominal tergites 2-6 extend full widths of segments 38
At least tergite 5 is much narrower than width of segment (e.g. Fig.
8.78b) Puliciphora (part)
38. Abdominal tergite 5 with semicircular flap (associated with a gland
opening), which may be small (e.g. Fig. 8.78a,b), in middle affront

242
 KEYS TO GENERA

marglll 39
No such flap (or gland opening) at front of abdominal tergite
5 Coridophora (part)
39. Anterior flap at front of abdominal tergite 5 less than one quarter of
width of tergite (e. g. Fig. 8. 78b) Puliciphora (part)
Anterior flap at front of abdominal tergite 5 more than half width of
tergite (Fig. 8.18a) Adenophora
40. Frons without bristles between supra-antennals and those on vertex 41
Frons with at least two bristles between supra-antennals and those on
vertex. (With gland complex in middle of front margin of abdominal
tergite5,e.g. Fig. 6.14.) 44
41. Gland opening in middle of front margin of abdominal tergite 5,
usually associated with small semicircular flap (e. g. Fig. 6.14)
Pachyneurella
and Puliciphora (part)
Note: These two genera can only be separated in the male sex, in our
present state of knowledge).
No such gland complex at front of abdominal tergite 5 42
42. Top of thorax with only four strong bristles near rear margin, there
being none on middle third Coridophora (part)
With six strong bristles near rear of thorax, there being a pair in middle
third 43
43. Viewed from above, back of head concave Homalophora
Back of head not concave Penthaplus
44. Top of thorax with 4-6 bristles near hind margin 45
Top of thorax with only a single pair of bristles at rear Placophorina
45. Eyes greatly reduced, their greatest diameter being less than half that of
third antennal segment Arrenaptenus
Eyes with greatest diam~ter clearly more than half that of third antennal
segment 46
46. Viewed from above, diameter of eye at most only slightly greater than
that of third antennal segment, usually subequal or less. There is a
bristle, sometimes on a distinct papilla, on side of thorax towards rear
(representing the last vestige of the ancestral wing). (Abdominal
tergites as Fig. 6.14.) Puliciphora (part)
Viewed from above, diameter of eye clearly greater than that of third
antennal segment. There is no bristle on side of thorax towards rear (i. e.
in the position that would have been occupied by the wing in the
ancestral state) Myrmomyia
47. Wings with at least membranous area (part bearing the thin veins)
reduced 48
Wings fully developed (e.g. Fig. 1.1) 84
Note: Borderline cases, where the membranous area is only reduced a
little (e.g. Fig. 6.2b) are keyed both ways.
48. Wings represented by intact, but reduced, wings or rudiments (e.g.
Figs6.2c,d, 6.3-6.6, 8.26, 8.42d, 8.52a, 8.55a, 8.79d and 8.90) 49
Wings represented by stumps, the membranous area having been
largely shed, leaving a ragged outer margin (e.g. Figs 6.1,8. 77a) 84

243
 IDENTIFICATION

49. Wing membrane greatly narrowed, but not obviously shortened (e.g.
Fig. 6.3) 50
- Wing otherwise 51
50. Costa narrow (no wider than vein 3) with at most two rows of costal
cilia, which are clearly longer than width of costa in outer half. Veins
4-6 (first three thin veins) evident. The tergite rudiments associated
with gland opening on abdominal segment 5 relatively small
Chelidophora
- Costa broad, with at least three irregular rows of hairs which are
relatively short. Veins 4-6 not evident (e.g. Fig 6.3). Semicircular flap
associated with gland opening on abdominal segment 5 relatively large
(Fig.8.45b) Franssenia
51. Wing rudimentary in that the membrane (i.e. part beyond end of costa
and behind vein 3) is greatly reduced or absent. When some of
membrane present costal index exceeds 0.9 52
- Wing complete, even if membrane is shorter than usual. Costal index
always less than 0.9 84
52. Strongly dorso-ventrally flattened and abdominal segments 1-6 largely
covered by single large tergite; antennae in pits (Figs 8.26 and 8.90) 53
- If strongly dorso-ventrally flattened, then abdominal tergites 1-6
largely covered by six separate tergites or 3-6 reduced so that much of
dorsal face is membranous. Antennae usually clearly not in pits 54
53. As Fig. 8.90, with head very much broader than long and proboscis
shorter than head. Hind femur with at least one (usually two) pre-apical
anterior spines (Fig. 8.91 a) Thaumatoxena
- Head and thorax as Fig. 8.26, with proboscis longer than head. Hind
femur without pre-apical anterior spines Assmutherium
54. Six tergites evident and entirely covering dorsal face of abdomen. Frons
with anterior face at right angle to top face (Fig. 8.23b,c) and sometimes
latter greatly shortened (Fig. 8.24). Wing rudiments simple
(Fig. 6.4) Aenigmatistes
- Without this combination 55
55. Without wing rudiments or they are fused to postero-Iateral corners of
thorax, and do not articulate with it (e.g. Figs 8.22a and 8.44) 56
- Wing rudiments present, and articulating with thorax 58
56. Diameter of eye greater than that of third antennal segment (Fig. 8.22a).
At rear of abdominal segment 7 with a series of strong hooks each side
(Fig. 8.21 b,c) (Hind tibia as Fig. 8.60b.) Aenigmatias
- Diameter of eye less than that of third antennal segment. No strong
hooks at rear of abdominal segment 7 57
57. Palps globose (Fig. 8.44b). Mid femur with antero-ventral row of
strong, dark, spine-like bristles in outer half (Fig. 8. 43b, c) Epicnemis
- Palps flattened (Fig. 8.23a). Antero-ventral row of bristles on outer half
of mid tibia weaker and no darker than adjacent hairs Aenigmatopoeus
58. Wing rudiment with very long, strong, feathered bristles on costa (e.g.
Figs 8.15b and 8.17b) 59
- No such bristles on costa 61
59. Wing with radial veins distinct and thin veins also evident (Fig. 8.17b).

244
 KEYS TO GENERA

Normal halteres present. Ocelli present Adelopteromyia (part)

Radial veins obscure and thin veins not discernible (e.g. Fig. 8.15b).
Normal halteres may be present, but usually strongly reduced or
absent. Ocelli absent 60
60. Frons with only four supra-antennal bristles, arranged two in front and
two behind, at front of frons between moderately separated antennae.
Scutellum with pair of hairs, weaker than pre-scutellar dorso-central
bristles, or bare Xanionotum
- Frons with at least six bristles at front, arranged in a single transverse
row, between widely separated antennae. Scutellum with pair of
bristles at least as strong as pre-scutellar pair of dorso-centrals
Acontistoptera
61. Wing rudiment with costa and radial veins distinct 62
Wing rudiment more reduced, the radial vems (at least) being not
discernible 67
Note: Variable cases are keyed both ways.
62. Proboscis clearly longer than head. Ocelli absent. A long bristle at apex
of costa and two similar bristles on vein 3 (Fig. 8.75a). (Hind tibia as
Fig. 8. 60c.) Psyllon:yia
Without this combination 63
63. Mesopleuron with one or more hairs and/or bristles 64
Mesopleuron bare 65
64. Mesopleuron with very strong, very long, bristle (about as long as
costa), and sometimes a few minute hairs as well. Wing somewhat
rounded and with very long costal bristles (Fig. 6.2d) Hypogeophora
Bristles on mesopleuron much shorter and finer. Wing more than 2.5
times as long as greatest breadth and with short costal hairs only
Gymnophora (part)
Note: If palps are unusually large and two-segmented (Fig. 8.37a, b) and
abdominal tergites are reduced to lateral remnants of 1 and 2 only (Fig.
8.37c), go to couplet 248.
65. Abdominal tergite 2 at least 1.5 times as broad as long, and covering at
least three-quarters breadth of segment 66
Abdominal tergite 2 with breadth at most only 1.2 times length, and
covering at most only halfbreadth of segment Commoptera
66. Wing rudiment relatively long and narrow (greatest breadth being
about one-third oflength) and veins 1 and 3 clearly defined Thalloptera
Wing rudiment usually relatively short and broad (greatest breadth
being at least two-fifths oflength) and veins 1 and 3 poorly defined (e.g.
Figs 6.5a, 8.55a) 67
67. Note: This couplet is reached by two different routes to allow for
differences in the wing rudiments:
- Wing rudiment long and narrow, with narrow membrane behind
hairy, basally broadened, costa (Figs 6.5a, 8.81d). Proboscis clearly
longer than head (e. g. Figs 8.80b, 8.81a) Rhynchomicropteron
- Wing rudiment and proboscis otherwise 68
68. Cerci normally developed. Abdominal tergites 1-5 all broader than
long and extending full widths of segments (e.g. tergites 4--6 as Fig.

245
 IDENTIFICATION

8.55b,c). Thorax much broader than long (e.g. Fig. 8.55d). Wing
rudiment simple, with long terminal bristle (Fig. 8.55a)
Mesopathusa (part)
Without this combination 69
69. Abdominal tergites 1-5 wider than long and extending full widths of
segments. Tergite 6 a little narrower, but still wider than long. Ocelli
absent. Diameter of eye sometimes less than that of third antennal
segment. Wing rudiment a short lobe with up to six costal cilia, the end
one being as long as rudiment itself. Sometimes only this bristle is
present. Abdomen ending in sharply pointed, sclerotized, ovi-
positor Borgmeieriphora
With a different combination 70
70. Wing rudiment long and rod-shaped (being longer than hind femur).
Mid tibia with dorsal and antero-dorsal bristle in basal third and second
dorsal bristle near middle. Abdomen with tergites 1-6 present, but
progressively narrower from 2 to 6. Scutellum absent Contopteryx
- Not with this combination 71
71. Wing rudiment an elongated irregular triangle (e.g. Figs 6.5b and
8.52a). Abdominal tergite 2 clearly broader than long and extending
most, ifnot all, width of segment (e.g. Figs 6.6 and 8.52b) 72
Wing rudiment otherwise. Abdominal tergite 2 usually extending
much less than full width of segment and sometimes longer than
broad 73
Note: Since completion of this key, Brown (1993a) has described a new
genus Cootiphora, which runs to this couplet. It has no ocelli; widely
separated antennae; a dorso-ventrally flattened body, with dorsally-situated
anterior thoracic spiracles; a transversely-triangular wing rudiment tapering
rearwards, whose greatest breadth is more than wing length; and abdominal
tergites 2-6 extending full widths of segments.
72. Ocelli absent and eyes greatly reduced; abdominal tergite 2 extends full
width of segment; no other tergites on abdomen (Fig. 6.6). Mid and
hind femora each with a near-dorsal pre-apical spine (Fig. 8.51c). Mid
and hind tibiae with near-dorsal, spine-like, bristles (Fig. 8.51 b). Wing
rudiment as Fig.6.5b Maculiphora
Ocelli present. Eye with diameter at least as great as that of third
antennal segment. Abdominal tergites as Fig. 8.52b, or with tergites 3,4
present or absent. Mid and hind femora without pre-apical spines. Mid
and hind tibiae without near-dorsal, spine-like, bristles. Wing rudi-
ment as Fig. 8.52(a) Lepidophoromyia (part)
73. Wing rudiment paddle-shaped (Fig. 8.79d). Abdomen with strong
bristles arising from sclerotized discs, which may be large (e.g. Fig.
8.80a) Rhabdomyia (part)
Wing rudiment not paddle-shaped. Abdominal bristles usually much
weaker and only very rarely arising from sclerotized discs 74
74. Ocelli absent 75
Ocelli present 79
Note: Variable cases are keyed both ways.
75. Thorax clearly longer than broad and sharply narrowed behind, before

246
 KEYS TO GENERA

reaching slightly broader (confluent) scutellum. Postero-laterally rear

margm of head each side is extended rearwards as a rounded
lobe Lenkoa
Thorax usually a little broader than long or length and breadth
subequal. No rearwardly-directed lobe each side of head 76
76. Eyes reduced, so that diameter of third antennal segment is clearly
greater than diameter of eye. (Wing rudiment a long, narrow, hairy
appendage.) 77
Diameter of eye subequal to or greater than that of third antennal
segment 78
77. Abdominal tergite 2 clearly longer than wide Aemulophora
Abdominal tergite 2 clearly wider than long Idiophora
78. Wing rudiment very short (at most only four times as long as broad).
Abdominal segments 3 and 4 devoid oftergites
Lepidophoromyia (part)
Wing rudiment clearly longer. At least tergite 4 is present, even if
small 79
79. Abdominal tergite 2 clearly broader than long 80
Abdominal tergite 2 as long as broad or longer (e.g. Fig. 8.42a), but
sometimes very pale 82
80. Abdominal tergites 3-5 well developed, but progressively narrower,
and 5 with gland complex (Fig. 6.14) 81
Abdominal tergites 3-5 absent. (Tergite 2 narrower at rear than at
front. Wing rudiment at most only four times as long as wide.)
Colobomeles
81. Wing rudiment a very small finger-like structu.re (at most only three
times as long as broad) bearing minute hairs only. Pulicimyia
- Wing rudiment a rounded, membranous lobe bearing hairs and at least
one bristle at, or near, tip Puliciphora (part)
82. Wing rudiment reduced to halt ere-like structure (W in Fig.
8.42d) Ecitophora (part)
Wing rudiment a long, narrow, hairy appendage (e. g. Fig. 8. 42b) 83
83. A pair of scutellar hairs present at rear of thorax (S in Fig. 8.42d)
Ecitophora (part)
No scutellar hairs present Ecitomyia
84. Wing with only two thin veins (e.g. Fig. 8.63d); or, if membranous
region bearing thin veins has been shed, tip of abdomen is deflected
ventrally (e.g. Fig. 5.1) 85
With at least three thin veins; or, if membranous region has been shed,
tip of abdomen directed rearwards 108
85. First antennal segment not elongated 86
First antennal segment elongated (e.g. Fig. 8.46b) 108
86. No gland opening evident in middle of front margin of abdominal
segmentS 87
A distinct gland opening evident in middle of front margin of
abdominal segment 5 (as Fig. 8.45b, but smaller) 108
87. Palp enlarged apically and without bristles at tip Pseudotermitoxenia
If palp a little enlarged apically, then at least one bristle at tip 88

247
 IDENTIFICATION

88. At least 40 fine bristles on thoracic scutum. (Tip of costa extensively

overreaches tip of costa.) Termitoscrofo
- With far fewer fine bristles on thoracic scutum 89
89. Arista long, but sparsely, haired (e.g. Fig. 8.84a) to very long-haired
(e.g. Fig. 8.64a) 90
- Arista short, and more densely, haired (e.g. Fig. 8.64b) 95
90. Abdomen with tube-like appendages in anterior half 91
- Abdomen without such appendages 92
91. Some of hairs on sides of abdomen arising from discs (as in Figs 8.86a
and 8.87b) whose rims are made up of a series of outwardly projecting
lobes (appearing as dashes in plan view). Proboscis normal
Termitosphaera
- No abdominal hairs arising from such discs. Ventral face of base of
proboscis (postmentum) grossly inflated Cheiloxenia
92. Tip of costa clearly overreaching tip of vein 3 (e.g. Fig. 8.85a). Anterior
thoracic spiracles not enlarged 93
- Tip of costa little, if any, overeaching tip of vein 3 (e. g. Fig. 8.77a).
Anterior thoracic spiracles enlarged (e. g. Figs 8.64d and 8. 77c).
(Antenna as Fig. 8.64a, front metatarsus as Fig. 8.64c, proboscis as Fig.
8.77 d.) Ptochomyia
93. Head as broad as long Termitomyia
- Head longer than broad 94
94. Abdominal tergite 2 present Alamira
- Abdominal tergite 2 absent (Fig. 5.1) Termitophilomyia
95. Costa extensively overreaches tip of Vein 3, and latter normal (i. e.
almost parallel-sided) (e.g. Fig. 8.86c,d) Termitostroma
- Costa ends at or only a little beyond tip of vein 3, or if somewhat
overreaching tip then latter is more egg-shaped (e.g. Fig. 8.88d) 96
96. Proboscis relatively short (as in Fig. 8.77d). Mesopleuron always with
some fine bristles 97
- Proboscis relatively long (e.g. Fig. 8.64e). Mesopleuron frequently,
but not always, bare 99
97. Palps long and club-shaped (Fig. 8.69b). (Front leg and wing as Fig.
8. 69c, d.) Perittophora
- Palps short and more leaf-like 98
98. Vein 3 near normal (i.e. almost parallel sided), even if somewhat
obscure (Fig. 8.84e) Termitosagma
- Vein 3 more egg-shaped and distinct (as Fig. 8.83a). 'Perissa' (part)
99. Palps short and rounded and basiproboscis long (Fig. 8.64e). (Antenna
as Fig. 8.64b, wing as Fig. 8.63d.) Necperissa
- Palps longer and more tapered apically (as in Fig. 8.77d) and
basiproboscis usually shorter 100
100. Vein 3 almost parallel-sided. With four bristles on scutellum. Abdomi-
nal segment 2 not inflated to form a 'collar' Paratermitoxenia
- Without this combination 101
101. Propleuron bare 102
- Propleuron with at least one bristle in lower half and usually a second in
upper half 103

248
 KEYS TO GENERA

102. Many fine bristles on basal segments of abdomen situated on saucer-

like discs (as Figs 8.86a and 8.87b), whose rims are made up of a
series of projecting lobes (appearing as dashes in plan view)
Termitoxenia
- No saucer-like discs present at bases of any of fine bristles on
abdomen Odontoxenia
103. Apical section of hind femur inflated (Fig. 8.33c) 104
Apical section of hind femur normal 107
104. Some of fine bristles on basal segments of abdomen situated on
saucer-like discs (e. g. Fig. 8.33b), whose rims are a series of outwardly-
projecting lobes (appearing as dashes in plan view) 105
No abdominal bristles situated on such discs 106
105. Mesopleuron bare Javanoxenia
Mesopleuron with one or more (usually two) fine bristles. (Head as Fig.
8. 33d.) Clitelloxenia
106. Back of head extended rearwards so that it faces upwards (as in Fig.
8.33d) Indoxenia
Back of head normal (i. e. facing rearwards) Ceylonoxenia
107. Some of fine bristles on basal segments of abdomen are situated on
circular discs of crowded micropapillae (Fig. 8.82e) Syntermitoxenia
Some of fine bristles on basal segments of abdomen set on saucer-like
discs (as Figs 8.86a and 8.87b), whose rims are made up of a series of
outwardly-projecting lobes (appearing as dashes in plan view) 'Perissa'
(part)
108. Vein 3, apart from 1 to 3 (usually 1) hairs at base, with a row of, usually
finer, hairs along upper face 109
Vein 3 with 0-3 hairs at base only 126
109. Vein 1 also with row of hairs along upper face. (With pair of bristles, a
dorsal and a near-dorsal, in basal third of mid tibia; mesopleuron with
hairs.) Phlebothrix
- With at most a single hair on vein 1 110
110. Veins 4-6 (first three thin veins) also with rows of hairs (e.g. Fig.
8.62a) 111
- Veins 4-6 without rows of hairs 112
111. Vein 1 with single hair; scattered erect hairs on membrane between
veins; vein 3 unforked. Hind tibia without dorsal longitudinal hair
palisade. Mid tibia without differentiated bristles in basal third
Dorsochaeta
Vein 1 bare; no scattered hairs on membrane; vein 3 forked (Fig. 8.62a).
Hind tibia with dorsal longitudinal hair palisade. Mid tibia with pair of
bristles (one dorsal and one near-dorsal) in basal third. (Head as Fig.
8.62b.) Myopiomyia
112. A pair of bristles (one dorsal and one near-dorsal) in basal half of mid
tibia 113
No isolated bristles in basal two-thirds of mid tibia 123
113. Vein 3 forked 114
- Vein 3 unforked 117
114. Mesopleuron bare. Hind tibia without longitudinal hair palisades 115

249
 IDENTIFICATION

Mesopleuron with hairs. Hind tibia with two longitudinal, dorsal or

near-dorsal, hair palisades Borophaga (part)
115. Labrum more than three times as long as greatest breadth Burmophora
Labrum much shorter 116
116. With more than ten hairs on vein 3 Anevrina
With fewer than ten hairs on vein 3 127
117. Hind tibia with one or two dorsal, or near-dorsal, longitudinal hair
palisades (as Fig. 8.4d,e) 118
Hind tibia without longitudinal hair palisades, but there may be row of
spaced, differentiated, hairs (as Fig. 8.4c) 120
118. Frons as broad as, or broader than, high (long) 119
Frons clearly higher (longer) than broad Trineurocephala
119. Posterior oceIli clearly closer to eyes than to anterior oceIlus, and the
oceIlar region usuaIly clearly bounded in front by a sinuous furrow (as
Fig. 8.94d) Stichillus
Posterior oceIli clearly closer to anterior oceIlus than to eyes, and oceIlar
region not clearly demarcated in this way in front Borophaga (part)
120. Frons with one pair of bristles, the supra-antennals only. Maximum
breadth of wing sub equal to length of costa Euryophora
Frons with at least 14 bristles. Maximum breadth of wing less than
length of costa 121
121. Hind tibia with transverse combs (Fig. 8.46c). Mesopleuron with a
strong bristle as well as hairs Hypocera
Hind tibia without transverse combs. Mesopleuron without a strong
bristle and sometimes without hairs also 122
122. Mesopleuron with hairs. Abdominal tergites 5 and 6 fused, to give only
five tergites on segments 1--6 Plethysmochaeta
Mesopleuron bare. With six tergites on abdominal segments 1--6
Kuenburgia
123. Costa clearly less than half length of wing. Eight bristles on frons
between supra-antennals and oceIli. Body not obviously flattened
dorso-ventrally, so anterior thoracic spiracles are on sides Brachyselia
Costa clearly more than half wing length. With at most a single pair of
bristles on frons below (in front of) anterior ocellus. Body somewhat
flattened dorso-ventraIly, so that anterior thoracic spiracles are on
top 124
124. Costa clearly longer than greatest breadth of wing. Anterior oceIlus as
close to front margin of frons as to hind margin. Some differentiated
fine bristly hairs on frons. All tibiae with a dorsal longitudinal hair
palisade Misotermes
Costa sub equal to or shorter than greatest breadth of wing. Anterior
ocellus clearly closer to hind margin than to front margin of frons. No
differentiated bristly hairs on frons. All tibiae lack dorsal longitudinal
hair palisades 125
125. Greatest diameter (height) of eye at least twice length of third antennal
segment, and viewed from above eyes extend full length of
frons Microplatyphora
Greatest diameter of eye less than 1.5 times length of third antennal

250
 KEYS TO GENERA

segment, and viewed from above eye is less than half length of
frons Euryplatea
126. Antennae partly concealed in pits adjacent to eyes (Fig. 8.68b). Palps
short with long, unpigmented, bristles with frayed tips in addition to
normal black bristles (Fig. 8.67b). Costa with short hairs and long
bristles (Fig. 8. 68a). (Wing membrane may be shed.) Palpiclavina
- Without this combination 127
127. Mid tibia with a pair of bristles (one dorsal and one near-dorsal) in basal
half(as in Fig. 8.32a) 128
- Mid tibia without a pair of bristles in basal half, but occasionally a si'lgle
bristle is present 154
128. Hind tibia with at least one dorsal, or near-dorsal, longitudinal hair
palisade (e.g. Fig. 8.39f) 129
- Hind tibia without longitudinal hair palisades, but there may be row of
spaced, differentiated, hairs (as Fig. 8.4c) 134
129. Vein 3 forked 130
- Vein 3 unforked 132
130. Mesopleuron with hairs. (Only one longitudinal hair palisade on hind
tibia. Wing with single axillary bristle. Abdominal tergite 7 typically
well developed laterally, but middle third, or more, absent - e.g. Fig.
8.40c.) 131
- Mesopleuron bare 281
131. Scutellum with six bristles, of which middle one each side is more than
twice as long as other two. Bristle at base of vein 3 longer than distance
between point ofinsertion and the fork of vein 3 Aenictomyia
- These bristles much shorter, and usually only four on scutellum. When
there are six the middle one each side is only as long as the one
behind Dohrniphora
132. Mesopleuron with hairs Abaristophora
and Antipodiphora
Note. It is not possible, in our present state of knowledge, to distinguish
these two genera in the female sex.
- Mesopleuron bare 133
133. Hind tibia with two, dorsal or near-dorsal, longitudinal hair palisades
(Fig. 8.39f). Palp two-segmented (as Fig. 8.41 b,c) Diplonevra (part)
- Hind tibia with single, dorsal, longitudinal hair palisade. Palp single-
segmented. (Typically vein 1 as Fig. 8. 49a.) Hypocerides
134. Vein3forked 135
- Vein 3 unforked 141
135. Mesopleuron with hairs 136
- Mesopleuron bare 137
136. Median furrow present on frons. Vein 2 and Sc weakly developed and
latter usually not reaching vein 1 Coniceromyia (part)
- Median furrow absent. Vein 2 and Sc well developed and tip of latter
confluent with vein 1 Chaetopleurophora
137. Frons without a median furrow 138
- Median furrow present on frons Coniceromyia (part)
138. Bristle at tip of each fifth tarsal segment dorso-ventrally flattened (Fig.

251
 IDENTIFICATION

8.94c). Abdominal tergite and sternite of segment 7 as Figs

8. 94(a, b) Spiniphora
Bristle at tip of each fifth tarsal segment normal (i.e. round in
cross-section). Tergite and sternite of abdominal segment 7 other-
wise 139
139. The base of vein 2 missing, so that it is not attached to vein 3 to form a
complete fork Kierania (part)
and Triphleba (part)
Note: It is not possible to separate these two genera in our present state of
knowledge. Kierania is only known from New Zealand. All Triphleba species
with this feature are only known from the Holarctic Region.
Vein 2 complete 140
140. Frons with medio-Iateral bristles absent but antero-Iaterals
present Tarsocrates
Medio-Iaterals present and usually antero-Iaterals also (e.g. Fig.
8.1a) Triphleba (part)
141. The single bristle on axillary ridge of wing has very long, branched,
hairs (Fig. 8.29a) Beyermyia
Axillary bristle or bristles with only short hairs or none 142
142. Mesopleuron with hairs 143
Mesopleuron bare 144
143. Frons without median furrow Mannheimsia
Frons with distinct median furrow Coniceromyia (part)
144. Median furrow offrons present 145
Median furrow absent 147
145. One or more dorsal bristles on mid tibia beyond one in basal pair of
bristles. Usually more than one bristle on axillary ridge of wing. (Very
black species.) Phora
Only dorsal bristle on mid tibia is one in basal pair of bristles. Only one
axillary bristle on wing 146
146. Antial, antero-Iateral, pre-ocellar and medio-Iateral bristles present on
frons Coniceromyia (part)
Antero-Iateral bristles missing; other bristles present 147
147. Hind tibia with two or more dorsal or antero-dorsal bristles in upper
two-thirds 148
Hind tibia with at most only a single near-dorsal or antero-dorsal bristle
in uppertwo-thirds 151
148. Hind tibia with two dorsal and one or two antero-dorsal bristles in
upper two-thirds (e.g. Fig. 8.32a) 149
Hind tibia at most with only one dorsal and with one to three
antero-dorsal bristles in upper two-thirds 150
149. Hind tibia with one (Fig. 8.32a) or (rarely) two antero-dorsal bristles
(the second being near lower dorsal bristle). Vein Sc usually clearly not
reaching vein 1 (Fig. 8.31c), and always free when there are two
antero-dorsals on hind tibia Conicera
Hind tibia with two antero-dorsal bristles in upper two-thirds. Tip of
vein Sc reaches vein 1 and fuses with it Darwiniphora
150. Costal section 1 much longer than section 2, as veins 1 and 3 are closely

252
 KEYS TO GENERA

approximated and subparallel (Fig. 8.65c). Hind tibia as Fig.8.65b.

(Frons strongly prolongated in front - Fig. 8. 65a.) Obscuriphora
Costal section 1 sub equal to or clearly shorter than section 2, as the tips
of veins 1 and 3 are widely separated. The two to three antero-dorsal
bristles on hind tibia are more widely spaced Gymnoptera
151. A single near-dorsal bristle in upper two-thirds of hind tibia 152
- No differentiated bristle in upper two-thirds of hind tibia 153
152. With only six bristles on frons between the two supra-antennals and
ocelli, the antero-Iaterals being absent Bothroprosopa
With eight bristles on frons between the two supra-antennals and ocelli
(e.g. Fig. 8.1a) Kierania (part)
and Triphleba (part)
Note: See note in couplet 139.
153. Arista apical in position Billotia
Arista in dorsal, pre-apical position Triphleba (part)
154. Hind tibia with at least one dorsal, or near-dorsal, longitudinal hair
palisade (as Fig. 8.4d,e) 155
Hind tibia without longitudinal hair palisades, but there may be a row
of spaced, differentiated, hairs (as Fig. 8.4c) 247
155. Vein 3 unforked 156
Vein 3 forked 182
Note: Some genera are keyed both ways, especially when the base of vein
2 is obscure.
156. A clearly demarcated, darkened, oval patch on wing between bases of
veins 4 and 5 (the first two thin veins). Veins 1 and 3 subparallel or even
convergent, in basal third or more. Phymatopterella (part)
- Without this combination 157
157. Mid and hind femur each with pair of strong spines on anterior face in
pre-apical position (similar to Fig. 8.91a) Colyeria
- No such spines on mid and hind femora 158
158. Front tarsal segments 1-4 or 1-3 asymetrically widened (Fig. 8.72a) and
tarsal segment 5 of all legs tapered (Fig. 8.72c,d). Mesopleuron with a
few hairs Plastophorides
- Front tarsi without asymetrically widened segments. Fifth tarsal
segments rarely tapered; but when they are, mesopleuron is bare 159
159. Eyes greatly enlarged and almost meeting below, and lower facets
(ommatidia) markedly larger than those above (Fig. 8.28a). Legs
slender (Fig. 8.18b). Ovipositor of elaborate design (e. g. Fig.
2.5) Auxanommatidia (part)
Without these features combined 160
160. Body somewhat dorso-ventrally flattened, so that anterior thoracic
spiracles are dorsal in position 161
- Body not modified in this way; anterior thoracic spiracles on sides
162
161. Apical section of third antennal segment elongated, tapered and bearing
apical arista. Scutellum with four strong bristles Platydipteron
Third antennal segment more oval, not tapered apically, and aristajust
pre-apical. Scutellum with ten hairs on margin. (Hair palisade of hind

253
 IDENTIFICATION

tibia complete in basal part, but more intermittent distally as tranverse

combs more developed.) Melittophora
162. Costa less than third of wing length and distal half or more of section 3
turned rearwards, through a right angle, to meet tip of vein 3 well
behind wing margin Stenoneurellys
Costa not modified in this way 163
163. Mesopleuron with hairs 164
Mesopleuron bare 165
164. Veins 1 and 3 subparallel (section 1 being about three times length of
section 2). Abdominal tergite 6 at least eight times as broad as long
(being only about as long as tergite 1) and with small median
emargination in front margin Syneura (part)
Veins 1 and 3 divergent (section 1 being clearly less than three times
length of section 2). Abdominal tergite 6 longer and narrower
Megaselia (part)
165. Abdominal tergite 6 with an embayment of front margin which
embraces a roughly pear-shaped, hairless, sclerite (Fig. 8.93a). Intern-
ally Dufour's crop mechanism not only very short and wide, and
usually darkly sclerotized, but also with pair oflong tubules extending
from middle into lumen of crop (Figs 6. 8a and 6. 9b) Woodiphora (part)
Abdominal tergite 6 and Dufour's crop mechanism (when present) not
modified in these ways 166
166. Costa less than two-fifths of wing length 167
Costa more than two-fifths of wing length 174
Note: Variable genera are keyed both ways.
167. Only two bristles on front (lower) half of frons. (Third antennal
segment elongate with pre-apical, dorsal arista; veins 1 and 3 sub-
parallel.) Styletta
At least four bristles on front half offrons 168
168. Third antennal segment more than twice as long as broad, with dorsal
arista clearly before tip Tubiceroides (part)
Third antennal segment usually clearly less than twice as long as broad;
but if nearly twice as long, then arista is apical or, occasionally,
absent 169
169. Supra-antennal bristles absent. Ovipositor asymmetrical and complex
(usually with one or two strong black spines; venter of abdominal
segment 6 often with well-developed sternite and frequently with
conspicuous bristles and/or spines) Cremersia (part)
At least one pair of supra-antennal bristles present. Ovipositor
symmetrical 170
170. Abdominal tergites 5 and 6 both very short, and 6 completely encircling
segment Trucidophora
Not thus 171
171. Abdominal sternite 6 present and bearing bristles; modified ovipositor
well developed and sometimes bizarre (e. g. Fig. 2.6) Pseudacteon (part)
No sternite on venter of abdominal segment 6 172
172. Terminal segments of abdomen developed as a somewhat dorso-
ventrally flattened ovipositor 173

254
 KEYS TO GENERA

Terminal abdominal segments not so modified Megaselia (part)

173. Costa at most only one-third of wing length Pradea
Costa more than one-third of wing length Pseudacteon (part)
174. Epistoma (clypeus) inflated between bases of antennae and extending to
front of frons Apodicrania
Epistoma not unusually inflated 175
175. Hind tibia, in addition to dorsal hair palisade, with irregular antero-
dorsal palisades in basal third. No supra-antennal bristles. Vein Sc
obscure or absent. Median furrow of frons obscure or absent. Third
antennal segment elongated and tapered, with an apical or subapical
arista. Abdominal tergites wide, extending onto upper parts of
flanks Melaloncha (part)
Without this combination 176
176. Third antennal segment about twice as long as greatest breadth,
strongly tapered, with apical arista. With two to four supra-antennal
bristle Ceratoconus
Without both characters in combination 177
177. Third antennal segment oval to conical 178
Third antennal segment globose, but sometimes slightly pointed (in
latter case tip of vein 4 deflected rearwards) 179
178. A brown sternite bearing hairs present on abdominal segment 6 (e.g.
Fig. 2.6) Pseudacteon (part)
No sternite on abdominal segment 6 Apocephalus (part)
and Pseudacteon (part)
Note: A few Apocephalus species have an unforked vein 3, as is typical of
Pseudacteon species lacking a sternite on segment 6. T1!e former cannot be
distinguished with confidence, in our present state of knowledge, unless
associated with their males.
179. Costal index exceeds 0.7. Veins 4 and 5 widely separated, vein 4 being
very pale and originating near apex of vein 3; base of vein 5 originating
well before apex of vein 1. Abdominal tergites 3 and 4 fused to form a
single elongated tergite Parasyneura
Without this combination 180
180. Third antenna I segment slightly pointed; costa more than halflength of
wing; with elongated ovipositor with long narrow cerci; tip of vein 4
deflected rearwards and running parallel to wing margin for short
distance Ectochaeta
Not with this combination 181
181. Without row of differentiated postero-dorsal hairs behind hair palisade
of hind tibia Beckerina (part)
With differentiated row of postero-dorsal hairs on hind tibia
Megaselia (part)
182. Mesopleuron bare 183
Mesopleuron with hairs, and sometimes bristles as well 224
183. Front tibia with dorsal longitudinal hair palisade, as on mid and hind
tibiae. Fore tarsus with only four segments Neodohrniphora
Without these two features in combination 184
184. Front tarsus with only two, long, segments, the second usually being

255
 IDENTIFICA nON

tapered 185
- Front tarsus with five segments 186
185. Venter of abdominal segment 6 with distinct, semicircular or horse-
shoe-shaped, sternite bearing bristles on hind margin. (Costal index at
least O. 32, usually 0.34 or more.) Dacnophora (part)
- Venter of abdominal segment 6 without distinct sternite. (Costal index
at most 0.36, usually 0.33 or less. Lower ommatidia of each eye a little
larger than upper ones.) Myrmosicarius
186. Eyes greatly enlarged and almost meeting below, and lower facets
(ommatidia) larger than those above (e.g. Fig. 8.28a). Legs slender (Fig.
8.18b). Ovipositor elaborate (e. g. Fig. 2.5) Auxanommatidia (part)
Not with these features combined 187
187. Fork of vein 3 very large, so that costal section 3 is about four times as
long as section 2. Vein Sc strongly developed and its tip fusing with
vein 1. Axillary ridge with at least five bristles. Abdominal tergite 6
almost divided in two by anterior V-shaped notch. On each flank at
boundary of abdominal segments 5 and 6 there is an eversible
finger-like process Dicranodeina
Without this combination 188
188. First half to two-thirds of costal section 1 clearly wider than rest of
costa, and with bristle situated dorsally at end of wide part at least as
long as rest of costa. Fork of vein 3 large, with vein 2 as strongly
developed as veins 1 and 3. Costal cilia of section 3 longer than vein 2.
(The wing membrane may be shed.) Pheidolomyia
Not thus 189
189. With 16-20 supra-antennal bristles; otherwise frontal bristles normal.
Frons broader than long. (Costa only about one-third of wing length.
Ovipositor short, broad, with narrowly rounded tip.)
Apocephalus (part)
Usually with fewer than 14 supra-antennals (usually 2-4), but if as
many as 16 the frons is as long as broad 190
190. Hind tibia with single, dorsal, hair palisade 191
- Hind tibia with several, irregular, hair palisades (Fig. 8.73a) 194
191. Six or more supra-antennal bristles 192
- None to four supra-antennal bristles 193
192. With differentiated row of postero-dorsal hairs behind hair palisade on
hind tibia Johowia (part)
- Without differentiated postero-dorsals on hind tibia Beckerina (part)
193. With dark oval patch on wing between tip of vein 3 and base of vein
4 Phymatopterella (part)
- Any dark marks on wing either are traversed by veins or are between
bases of veins 4 and 5 194
194. With more than one longitudinal hair palisade on hind tibia, even if
those on anterior face only present on basal third 195
With only one, dorsal, hair palisade on hind tibia 197
195. Several irregular hair palisades on anterior face of mid and hind tibiae,
those on hind leg running full length of tibia (Fig. 8.73a) Plectanocnema
Any irregular hair palisades on anterior faces of mid and hind tibiae are

256
 KEYS TO GENERA

restricted to basal halves or less; but regular, even if intermittent,

palisades may be present beyond basal third (as Fig. 8. 92c, d) 196
196. Irregular hair palisades of anterior face of hind tibia restricted to basal
third. Without supra-antennal bristles. Vein Sc obscure or absent.
Abdominal tergites wide, extending onto upper parts of flanks.
Terminal segments obviously developed as an ovipositor
Melaloncha (part)
- Without this combination. Abdominal tergite 6 as Fig. 8.93a, and no
obviously developed ovipositor behind. (InternaIly, Dufour's crop
mechanism not only very short and heavily sclerotized, but also with
pair of long tubules extending from middle into lumen of crop - Figs
6.8a and 6. 9b) Woodiphora (part)
Note: One Neotropical species assigned to Beckerina (B. nudipleura) wiIl
also key out here. However, it seems likely that it is reaIly a Woodiphora
species. Its description differs from that of a typical Woodiphora by its more
acute angle between veins 2 and 3. Its transference to the latter genus would
require reporting of a notopleural cleft in the male and the above type of
Dufour's crop mechanism in the female.
197. The distal section of proboscis (distiproboscis) long and narrow (being
longer than length of head) Rhyncophoromyia (part)
If proboscis longer than head, then this not primarily due to elongation
of distproboscis (labrum plus labeIla) 198
198. Epistoma (clypeus) longer than broad. Membranous region in front
bearing palps also elongated, with result that latter originate weIl in
front of (below) head. Abdominal tergite 2, and sometimes 3 also,
elongated; tergites 3-5 or 3-6 shortened and of complex structure
associated with gland opening. Segments 3-6 usuaIly with one or more
finger-like processes on flanks Trophithauma
Without this combination 199
199. Third antennal segment more than twice as long as greatest breadth and
with pre-apical, dorsal, arista. Frontal bristles all close to margins (six at
front; two each side, one near middle and one near top of each eye; and
four on vertex). Base of vein 2 df'tached from vein 3 Tubiceroides (part)
Notso 200
200. Wing with triangular dark patch between bases of veins 4 and 5 201
- With no such dark patch 202
201. Base of triangular dark patch on wing against hind margin of vein 3.
There is a smaIl oval sweIling near base of vein 3 at base of vein
6 Physoptera (part)
Base of triangular dark patch clearly separated from vein 3. Without
oval swelling near bases of veins 3 and 6 Phymatopterella (part)
202. Wing with small oval swelling near base of vein 3 at base of vein 6. Basal
half of hind tibia with postero-ventral row of differentiated hairs,
which are either shorter or longer than adjacent hairs. Costa more than
half wing length Physoptera (part)
- Not with this combination 203
203. Hind tibia with differentiated row of antero-dorsal hairs as well as
postero-dorsal row (the two rows separated by dorsal hair

257
 IDENTIFICA TION

palisade) 204
- Hind tibia at most with postero-dorsal row of differentiated hairs
ocly 2ffi
204. Palp bristles and supra-antennal bristles only about as strong as upper
occipital bristles. Abdominal tergite 3 frequently with a pair of oval to
round papillae near front margin (Fig. 8.70a) Phalacrotophora (part)
- Either the palp or supra-antennal bristles or both clearly longer than
upper occipitals. Without such papillae on abdominal tergite 3
Megaselia (part)
205. Last tarsal segments on both front and middle legs, at least, longer than
fourth segments and usually tapered (Figs 8. 4b and 8. 56b, c) 206
- Last tarsal segments shorter than fourth segments on all legs or else
longer on front pair only 208
206. Palps with bristles, even if they are short. Costa at most only two-fifths
of wing length 207
- Palps without bristles. Costa just over two-fifths of wing length.
(Supra-antennal bristles absent. Abdominal tergites, ovipositor, and
fifth tarsal segments as Fig. 8. 79a-c.) RazorJemora
207. Third antennal segment pear-shaped, with apical arista and conus
(apical bulb) of second segment inserted into side, not base, of third
segment Epacteon
- Third antennal segment bean-shaped with dorsal, pre-apical, arista and
conus of second segment inserted into base of segment in normal way
(e.g. Figs 8.56d and S.57a). (Ovipositor well developed - e.g. Fig.
S. 58.) Microselia
20S. Abdominal segment 6 with distinct sternite bearing bristles 209
- Abdominal segment 6 without sternite 212
209. Only four bristles at front margin offrons. Fifth tarsal segment offront
leg frequently longer than fourth and tapered (as Fig. S.56b) 210
- At least six bristles at front margin of frons. Fifth tarsal segment not
elongated or tapered Synclinusa
210. Ovipositor symmetrical 211
- Ovipositor asymmetrical and of complex structure (usually with one or
two strong black spines) Cremersia (part)
211. Costa clearly less than half wing length Dacnophora (part)
- Costa at least half wing length Macrocerides (part)
212. Hind tibia without postero-dorsal row of differentiated hairs, the hairs
in this position being no different from adjacent ones of posterior
face 213
- Hind tibia with row of distinctly differentiated postero-dorsal hairs,
even if they are fine 214
213. Abdominal tergite 6 as Fig. S.93(a). Internally Dufour's crop mechan-
ism not only very short and heavily sclerotized, but also with pair of
long tubules extending from middle into lumen of crop (Figs 6. Sa and
6.9b) Woodiphora (part)
- Abdominal tergite 6 otherwise. Without Dufour's crop mechanism
Beckerina (part)
214. With only four bristles at front margin offrons. Ovipositor asymmetri-

25S
 KEYS TO GENERA

cal and complex (usually with one or two strong black spines)
Cremersia (part)
Without this combination 215
215. Third antennal segment conical, with apical arista. With only one
bristle on axillary ridge of wing Procliniella
Without these two features in combination 216
216. Costa less than halflength of wing 217
- Costa at least halflength of wing 220
Note: Borderline cases are keyed both ways.
217. Terminal abdominal segments (beyond tergite 6) clearly modified into
dorso-ventrally flattened basal part embracing pointed ovipositor
beyond 218
Terminal abdominal segments not obviously modified as specialized
ovipositor 219
218. Supra-antennal bristles as strong as other bristles at front offrons
Neophora
Supra-antennals clearly shorter and weaker than other bristles at front
offrons Apocephalus (part)
219. Only two supra-antennal bristles. Costa only about one-third of wing
length. Abdominal tergite 6 reduced and much wider than long
Syneura (part)
Without these three features in combination Megaselia (part)
220. The two supra-antennal bristles as strong as antials. Third antennal
segment conical with apical, or slightly subapical, arista. Base of
ovipositor broader than long, embracing the pointed distal portion
(ovipositor proper). With long bristles at rear of segment 6, on sides and
below Anaclinusa
Without this combination 221
221. Terminal abdominal segmencs (beyond tergite 6) clearly modified into
dorso-ventrallyflattened basal part embracing pointed ovipositor (e. g.
Fig. 2.5) 222
Terminal abdominal segments not obviously modified to form special-
ized ovipositor 223
222. Frons clearly broader than long; with only two, strong, supra-antennal
bristles in nearly straight transverse row with antials and antero-Iaterals
at front offrons. Antials close to antero-Iaterals Lecanocerus
Frons usually as long as broad or longer; with antials near mid line and
two to four reduced supra-antennals below these or absent
Apocephalus (part)
223. With only two supra-antennal bristles. Postero-dorsal hairs of hind
tibia only weakly differentiated from adjacent hairs. Internally without
Dufour's crop mechanism Beckerina (part)
Without first two features in combination. Internally Dufour's crop
mechanism is present (e.g. Fig. 6.8b) Megaselia (part)
Note: If there is a small swelling near base of vein 3 at base of vein 6 and
costa exceeds half wing length, it may be a Physoptera species without ventral
hairs on hind tibia (see couplet 202).
224. With tube-like elongation of basiproboscis (largely embracing

259
 IDENTIFICATION

distiproboscis) so that reduced palps are borne at tip of proboscis,

which is clearly longer than head. (Third antennal segment globose.
Costa at least half wing length. Abdominal tergite 6 longer than any
other tergites and with small notch in front margin.) Calamiscus
- Proboscis usually much shorter and palps always at base 225
225. Wing margin incised at tip of vein 5, and the posterior lobe bordering
incision with dense patch of minute hairs. Veins 4 to 6 not thin (as in
most phorids), but as thick as vein 1 Menozziola (part)
- Without these features in combination 226
226. Eight bristles on scutellum 227
- Two to six bristles on scutellum 228
227. In addition to rest of normal bristles on frons, with only four to six
differentiated supra-antennals Egregiophora
- In addition to rest of normal bristles on frons, with four median
supra-antennals and several additional differentiated bristles either side
of these Johowia (part)
228. Distiproboscis (labrum plus narrow labella) elongated, exceeding
length of head. Labrum almost parallel-sided 229
- Distiproboscis shorter than head, or if longer then labrum is much
broader basally and tapers towards its tip 230
229. Labrum broader (e.g. Fig. 8.2c) and cerci well developed (e.g. Fig.
8.6b) Megaselia (part)
- Labrum narrower. Cerci vestigial Rhyncophoromyia (part)
230. At least six differentiated supra-antennal bristles 231
- At most only four supra-antennals 232
231. With six of eight supra-antennals on sides of a narrow forward
projection of middle of front margin of frons (between bases of
antennae). Only two bristles on axillary ridge of wing. Abdominal
tergites 1-5 very wide but short, but 3 being narrower than others.
Tergite 6 reduced to small semicircle associated with gland
opening Lobochaeta
- Additional supra-antennals either side of normal ones. At least four
axillary bristles. Abdominal tergites otherwise Johowia (part)
232. Hind tibia with row of differentiated antero-dorsal hairs as well as
postero-dorsals (the two rows being separated by hair palisade) 233
- Hind tibia without differentiated row of antero-dorsals 235
233. Basal half of hind tibia with differentiated row of very short antero-
ventral hairs. Wing with small oval swelling near base of vein 3 at base
of vein 6 Physoptera (part)
- Without this combination 234
234. Palp bristles and supra-antennals only about as strong as upper-
occipital bristles. Abdominal tergite 3 frequently with pair of oval to
round papillae near front margin (Fig. 8.70a) Phalacrotophora (part)
- Either palp or supra-antennal bristles or both clearly longer than upper
occiptals. Without such papillae on abdominal tergite 3
Megaselia (part)
235. Second quarter of hind tibia with differentiated near-ventral row of
longer hairs. Wing with small oval swelling near base of vein 3 at base of

260
 KEYS TO GENERA

vein 6. Abdominal tergites 3 and 4 greatly reduced, so that dorsal faces

of these two segments largely membranous Physoptera (part)
- Not with this combination 236
236. More than 12 bristles on axillary ridge of wing. Arista with very short
and widely spaced hairs (Fig. 8.74c), appearing to be bare at low
magnifications. Three bristles on humerus, the middle one being above
anterior thoracic spiracle. All tarsal claws with pre-apical tooth (Fig.
8. 74b) Pseudohypocera
Without these features in combination 237
237. Frons clearly longer than broad; with five bristles each side, one above
the other and close to eye, the only other bristles being pre-ocellars and
ocellars. Scutellum with six bristles, the middle one each side being
longest and strongest. Costa only about one-third of wing length.
Terminal abdominal segments dorso-ventrally flattened and embracing
a sharply-pointed ovipositor Stenodicrania
- Notso 238
238. Fifth tarsal segments on all legs longer than fourth segments and
strongly tapered (as Fig. 8.56b). With sharply-pointed ovipositor and
vestigial cerci. Abdominal tergite 6 with conspicuous anterior median
notch. Mesopleuron with fine hairs but no bristles. Costa less than
two-fifths of wing length Lepta
Without this combination 239
239. Frons clearly broader than long and without median furrow; with four
median bristles near front and three each side (one above the other) near
eye. Third antennal segment pear-shaped with apical arista. Terminal
segments of abdomen long and narrow with sharply-pointed ovi-
positor. Mesopleuron with hairs and one long bristle Acanthophorides
Without these features in combination 240
240. Mesopleuron with strong bristle as well as hairs 241
Mesopleuron with hairs only 243
241. Abdominal tergites 5 and 6 clearly shorter and narrower than tergite 4.
A pair of finger-like processes on sides of segment 5 (e.g. Fig.
2.6a) Menozziola (part)
Not with both these features 242
242. Ovipositor not well developed but cerci, bearing hairs, well developed
(e.g. Fig. 8.6b) Megaselia (part)
- Ovipositor well developed but cerci vestigial and without hairs (e.g.
Fig.8.38b) Diocophora,
Macrocerides (part)
and Megaselia (part)
Note: The females of Macrocerides species, whose males have a bristle on
the mesopleuron, have not yet been described. A few Megaselia species with
modified ovipositors cannot be distinguished from Diocophora, in our present
state of knowledge, apart from one Diocophora with elongated, tapered, fifth
tarsal segments, one with bifid tarsal claws and one whose vein 4 ends behind
the wing tip.
243. Ovipositor blade-like (laterally compressed) with well developed cerci
below (Fig. 8.51a) Megaselia (part, subgenus Kerophora)

261
 IDENTIFICATION

Ovipositor not thus 244

244. Terminal segments of abdomen (behind semicircular tegite 6) dorso-
ventrally flattened, broadening apically and with concave hind margin.
Ovipositor much narrower and normally withdrawn Ceratophoromyia
- Terminal abdominal segments otherwise 245
245. Frons clearly wider than long. Costa just over half wing length. Only
two bristles on axillary ridge of wing. Third antennal segment oval,
with pre-apical arista. With well developed ovipositor embraced by
modified terminal segments Brachycephaloptera
Without this combination 246
246. With well-developed ovipositor and cerci vestigial or absent. Third
antennal segment short-conical with pre-apical arista. Wing with vein
Sc well developed, more than six bristles on axillary ridge and costa
more than halflength Macrocerides (part)
Not with this combination Megaselia (part)
247. Mesopleuron with hairs and wing vein 3 forked 248
Mesopleuron bare and vein 3 forked or unforked 250
248. Palps very large, two-segmented, and with scattered hairs but no
strong bristles (Fig. 8.37a,b). Fork of vein 3 large; wing membrane may
be shed (Fig. 6.1a-c). Remnants of tergites present on abdominal
segments 1 and 2 only (Fig. 8.37c) Dicranopteron
Not thus 249
249. Without supra-antennals, antials, antero-Iaterals and pre-ocellar
bristles, and usually medio-Iaterals as well. A notopleural cleft present
(Fig.8.47a,b) Gymnophora (part)
- With four supra-antennal bristles and other normal bristles on frons.
Without notopleural cleft. (Fork of vein 3 large. ) Tropophleba
250. Wing vein 3 forked 251
Vein 3 unforked 260
251. An isolated bristle in basal third of mid tibia Triphleba (part)
Without such a bristle 252
252. Wing with vein 6 deflected forwards opposite basal bend of vein 5, so
that space between two veins is constricted (as Fig. 8.56a). No bristles
on axillary ridge Metopina (part)
Vein 6 not deflected in this way. At least one bristle on axillary
ri~ ill
253. Third antennal segment clearly longer than greatest breadth, strongly
tapered from broad base (the tapered part being at least one-third of
length), with apical arista Corynusa
Third antennal segment either globose, with pre-apical arista, or if
somewhat pear-shaped with apical arista it is scarcely, if any, longer
than greatest breadth 254
254. End of costa clearly overreaches tip of vein 3 a short distance. With only
two axillary bristles, which are shorter than costal cilia on costal section
3 Allochaeta
Without this combination 255
255. Apart from bristles on vertex, with only eight or fewer bristles on
frons. Median furrow offrons absent 256

262
 KEYS TO GENERA

- Apart from bristles on vertex, with 10 or 12 bristles on frons. Median

furrow present, but sometimes obscure 258
256. Only four or six bristles on frons. Breadth of frons clearly more than
twice length from anterior ocellus to front margin. Macroselia
- Eight bristles on frons. Frons breadth not twice midline length 257
257. With only one or two bristles on axillary ridge of wing Rhopica (part)
- With at least three bristles on axillary ridge Aphiura
258. Proboscis long and slender (the distal part being longer than frons).
Fork of wing vein 3 large (costal section 3 being as long as, or longer
than, section 2); axillary ridge with only two bristles Pericyclocerina
- Without this combination 259
259. Abdominal tergites 6 and 7 as Fig. 8.93(a). Internally Dufour's crop
mechanism not only very short and heavily sclerotized, but also with
pair of long tubules extending into lumen of crop (Figs 6.8a and
6. 9b) Woodiphora (part)
Abdominal tergites 6 and 7 otherwise. Without Dufour's crop mechan-
ism Beckerina (part)
260. Wing with very long bristles on costa (Fig. 8.17b)
Adelopteromyia (part)
- Costa with shorter, finer, bristles or hairs. When some long bristles
present in outer half, then section 1 (at least) with hairs 261
261. Fifth tarsal segment on all legs longer than fourth and a little to strongly
tapered (e.g. Fig. 8.50b,c) 262
- Fifth tarsal segments not longer than fourth segments 263
262. All tibiae with near-dorsal pre-apical bristle in last quarter (and
sometimes one or two additional near-dorsals above this one). Outer
half of costa with 4-5 long bristles (some being longer than vein 3).
(Wing membrane may be shed.) Third antennal segment globose with
apical arista. Abdominal tergites 1-6 absent. Ovipositor not obviously
modified Termitophorides
- Tibiae without pre-apical near-dorsal bristles. Wing (Fig. 8.50d) with
short costal cilia only. Third antenna I segment elongated and without
arista (Fig. 8.50a). Abdominal tergites 1-6 present. Ovipositor as Fig.
2.7 Iridophora
263. First antennal segment elongated and third segment conical, with
pre-apical arista; proboscis narrow and elongated (Fig. 8.46b). Without
tergites on abdominal segments 1-6 (Fig. 8.46a). (Wing membrane
may be shed- Fig. 6.1d.) Eutermiphora (part)
Without this combination 264
264. Third antennal segment globose, with apical arista. More than 30
bristles on thoracic scutum (mesonotum). No tergites on abdominal
segments 1-6. Costa clearly less than one-third of wing length and with
two longer bristles in outer half, preceded by shorter hairs
Sphinctomyia
- Not with these features combined 265
265. Third antennal segment with apical half developed as long tubular
extension bearing widely-spaced long hairs, contrasting with very
short hairs of apical arista. (Only four bristles on frons in front of

263
 IDENTIFICATION

anterior ocellus, two being supra-antennals; costa at least half wing

length.) Dahliphora (part)
Third antennal segment globose, without apical extension bearing long
hairs 266
266. At least two bristles on axillary ridge of wing 267
Without axillary bristles 269
267. At most only ten bristles on frons in front of anterior ocellus, including
two supra-antennals 268
Twelve bristles on frons in front of anterior ocellus, including four
supra-antennals. (Dufour's crop mechanism wider than long, and with
pair of long tubules extending into lumen of crop - e.g. Fig.
6. 9b.) Woodiphora (part)
268. Only eight bristles on frons in front of anterior ocellus. Fewer than four
bristles on axillary ridge of wing Rhopica (part)
Ten bristles on frons in front of anterior ocellus. More than five axillary
bristles Haplophleba
269. Thoracic scutum (mesonotum) more than 2.5 times as long as broad
and tapering towards distinctly separated scutellum. Frons about twice
as long as wide, with four supra-antennals, a pair of reduced
pre-ocellars and bristles on vertex only. Abdominal tergites 1-5
present, with front half of 5 developed as semicircular flap
Stenophorina
Without these features combined 270
270. At least abdominal tergite 2 present 271
Abdominal tergites 1-6 absent, apart from rudiment of 5 associated
with gland opening. (Without bristles on frons between anterior ocellus
and two supra-antennals. ThoraClc scutum clearly wIder at front than at
rear. Wing membrane may be shed.) Termitophorina
271. With only tergite 2 and rudiment of 5, associated with gland opening,
present on abdominal segments 1-6 272
With at least four tergites present on abdominal segments 1-6 273
272. Abdomen with numerous long bristles (longer than tergite 2). Wing
membrane may be shed; when present costa at most only one-third of
wing length and only two thin veins present. Costal cilia on section 2
longer than vein 1 Echidnophora
- Abdomen with short hairs only, which are only a little longer at hind
margins of segments. Wing membrane not shed; costa more than half
wing length and three thin veins present. All costal cilia shorter than
vein 1 Bolsiusia
273. Costa less than one-third of wing length. Only one bristle on palp.
Only two supra-antennal bristles and two pre-ocellars on frons, apart
from those on vertex. Terminal segments of abdomen developed as
down-curved beak-shaped ovipositor Brachyphlebina
- Costa about half, or more, of wing length. Without other characters in
combination 274
274. Wing as Fig. 5.2(b). Eyes absent or represented by one or two reduced
ommatidia only. Ocelli absent Metopina (part)
Wing with thin veins evident. Eyes and ocelli present 275

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 GUIDE TO LITERATURE FOR SPECIES IDENTIFICA nON

275. Only four bristles, including supra-antennals, on frons In front of

anterior ocellus 276
- At least six bristles on frons in front of anterior ocellus 278
276. Vein 6 bent towards basal curve of 5, so that space between veins is
constricted in middle (e.g. Fig. 8.56a) 277
Veins 5 and 6 not thus (e.g. Figs 8.34a and 8.35a). (With four
supra-antennals as only bristles on frons, apart from those on vertex.
Abdominal tergites as Figs 8.34b and 8.35b.) Crasilla
277. Two supra-antennals and two pre-ocellar bristles on frons, apart from
six bristles on vertex Dahliphora (part)
Frons with four supra-antennals only, apart from those on vertex
Metopina (part)
278. Frons at least three times as broad as long. ('pre-ocellar' bristles level
with posterior ocelli. 'Thin' veins 4-6 may be almost as thick as outer
third of costa.) Holopterina
Frons less than three times as broad as long 279
279. Vein 6 bent towards basal curve of vein 5, so that space between veins is
constricted in middle (e. g. Fig. 8.56a) Metopina (part)
- Vein 6 not bent in this way (indeed vein 6 may be abbreviated or absent,
in association with narrowing of basal half of wing) 280
280. Wing broadest at level of tip of costa or beyond (Fig. 8.43a). Abdominal
tergite 2 at most only twice as wide as long Ecitoptera
Wing broadest before level of tip of vein 1. Abdominal tergite 2 more
than four times as wide as long Cataclinusa
281. Head, thorax and abdomen somewhat flattened dorso-ventrally, with
anterior thoracic spiracles dorsal in position. Eyes somewhat reduced.
Hind tibia with antero-ventral row of spines before apical spur
Synaptophora
Without this combination. (With two or more longitudinal hair
palisades on hind tibia - Fig. 8.39£ Wing with two or more axillary
bristles.) Diplonevra (part)

8.3 GUIDE TO LITERATURE FOR SPECIES

IDENTIFICA TION

The following review gives the number of known species in each genus and
cites the keys or descriptions that will allow identification of the species. It has
been estimated that perhaps as few as one-tenth of the world's scuttle fly
species have been described and named. Furthermore the limits of distri-
bution for the described species are poorly known. There is therefore a high
probability of turning up species new to science or new to a region.

Abaristophora Schmitz
Two Palaearctic and one Nearctic species are known (Schmitz, 1951c;
Michailovskaya, 1988; Brown, 1988).

Acanthophorides Borgmeier
The seven N eotropical species are keyed by Borgmeier (1967h).

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 IDENTIFICA TION

Acontistoptera Brues
The one Nearctic and three Neotropical species are keyed by Borgmeier
(1963d).

Adelopteromyia Schmitz
The three or four Neotropical species (one species may be merely the
short-winged morph of another) are keyed by Disney and Kistner (1990b).

Adenophora Borgmeier
The single Neotropical species, known only in the female sex, is covered by
Borgmeier (1960b).

Aemulophora Borgmeier
The single Neotropical species, known only in the female sex, is covered by
Borgmeier (1938b).

Aenictacantha Disney
Two species from Malaysia and Sulawesi, known only as females, are
covered by Disney (1991m).

Aenictomyia Brues
The single Oriental species, known only in the female sex, is covered by
Borgmeier (1967a).

Aenigmatias Meinert
A dozen species are known from the Nearctic and Palaearctic Regions. The
former are keyed by Borgmeier (1963d) and the latter by Schmitz (1955b),
supplemented by Disney (1983a) for males and (1993c) for females.

Aenigmatistes Shelford
The 20 Afrotropical species are keyed by Disney (1991k).

Aenigmatopoeus Schmitz
The females of the six Afrotropical species are keyed by Borgmeier (1963e).
Colyer (1958) describes the only known male.

Alamira Schmitz
The two Afrotropical species, known only in the female sex, are keyed by
Disney (1990j). A third species, A. peckorum, is a species of Termitostroma (see
below).

Allochaeta Borgmeier
The four Brazilian species are keyed by Borgmeier and Prado (1975).

Anaclinusa Borgmeier
The single Neotropical species is covered by Borgmeier (1969b, 1971a).

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 GUIDE TO LITERATURE FOR SPECIES IDENTIFICATION

Anevrina Lioy
The five Palaearctic species are covered by Schmitz (1941 c) and Disney
(1983a). The nine Nearctic species are keyed by Borgmeier (1963d). Beyer
(1958a) describes three species from Burma.

Anticofimbria Schmitz
The male of the single known species from Japan is described by Schmitz
(1951e). The female is unknown.

Antipodiphora Schmitz
Six species from New Zealand are keyed by Schmitz (1939a). Undescribed
species occur in the Oriental Region.

Aphiura Schmitz
Three Australasian species are discussed by Borgmeier (1967a), along with an
unnamed female that probably belongs to some other genus.

Apocephalus Coquillett
There are certainly over 120 species in this genus, which is known from the
Australasian, Oriental, Nearctic and Neotropical Regions. Brown is cur-
rently carrying out revisionary studies. Until these are complete, specimens
can be keyed with Borgmeier (1971 a), supplemented by Prado (1976) and
Disney (1980i, 1981d, 1982k). A Nearctic species assigned to a new genus,
Zyziphora (Peterson and Robinson, 1976), has been transferred to Apo-
cephalus by Brown (1992b). It is also figured by Peterson (1987). Brown
(1993b) provides a revised key to Nearctic and Neotropical species of the
subgenus Mesophora.

Apodicrania Borgmeier
The three Neotropical species are keyed by Borgmeier (1963e).

Apterella Borgmeier
A single species from Costa Rica is known only in the female sex (Borgmeier,
1935a).

Apterophora Brues
The five Neotropical species are discussed by Prado (1976), allowing
identification of the males. In our present state of knowledge, females can
only be named when associated with their males.

Aptinandria Schmitz
Schmitz (1921£) describes the female of the single Afrotropical species. The
male has not been described, but it was included in a key by Beyer (1965b).

Aristocerina Borgmeier and Prado

A single Neotropical species, known only in the male sex, is described by
Borgmeier and Prado (1975).

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 IDENTIFICATION

Arrenaptenus Schmitz
Two Afrotropical species are known (Schmitz, 1958e; Borgmeier, 1959).

Assmutherium Schmitz
A single Oriental species, known only in the female sex, is covered by Disney
(1991m).

Auxanommatidia Borgmeier
The validity of this genus is called into question by observations on the
significance of ommatidial size differentiation (Disney and Schroth, 1989).
The single Nearctic species is covered by Beyer (1963), Borgmeier, (1963d)
and Barnes (1991b). Females of four Neotropical species are keyed by
Borgmeier and Prado (1975) and the males of two species by Borgmeier
(1971a), in which the generic assignment of a further three Neotropical
species and the Nearctic species is called into question. An Afrotropical
species is described by Disney (1981h).

Beckerina Malloch
This genus is recorded from all regions except the Afrotropical. The 19
known species are keyed by Borgmeier (1971a) with a Neotropical addition
by Borgmeier and Prado (1975). The Nearctic species are keyed by
Borgmeier (1963d) with the addition of a 'Neotropical' species by Barnes
(1991a). The sole European species is covered by Disney (1983a) and the sole
New Zealand species by Schmitz (1939a).

Beyermyia Disney
The female of the single Afrotropical species is dealt with by Beyer (1965b)
and the male by Disney (1991m), who adds to the description of the female.

Billotia Schmitz
The single European species is covered by Schmitz (1953a).

Bolsiusia Schmitz
The single Oriental species, known only in the female sex, is covered by
Schmitz (1913a).

Borgmeieriphora Prado
The four Neotropical species are keyed by Brown (1993a).

Borophaga Enderlein
This genus is known from every region except the Australasian. Twenty-
two of the 28 known species are keyed by Beyer (1959a). The key to N earctic
species (Borgmeier, 1963d) includes two of the omitted species. Beyer
(1965b) adds an Afrotropical species and Borgmeier and Prado (1975) add a
Neotropical species. Colyer (1966) added an eastern Palaearctic species.
Otherwise the Palaearctic species are keyed by Schmitz (1951c), with the
British species keyed by Disney (1983a). Brown (1992b) has proposed
splitting up this genus into a number of separate genera (Godavaria,

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 GUIDE TO LITERATURE FOR SPECIES IDENTIFICATION

Latiborophaga and Peromitra, in addition to Borophaga in a more restricted

sense). As the polarities of a number of the postulated transformation series
advanced in support of this proposal are in contention, I consider it more
sensible to treat these new taxa as subgenera (at least until a consensus is
achieved). Brown adds a new Oriental species.

Bothroprosopa Schmitz
A single species from New Zealand is described by Schmitz (1939a).

Brachycephaloptera Borgmeier
A single Brazilian species is described by Borgmeier (1924a).

Brachycosta Prado
This genus is known from a single male from Brazil (Prado, 1976).

Brachyphlebina Borgmeier and Prado

A single species from Brazil is described by Borgmeier and Prado (1975).

Brachyselia Schmitz
A single Afrotropical species, known only in the female sex, is covered by
Schmitz (1929g).

Burmophora Beyer
This genus, known from Burma, Japan and North America, is sexually
dimorphic to the extent that the female was placed in a separate genus
(Crinophleba) from the male. There are four described species, but one may be
the opposite sex of another. The species are dealt with by Beyer (1958a),
Borgmeier (1967 c), Got6 (1983b) and Brown (1990a).

Calamiscus Borgmeier
The single Neotropical species, known only in the female sex, is covered by
Borgmeier (1960b).

Cataclinusa Schmitz
It is doubtful that the single Nearctic and single Neotropical species really
belong to the same genus. The former is dealt with by Borgmeier (1963d) and
the latter by Schmitz (1929g).

Ceratoconus Borgmeier
The five Brazilian species are covered by Borgmeier (1928c) and Prado
(1976).

Ceratophoromyia Beyer
A single Afrotropical species is described by Beyer (1959d).

Ceratoplatus Schmitz
The male of a single New Zealand species is described by Schmitz (1939a).

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 IDENTIFICATION

Ceratusa Borgmeier
The male of the single Neotropical species is described by Borgmeier (1967f).

Ceylonoxenia Schmitz
There are two described species from the Oriental Region, known only as
females; but I have at least three new species awaiting description. In the
meantime reference should be made to Borgmeier (1964a).

Chaetaspidia Borgmeier
The single species, known only from males from Costa Rica, is dealt with by
Borgmeier (1961a).

Chaetopleurophora Schmitz
Six Palaearctic species are keyed by Schmitz (1941c), with the four British
species also keyed by Disney (1983a). Six Nearctic species (including one
Holarctic species) are keyed by Borgmeier (1963d). Seven Neotropical
species are keyed by Borgmeier (1969b) and three Oriental species are
covered by Beyer (1958a) and Borgmeier (1962c). Brown (1992b) has
proposed raising two segregates to the generic level (Chaetocnemistoptera and
Neopleurophora). Until the grounds for doing so command consent, I
consider it wiser to treat these segregates as subgenera.

Cheiloxenia Delachambre
The single Afrotropical species, known only in the female sex, is covered by
Delachambre (1966).

Chelidophora Borgmeier
Two Neotropical species, known only in the female sex, IS covered by
Borgmeier (1950b).

Chonocephalus Wandolleck
More than 40 species have been described and named and several are known
but not named. The marked sexual dimorphism, along with the naming of
species based on the description of one sex only, has caused much confusion.
The recognition of both sexes of two cosmopolitan species, transported
around the world by man, has been recently clarified (Disney, 1991 h).
Borgmeier (1963d) keys the Nearctic species. Paulian (1958) deals with
species from Madagascar. Australasian and Oriental species are covered by
Schmitz (1929g), Bohart (1947), Borgmeier (1967f) and Disney (1986c).
Neotropical species are covered by Borgmeier (1967f), Borgmeier and Prado
(1975) and Disney (1980h). Pairs caught in copula or in reared series are
needed for the resolution of taxonomic problems in this genus. New species
should only be named when both sexes are available for description.

Clinochaeta Borgmeier
A single species, known only from males collected in Brazil, is covered by
Borgmeier (1923c).

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 GUIDE TO LITERATURE FOR SPECIES IDENTIFICATION

Clitelloxenia Kemner
Borgmeier (1964a) refers to six described species, known only as females,
from the Oriental Region. The genus is currently undergoing revision. At
least one synonym is suspected and three undescribed species are awaiting
description.

Colobomeles Borgmeier
The single Neotropical species, known only in the female sex, is covered by
Borgmeier (1927a).

Colyeria Borgmeier and Prado

The female of the single Neotropical species is covered by Borgmeier and
Prado (1975) and the male by Brown (1993a).

Commoptera Brues
The females of the one Nearctic and five Neotropical species are keyed by
Borgmeier (1971 a) and former also covered by Borgmeier (1963d). The male
of only one species is known (Borgmeier, 1926a); its characters are briefly
summarized by Borgmeier (1963d), but with the perpetuated error that the
subcosta (,mediastinal vein') is said to be absent, despite its being illustrated in
the original description.

Conicera Meigen
Twenty-five species are known. The Palaearctic species are covered by
Schmitz (1953a) and Disney (1983a, 19911). The Australasian and Oriental
species are keyed by Disney (1990k). Beyer (196~b) records the genus in the
Afrotropical Region, but no species has been described. The Nearctic species
are keyed by Borgmeier (1963d), but see also Disney (1983a). The
N eotropical species are covered by Schmitz (1929g) and Borgmeier (1968a,
1969a).

Coniceromyia Borgmeier
More than 30 species are known, mainly from the Neotropical Region but
with at least three in the Nearctic. Borgmeier (1963e) provides a key, which
needs supplementing with Borgmeier (1950a, 1969a, 1969b), Borgmeier and
Prado (1975), Prado (1976), Peterson and Arntfield (1971), Peterson (1982)
and Barnes (1991a).

Contopteryx Schmitz
A single N eo tropical specIes IS known (Schmitz, 1929i; Borgmeier and
Prado, 1975).

Cootiphora Brown
The single Neotropical species, known only in the female sex, is covered by
Brown (1993a).

Coridophora Schmitz
There are two Afrotropical species, known only as females, and covered by

271
 IDENTIFICA TION

Borgmeier (1959) (under Edaphomyia) and Disney (1982e). Since the latter
paper further material, indicating greater variation than previously sup-
posed, has allowed the following synonymy:
Coridophora convexa Schmitz, 1954b: 14.
Edaphomyia nigriventris Borgmeier, 1959: 194. Syn. nov.

Corynusa Schmitz
The male of the single Neotropical species was originally assigned to
Cyphometopis. It is described by Schmitz (1929i) and the female (as a separate
species) by Schmitz (1931b). Borgmeier (1967f) discusses the genus.

Crasilla Borgmeier
Three Oriental species, known only as females, are covered by Borgmeier
(1967a) and Disney and Kistner (1989b). It is possible that this genus is a
polyphyletic assemblage.

Cremersia Schmitz
Twelve species are Neotropical and two of these, plus an additional species,
are also Nearctic. The females have been keyed by Borgmeier (1961a). Males
can be named only when associated with their females.

Cryptophora Borgmeier
Two Neotropical species, known only as females, are covered by Borgmeier
(1935d) or Silvestri (1945), and Seevers (1941).

Cyphocephalus Borgmeier
The only known species was described from a single male from Costa Rica
(Borgmeier, 1967f).

Cyphometopis Borgmeier
Three species are known from males only, all from Brazil. Borgmeier and
Prado (1975) provide a key.

Cyrtophorina Borgmeier and Prado

A single species from Brazil is known only in the male sex (Borgmeier and
Prado, 1975).

Dacnophora Borgmeier
A key to the females of five Brazilian species is provided by Borgmeier
(1961a). The female of a Nearctic species is described by Brown (1988). Males
can be named only when procured in association with their females.

Dahliphora Schmitz
The males of the three Neotropical species and one Australasian species are
keyed by Borgmeier and Prado (1975). Females can only be named by
association with their males.

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 GUIDE TO LITERATURE FOR SPECIES IDENTIFICATION

Darwiniphora Schmitz
The two species, both from Chile, are covered by Borgmeier (1963e).

Diaclinella Borgmeier
The single species from Cuba, known only in the male sex, is covered by
Borgmeier (1962c).

Dichocerina Borgmeier
A single species from Brazil, known only in the male sex, is covered by
Borgmeier (1971 a).

Dicranodeina Schmitz
Schmitz (1929i) assigned a single Neotropical species, only known in the
female sex, to a separate subgenus of Tropophleba. The differences, however,
are such that I am herewith raising this subgenus to the generic rank.

Dicranopteron Schmitz
The five Oriental species, known only as females, are keyed by Disney and
Kistner (1990a).

Dinocercus Borgmeier
A single species from Brazil, known only in the the male sex, is covered by
Borgmeier (1971a).

Diocophora Borgmeier
The validity of this genus is doubtful (Disney and Kistner, 1989d). Two
Nearctic species are dealt with by Borgmeier's (1963d) key to females, which
also includes seven of the eight Neotropical species. The omitted species is
known only in the male sex. It is included in Borgmeier's (1959) key to the
five Neotropical species known as males.

Diplonevra Lioy
About 70 species have been described in this cosmopolitan genus. The
Palaearctic species are keyed by Schmitz (1949c), augmented by the key to
British species (Disney, 1983a). Borgmeier (1963d) keys the Nearctic species
and also (1969b) the Neotropical species. The Australasian and Oriental
species are keyed by Disney (1990e) and the Afrotropical species are covered
by Beyer (1959g, 1960a, 1965b).

Diplostylocera Borgmeier
The single Afrotropical species, known only in the male sex, is covered by
Borgmeier (1967b).

Distichophora Schmitz
The males only are known for one species from Australia (Borgmeier, 1963b)
and one from New Zealand (Schmitz, 1939a). It is doubtful, however, that
these really belong to the same genus.

273
 IDENTIFICATION

Dohrniphora Dahl
More than 100 species are known. The single Palaearctic species is virtually
cosmopolitan, through the agency of man. It is covered by Disney (1983a),
and every other key cited below. The Nearctic species are keyed by
Borgmeier (1963d), supplemented by Khalaf(1971) and Barnes (1991a). The
N eotropical species are keyed by Borgmeier (1960b), substantially aug-
mented by Borgmeier (1961a, 1969a, b), Borgmeier and Prado (1975), Prado
(1976) and Disney (1983h). The Afrotropical species are keyed by Beyer
(1959a), supplemented by Disney and Kistner (1989c) and Disney (1991a).
The Australasian and Oriental species are keyed by Disney (1990g).

Dorsochaeta Borgmeier and Prado

The single Neotropical species is known only in the female sex (Borgmeier
and Prado, 1975).

Dundophora Schmitz
A single Afrotropical species, known only in the female sex, is described by
Schmitz (1955a).

Echidnophora Schmitz
The single species, known only in the female sex, is known from the Oriental
Region and Sulawesi. Specimens from Sulawesi differ slightly from the
Oriental material, and may represent a distinct species (Disney and Kistner,
1988). The species is described by Schmitz (1916a) supplemented by Kemner
(1932a).

Ecitomyia Brues
Females of the eight Neotropical and one Nearctic species are keyed by
Borgmeier (1960b). The Nearctic species is also covered by Borgmeier
(1963d), who includes a description of the only known male.

Ecitophora Schmitz
The females of the ten Neotropical species are keyed by Borgmeier (1960b),
supplemented by Disney and Kistner (1989d).

Ecitoptera Borgmeier and Schmitz

The females of the 12 Neotropical species are keyed, along with the five
known males, by Borgmeier (1960a).

Ecituncula Schmitz
Borgmeier (1960b) keys the females of five of the seven known N eo tropical
species. Silvestri (1947a) and Borgmeier (1971a) cover the other two. The
only known male is described by Silvestri (1947a) as a 'Puliciphora'.

Ectochaeta Borgmeier
The single Neotropical species, known only in the female sex, is probably
only an aberrant species of Megaselia. It is described by Borgmeier (1958).

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 GUIDE TO LITERATURE FOR SPECIES IDENTIFICATION

Egregiophora Beyer
The single known species from Burma is covered by Beyer (1958a).

Epacteon Borgmeier
A single Australian species, known only in the female sex, is described by
Borgmeier (1963b).

Epichonocephalus Schmitz
A single species from the Bismarck Archipelago, and known only in the male
sex, is covered by Schmitz (1929g).

Epicnemis Borgmeier
Three species, from the Oriental Region, are covered by Disney (1991m).

Eremophora Borgmeier
The single species, from Panama, is known only in the male sex (Borgmeier,
1961a).

Eurycnemis Borgmeier
A single species, from Brazil, is known only in the male sex (Borgmeier,
1971a).

Euryophora Schmitz
A single species is known, from Madagascar. The female is described by
Schmitz (1915d) and the male by Schmitz (1929g).

Euryplatea Schmitz
The single Afrotropical species, known only in the female sex, is covered by
Schmitz (1941e).

Eutermiphora Lea
The single Australian species, known only in the female sex, is redescribed by
Disney (1989d).

Exochaeta Schmitz
A single species, from Argentina, is known only in the male sex (Schmitz,
1931b).

Franssenia Schmitz
Four species, from the Australasian and Oriental Regions, are known only in
the female sex. They are keyed by Disney and Kistner (1989b).

Gymnophora Macquart
More than 50 species are known. Both sexes of the British species are keyed
by Disney (1983a). Otherwise only males can be named in our present state of
knowledge. The Neotropical species are keyed by Brown (1987b). The
Nearctic species and many Palaearctic species are keyed by Brown (1987a).
Further Palaearctic species are covered by Brown (1989a), along with some
Oriental species.

275
 IDENTIFICATION

Gymnoptera Lioy
Two Palaearctic species are keyed by Disney (1983a). Three Australasian/
Oriental species are dealt with by Colyer (1956a). The single Neotropical
species, which only doubtfully belongs in this genus, is covered by
Borgmeier (1958).

Haplophleba Schmitz
A single Neotropical species, known only in the female sex, is covered by
Schmitz (1929i).

Hexacantherophora Schmitz
The single Afrotropical species, known only in the female sex, is described by
Schmitz (1914d).

Holopterina Borgmeier
A single Neotropical species, known only in the female sex, is covered by
Borgmeier (1928c, 1960b).

Homalophora Borgmeier
Six Neotropical species are known. Four are described by Borgmeier (1958)
and two are described by Borgmeier (1960b, 1969a).

Hylophora Borgmeier
Two Neotropical species are known only as females, and are covered by
Borgmeier (1960b).

Hypocera Lioy
The single Palaearctic species is covered by Lundbeck (1922), Schmitz
(1951 c) or Disney (1983a), and two N earctic species by Borgmeier (1963d).
The three Oriental species are covered by Beyer (1958a) and Borgmeier
(1967a).

Hypocerides Schmitz
Seven species are known. Three are Afrotropical, one Australasian, two
Neotropical and one from the latter two regions plus the Nearctic. A
preliminary key is provided by Disney (1991 m).

Hypogeophora Got6
A single Japanese species, known only in the female sex, is described by Got6
(1986b) (but it should be noted that he overlooked the reduced gland opening
on abdominal segment 5).

Idiophora Borgmeier
The single Neotropical species, known only in the female sex, is covered by
Borgmeier (1960b).

Indoxenia Schmitz
Three Oriental species, known only as females, are referred to by Borgmeier
(1964a). The genus is currently undergoing revision.

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 GUIDE TO LITERATURE FOR SPECIES IDENTIFICATION

Iridophora Disney
A single species is known from Sulawesi (Disney, 1986b).

Javanoxenia Schmitz
This genus is currently undergoing revision. Two Oriental species, known
only as females, are referred to by Borgmeier (1964a). Both sexes of a third
species assigned to Perissa are covered by Disney (1990j). It is transferred
herewith:
Javanoxeniageorgei (Disney, 1989g: 111.) Comb. nov.

Johowia Silva Figueroa

Seven N eo tropical species are known. Six are keyed by Borgmeier (1960b)
and a seventh species added by Borgmeier (1971 a).

Kierania Schmitz
It is probable that this genus is merely a slightly aberrant Triphleba. The single
species from New Zealand is covered by Schmitz (1939a).

Kuenbergia Schmitz
A single Palaearctic species, known only in the female sex, is covered by
Schmitz (1953a).

Lecanocerus Borgmeier
The single Nearctic species is possibly only an aberrant Megaselia. The male is
described by Borgmeier (1963d) and the female by Peterson (1987).

Lenkoa Borgmeier
A single Neotropical species, known only in the female sex, is covered by
Borgmeier (1969b).

Lepidophoromyia Borgmeier
The three Neotropical species, known only in the female sex, are covered by
Borgmeier (1925c, 1971 a) and Disney and Kistner (1989d). The genus is
probably polyphyletic, as presently conceived.

Lepta Schmitz
The single European species is described by Schmitz (1938m).

Leptilla Borgmeier
The single Australian species is known only in the male sex (Borgmeier,
1963b).

Lobochaeta Borgmeier
A single species from the Philippines, known only in the female sex, IS
covered by Borgmeier (1967a).

Macrocerides Borgmeier
Borgmeier (1969b) provides a key to the males of the seven Neotropical and

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one Afrotropical species. He commented that 'this genus is rather hetero-

geneous. It is quite possible that some of the species ... do not belong'.
Indeed the 'genus' is almost certainly polyphyletic and may only represent an
assemblage of Megaselia species with aberrant antennae. Beyer (1965b) covers
the Afrotropical species.

Macroselia Schmitz
The validity of this genus is doubtful. The three species are from Burma,
Chile and New Zealand. They are keyed by Beyer (1966).

Maculiphora Disney
The two species, from the Philippines and Australia, are covered by Disney
and Kistner (1991).

Mannheintsia Beyer
The single Afrotropical species is covered by Beyer (1965b).

Megaselia Rondani
Smith (1984) has described this genus as 'the Diptera enfant terrible'! At least
45% of the known Phoridae belong to this genus, with nearly 1400 described
species currently recognized. However, the complexities of this genus means
that they are often put to one side when new taxa are being described. The
best-known phorid fauna is that of the British Isles (Disney, 1983a, 1989a,
1991j, 1993c), in which about 70% of the species belong to Megaselia.
Furthermore the boundaries of the genus are by no means clear. This has been
discussed with reference to the Palaearctic fauna, with the return of some
species to the genus (Disney, 1990b).
It remains the case that a number of described genera of the tribe
Megaseliini are probably no more than somewhat atypical Megaselia. The
temptation to raise distinctive species groups to the generic level needs to be
strongly opposed. The type species of the genus belongs to a distinctive
segregate. Thus a policy of raising such groups to the generic level would be
likely to change the names of more than 1300 species in due course. Such a
programme of nomenclatural upheaval would serve the interests of nobody
but the pedant. The affinities of clusters of species within the genus can be
adequately expressed by means of subgenera and species groups, in due
course. Some poorly-characterized 'genera' currently excluded from Mega-
selia may be best transferred to (in many cases returned to) the genus. Other
well-defined 'genera', such as Gymnophora and Woodiphora, belong to the
same monophylum as Megaselia. Their exclusion, therefore, renders Mega-
selia paraphyletic. One could reduce these genera to subgenera of Megaselia.
While this would resolve the paraphyly, the nomenclatural upheaval is
certainly not justified (as argued in section 8.1 above). The above consider-
ations lead me to reject Brown's (1992b) proposal to designate a group of
Megaselia species as a new genus Myriophora. Even if there were consensus
regarding the validity and boundaries of this taxon, he makes no convincing
case for not treating it as a species group or a subgenus.
In preparing the keys to genera, it became apparent that the genus

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Pericyclocera Schmitz could not be characterized. The sole character, uniting

an otherwise disparate assemblage of Megaselia species, is the dorsal
deflection of the median part of the epistoma (clypeus) in the females. This
deflection is associated with a wider than usual separation of the antennae.
Borgmeier (1966a) added that the proboscis is short, the buccal opening large
and the hypostoma somewhat produced. The hypostomal character was
illustrated by Schmitz (1929g), but in fact is not found in all the species.
Indeed these additional features are found in many Megaselia species.
Furthermore, apart from the epistomal feature, the other characters are not
found in all the Pericyclocera species. For example, P. diptychogastra has a long
proboscis, very similar to that of Megaselia chlumetiae (Fig. 8.2c). However,
despite inclusion of P. diptychogastra in his key to world species, Borgmeier
(1967e) added the short-proboscis character to his diagnosis of the genus. He
then (1969b) added three further species primarily on the basis of a short,
stout, labrum, even though the degree of deflection of the epistoma was well
within the normal range for Megaselia. The evidence is that Pericyclocera is a
polyphyletic assemblage of Megaselia species in which convergent evolution
has given rise to a similar deflection of the epistoma in relation to varying
degrees of increased separation of the antennae. I have, therefore, no
hesitation in synonymizing Pericyclocera with Megaselia. The species in the
former genus are, therefore, hereby transferred to Megaselia. In the case of
Megaselia fioricola (Borgmeier, 1966a) Comb. nov. this has the effect of
causing a homonym with Megaselia fioricola Borgmeier, 1967a. The latter
species is therefore renamed (after the original collector) as follows:
Megaselia peterseni nom. nov.
= Megaselia fioricola Borgmeier, 1967a: 220.
A further 'genus' is rejected because it cannot be clearly distinguished from
Megaselia. Malloch (1914a) proposed the genus Paraphiochaeta for Megaselia
species with a row of differentiated antero-dorsal hairs on the hind tibia, in
addition to the postero-dorsals. This proposal was rejected (e.g. Schmitz,
1929g). Beyer (1958a) endeavoured to reinstate Paraphiochaeta as a subgenus.
Again this was rejected (e.g. Borgmeier, 1961a, 1968b). However,
Borgmeier (1961a) restored the genus for three Neotropical species with
antero-dorsals on the hind tibia, a longer than average costa, and other small
deviations from typical Megaselia species. However, every character cited is
to be found in numerous Megaselia species in other parts of the world. There
is no set of derived character states by which Paraphiochaeta can be
characterized. I therefore, herewith, abolish the latter genus. The three
species are assigned to Megaselia, two being returned to the genus and one
being a new combination:
Megaselia exorbitans (Borgmeier, 1961a: 82.) Comb. nov.
Brown (1988) proposed a new genus Kerophora. While these flies form a
distinct monophylum, it seems most probable that they are no more than a
segregate within Megaselia. The initial description overlooked Dufour's crop
mechanism in the females (see Disney, 1992b), thus giving rise to the
erroneous view that they should be excluded from the Megaseliini. The three

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Nearctic species are keyed by Brown (1988). Brown (1992b) has proposed
the transfer of Megaselia styloprocta and M. tristis to this genus. I agree that the
ovipositor of M. styloprocta resembles that of Kerophora. However, the male
epandrium and anal tube are very different (cf. Figs 120-123 in Brown, 1988,
and Fig. 368 in Disney, 1989a). I consider it as likely that the resemblance of
the ovipositors is due to convergence. I therefore consider it premature to
transfer these species to Kerophora. Indeed there are other species of Old
World Megaselia which are far stronger candidates for being assigned to
Kerophora. For example, the ovipositor of M. aristica is indistinguishable from
that of Kerophora and the hypopygium of the male is far closer to that of
Kerophora than that of M. styloprocta. I propose treating Kerophora as a
subgenus within Megaselia. Unfortunately this action creates two new
homonyms. Replacement names are as follows:
Megaselia brianbrowni nom. nov.
Kerophora Jerruginea Brown, 1988: 309, nec Aphiochaeta ferruginea Brunetti,
1912a: 84.
Megaselia brokawi nom. nov.
= Kerophora brunnea Brown, 1988: 310, nec Aphiochaeta brunnea Schmitz
1920c: 136.
Today the priority in the genus Megaselia must be to facilitate the naming
of specimens. It will be many years before a phylogenetic classification of the
subgroups within Megaselia can be achieved. Until then we should aim at a
stable nomenclature. Estimates suggest that only about 10% of the species
have been described for the world. While some females are very distinctive, it
is now clear that our knowledge of Megaselia taxonomy will be best advanced
by only describing new species on the basis of both sexes or the males only.
Descriptions based on females only are now an impediment. A prime
requirement for all regions of the world is user-friendly, adequately-
illustrated, keys to males. I review below the existing keys, and supplemen-
tary papers, required for each region of the world.

Afrotropical

Beyer (1965b) provides keys which, however, are seriously flawed by errors
and omissions. Bridarolli's (1951) pioneering paper is still a useful companion
(especially for the photographs of wings). Species in genera subsequently
synonymized with Megaselia also need to be considered (Colyer and Elberg,
1969; Disney, 1986b, 1990d). A species assigned to 'Pericyclocera' by Schmitz
(1940a) runs to couplet 7 at the top of page 51 in Beyer's key. Additional
species are covered by Borgmeier (1967b), Compton and Disney (1991) and
Disney (1978d, 19820, 1985b, 1989b, 1991a,b, 1991h).
Beyer (1965b) erected a new genus, 'Mollicauda', for two Afrotropical
species, one of which he had previously assigned to Megaselia. He dis-
tinguished the' genus' from Megaselia, in his generic key, on the basis of the
somewhat elongated, pear-shaped, third antennal segments in the males.
Perhaps aware that this is a character of degree only, he included the 'genus' in
the key to Megaselia species as well (loc. cit. p. 51). Here he distinguished it on

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the basis of the divergence of the antial bristles on the frons. Neither of these
characters is sufficient to exclude numbers of undoubted Megaselia species.
Indeed I can find no characters that will diagnose Beyer's genus. I
accordingly, herewith, synonymise Mollicauda with Megaselia. The two
species thus become:
Megaselia discrepans (Beyer). Comb. nov.
Megaselia subpyricornis Beyer. Orig. desig.

Australasian and Oriental

Borgmeier (1967a, e) provides keys (including a key to 'Plastophora' species

and a 'Pericyciocera' - 1967e - which are now included in Megaselia). These
keys need supplementing by Disney (1981k, 1982m, 1986b, 1987g, 1990d,
1991i, 1993b and 1994b), Disney et al. (1992), Got6 (1985a) and Johal and
Disney (in press).

Nearctic

Borgmeier (1964b, 1966a) provides keys, including two species of 'Peri-

cyclocera' which are merely aberrant Megaselia. Species formerly in 'Plasto-
phora' are covered by Colyer and Elberg (1969). The subgenus Kerophora (see
above) is keyed by Brown (1988). Important revisions and additions are
given by W. H. Robinson (1977, 1978b, 1981). Further additions are covered
by Robinson and Wisseman (1983), Disney (19810), Brown (1990b) and
Barnes (1991 a).

N eo tropical

A primary key is provided by Borgmeier (1962c). This needs augmenting

with supplementary keys (Borgmeier, 1969a, band 1971a, including a key to
species formerly placed in 'Plastophora' - on p. 65). Additions are covered by
Disney (19821, 1989h) and Wakeford and Disney (1994). The five species of
'Pericyc/ocera' covered by Borgmeier (1967e, 1969b) are only aberrant
Megaselia species. The three species of' Paraphiochaeta' will run to couplet 5
(page 301) of Borg meier's (1962c) key. Borgmeier (1958, 1961a) covers the
recognition of these species.

Palaearctic

The primary, but incomplete, monograph is the contribution of Schmitz to

Lindner's Die Fliegen der palaearktischen Region (Schmitz, 1956b, 1957a, b;
Schmitz and Beyer, 1965, 1974; Schmitz and Delage, 1974, 1981). Colyer and
Elberg (1969) cover species formerly placed in 'Plastophora'. The entire
British fauna is covered by a key to males (Disney, 1989a) supplemented by
Disney (1991j, 1993c). This key not only incorporates much revision of
European species but also adds many species not covered by Schmitz's
monograph. Additional Palaearctic species are covered by Lundbeck (1922),

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Colyer (1962, 1969), Delage and Lauraire (1970), Naumov (1979), Goto,
(1985a), Disney (1988h, 198ge,i, 19901, 1991i) and Cakar and Disney (1991).
It needs to be emphasized that undescribed species are likely to turn up in
almost any collection ofPhoridae. In many parts of the world the majority of
the species encountered may prove to be undescribed.

Melaloncha Brues
Thirty-two Neotropical species are known. Borgmeier (1971a) provides a
key, one species is more fully described by Borgmeier and Prado (1975) and
Barnes (1991a) has recorded one species from the Nearctic Region.

Melittophora Brues
The female of the single Neotropical species is covered by Brues (1928a) with
supplementary figures by Salt (1929). Brown (1993a) describes the male.

Menozziola Schmitz
Eight species have been described and a ninth reported. Beyer (1965b)
describes two Afrotropical species, Borgmeier (1961a) describes two Neo-
tropical species and Brown (1992b) transfers a third species to this genus,
from Stylusa (Borgmeier and Prado, 1975). Disney (1990b) keys the three
Palaearctic species. An undescribed species is reported from the Nearctic
Region (Brown, 1988; Brown et aI., 1991). Whether all these really belong to
a single genus remains to be demonstrated.

Mesopathusa Schmitz
Females of the three Afrotropical species are covered by Disney and Kistner
(1989c). I briefly characterize the undescribed male sex below, by describing
the first male for one of the known species.

Mesopathusa modesta Disney

Male: Frons brown, broader than high, and with three ocelli. Four
supra-antennal bristles, the lower being weaker and closer together. A pair of
pre-ocellar bristles level with the lower margin of anterior ocellus. A pair of
ocellar bristles and four other bristles on vertex. Median furrow absent, but
represented by a bare frontal vitta. With 3-4 bristles on each cheek and also on
each jowl. Third antennal segment globose, pale brown, with subterminal
arista. The latter pale brown, with the swollen base of segment 3 longer than
segment 1, which is longer than 2. The hairs of the flagellum of segment 3
well developed (about 0.015 mm long). Eyes hairy. Palps somewhat
broadened distally but narrowing to apex, being brown apically but paler
basally. With two strong apical bristles, and scattered hairs behind. Proboscis
relatively small, with brown labrum and paler, relatively simple labella.
Thorax brown, being darkest on top. Mesopleuron with a furrow and
bare. With no differentiated humeral bristle. With three bristles on noto-
pleuron (but only two on one side in one specimen). An anterior and a
posterior supra-alar, and a pre-scutellar dorsocentral bristle each side. The
scutellum with a posterior pair oflong bristles and an anterior pair of short,
fine, hair-like bristles.

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Abdomen with brown tergites bearing short hairs, which are longer at
hind margins. Tergite 6 longer than any of preceding tergites. Venter pale
brownish grey and with small, obscure, hairs on segment 4-6 only.
Hypopygium and anal tube brown and as Fig. 8.54(b,c).
Legs pale brown. All tibiae devoid of dorsal hair palisades and bristles,
apart from short apical spurs. Front leg as Fig. 8.59(c).
Wing as Fig. 8.59d. Length 1.49-1.61 mm. Costal index 0.56--0.58. Costal
ratios 0.66--0.71 : 1. Costal cilia O. 03-D. 04 mm long. Thick veins pale brown,
thin veins dark brown. No hair at base of vein 3 and no axillary bristles.
Membrane distinctly greyish-brown tinged. Haltere with dark greyish
brown knob.
Material examined: Several males, including pairs in copula, from Zim-
babwe, Harare, July and August 1991 (coli. Meg S. Cumming), at mounds of
the termites Odontotermes latericius and O. transvaalensis.

Metopina Macquart
More than 30 species of this cosmopolitan genus are known. Many
undescribed species are also known. Confusion has been caused by naming
species on the basis of descriptions of one sex only. Modern species
descriptions should be based on both sexes or the females alone. Males alone
should not be named. The discovery of polymorphism among the females of
one species (Disney, 1988d) revealed that in another species one morph had
been assigned to a separate genus!
A single Afrotropical species is redescribed by Disney and Kistner (1989c).
Other undescribed species are known from this region. The two species
known from Australia and New Zealand are covered by Disney (1994b). The
two Nearctic species are covered by Borgmeier (1963d). The Neotropical
species are keyed by Borgmeier (1969b), who also redescribes one species
under the generic name 'Typhlophorina'. Further species are added by
Borgmeier and Prado (1975). Amplified descriptions of some species are
given by Borgmeier (1969a) and Disney and Kistner (1989a). A distinctive
species, the only one known with a forked vein 3, was omitted by Borgmeier
(1969b). It is covered by Schmitz (1929g). The Palaearctic species are covered
by Disney (1983a) supplemented by Disney (1979j). The Oriental species and
tropical Australasian species are currently being revised, with the addition of
several new species. The published species are covered by Schmitz (1927e),
Hardy and Beyer (1964), Beyer (1966) and Disney (1988d, 1994b).

Microplatyphora Schmitz
The single Afrotropical species, known only in the female sex, is covered by
Schmitz (1915d).

Microselia Schmitz
This genus may only be a clade within Pseudacteon. It is still poorly known
and species recognitions are based on the female sex in the first instance. The
single Nearctic species and single Neotropical species are covered by Disney
(1982k), supplemented by Borgmeier (1969b). The three European species
are covered by Disney (1988c). The four Afrotropical species are covered by

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Disney (1983g, 1991a). I have an undescribed species from Papua New

Guinea.

Misotermes Schmitz
A species fromJava is covered by Schmitz (1938i) and a species from Thailand
by Borgmeier (1967e), who gives a key to the two females.

Morahania Schmitz
The single Afrotropical species, known only in the female sex, is described by
Schmitz (1958e).

Multinevra Disney
The male only of a single Afrotropical species has been described (Disney,
197ge). Several undescribed Afrotropical species are also known.

Myopiomyia Disney
A single species from Sulawesi, known only in the female sex, is covered by
Disney (1987g).

Myrmomyia Silvestri
The three Neotropical species, known only as females, are only doubtfully
distinct from Puliciphora. They are covered by Borgmeier (1958, 1960b) and
Silvestri (1911).

Myrmosicarius Borgmeier
The single Nearctic species is covered by Borgmeier (1963d). The females of
ten Brazilian species are keyed by Borgmeier (1931a) and another Brazilian
species is added by Borgmeier and Prado (1975).

N ecperissa Disney
A single Afrotropical species was described (as a Perissa) by Disney (1990j),
before the abolition of the Alamirinae, but has been subsequently placed in a
new genus (Disney, 1993a).

N eodohrniphora Malloch
This genus is in need of revision. Prado (1976) covers the currently
recognized species. These are seven Neotropical species and one Nearctic
species. The latter was placed in a separate genus (Borgmeieria) by Prado, but
was returned to Neodohrniphora by Brown (1988), who describes its female.
The male of this species is also covered by Borgmeier (1966a).

Neophora Borgmeier
The single Neotropical species is covered by Borgmeier (1958) with
additional notes by Borgmeier and Prado (1975).

Obscuriphora Disney
The single European species is described by Disney (1986a) and included in
the handbook on British Phoridae (Disney, 1989a).

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Odontoxenia Schmitz
The single Oriental species, known only in the female sex, is described by
Schmitz (1916a).

Oligophora Silvestri
The single Oriental species, known only in the female sex, is described by
Silvestri (194 7b).

Pachyneurella Brues
This Neotropical genus had long been synonymized with Puliciphora, when
Borgmeier (1969a) reinstated it. However, it was only readily distinguished
from the latter genus in the male sex, and on the basis of a single, seemingly
somewhat trivial, character. With the taxonomy of Puliciphora being based
primarily on the female sex the problem of recognition is compounded. I
suspect that Pachyneurella is merely a Neotropical subgenus within the
cosmopolitan genus Puliciphora.
The recognized species are covered by Borgmeier (1969a) and Borgmeier
and Prado (1975). However, the genus Puliciphora should also be considered
when trying to identify specimens, especially in the case offemales.

Palpiclavina Silvestri
The two Oriental species are keyed by Disney and Kistner (1992).

Palpocrates Schmitz
This 'genus' probably represents two further species of Triphleba with
enlarged and peculiar palps: there being several (subsequently discovered)
species ofNearctic and Palaearctic Triphleba with enlarged and aberrant palps
in the male sex. The discovery of the unknown females is likely to resolve
this. The two species, both from New Zealand, are covered by Schmitz
(1939a).

Paraphiura Beyer
This genus is known from a single male from Tasmania. It is described by
Beyer (1966).

Parasyneura Malloch
A single species from Java, known only in the female sex, is described by
Malloch (1912b) supplemented by Borgmeier (1962c).

Para term ito xenia Beyer

The single Afrotropical species, known only in the female sex, is described by
Beyer (1965c) and more fully illustrated by Kistner (1982).

Paurophora Silvestri
A single Brazilian species, known only in the female sex, is covered by
Silvestri (1947a).

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 IDENTIFICATION

Pelidnophora Borgmeier
Two species are both known only from males from Brazil. They are covered
by Borgmeier (1969b).

Penthaplus Schmitz
A single Afrotropical species is described by Schmitz (1958e).

Pericyclocerina Borgmeier
A single Neotropical species, known only in the female sex, is described by
Borgmeier (1971a).

Perissa Borgmeier
This 'genus' has proved to be the 'missing' males of the Termitoxeniinae
(Disney and Cumming, 1992). One Oriental species remains in the 'genus'
by default, until it can be linked up with its termitoxeniine female. The
Termitoxeniinae are currently being revised. In the interim 'Perissa' species
not yet re-assigned to their termitoxeniine genera are covered by Disney
(1990j).

Perittophora Disney
The single Afrotropical species, known only in the female sex, is described by
Disney (1990j).

Phalacrotophora Enderlein
About 50 species are known in this cosmopolitan genus. The New World
species are keyed by Borgmeier (1971a), who also keys the Nearctic species
separately (Borgmeier, 1963d). The Australasian, Oriental and Japanese
species are keyed by Borgmeier (1967e). The Afrotropical species are covered
by Beyer (1965b) supplemented by Schmitz (1932e). British species are
covered by Disney (1983a), and the rest of European species by Delage and
Lauraire (1974b) and Disney (1979k).

Pheidolomyia Schmitz
Three species are known from Brazil. The females are keyed by Borgmeier
(1956), who also describes the male of one species.

Phlebothrix Borgmeier
The single Afrotropical species is covered by Schmitz (1929g), under its
discarded generic name Nossibea.

Phora Latreille
More than 50 species are known, with most being north temperate in
distribution. The Nearctic species are keyed by Borgmeier (1963d), includ-
ing Holarctic species. Beyer (1965b) deals with an Afrotropical species and
(1958a) with an Oriental species. The European species are covered by
Disney (1983a) supplemented by Schmitz (1953a, 1955b) and Disney (1989a,
1989c, 1993c). The Asian species are covered by Schmitz (lac. cit.) sup-
plemented by Goto's (1986a) key to Japanese species, and additions by
Zaitzev (1977) and Michailovskaya (1986).

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Phymatopterella Brues
The five Neotropical species are keyed by Borgmeier (1971a) and a Nearctic
species is described by Barnes (1990b).

Physoptera Borgmeier
There are 13 known species in this genus, two Nearctic species keyed by
Borgmeier (1966a) and also, along with 11 Neotropical species, by
Borgmeier and Prado (1975). Whether this is really a distinct genus, or
merely a subgenus within Megaselia, remains to be demonstrated.

Placophorina Borgmeier
A single Neotropical species, known only in the female sex, is redescribed by
Borgmeier (1958).

Plastophorides Beyer
The females of the three Afrotropical species are keyed by Beyer (1959f). The
male of the type species is described by Disney (1991a), who also amplifies
the description of the female. An Oriental species is described by Disney
(1993b).

Platydipteron Borgmeier and Prado

The single Neotropical species, known only in the female sex, is covered by
Borgmeier and Prado (1975).

Plectanocnema Schmitz
The single European species is covered by Schmitz (1956b) and Disney
(1983a).

Plethysmochaeta Schmitz
This genus is currently being revised. There are supposedly eight Afro-
tropical species, but with several known in only one sex, one from Malaya
and one from Java. The last two are poorly known, but are treated by
Borgmeier (1967a). There is no key to the Afrotropical species. Reference
should be made to Schmitz (1953d, 1958e) and Beyer (1965b) in the first
instance.

Pleurophorina Borgmeier
This genus is known from a single male from Costa Rica (Borgmeier,
1969b).

Postoptica Disney
This single species is known from males only (Disney, 1987a), being
recorded from Sulawesi and the Philippines.

Pradea Borgmeier
The two N eo tropical species are keyed by Prado (1976).

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 IDENTIFICATION

Procliniella Borgmeier
The single Neotropical species, known only in the female sex, is covered by
Borgmeier (1925c, 1926c).

Pseudacteon Coquillett
More than 40 species are known in this genus. The females of the Nearctic
and Neotropical species are keyed by Borgmeier (1969b), which needs
supplementing by Borgmeier and Prado (1975) and Disney (1982k, 1991 f).
The Palaearctic species are keyed by Schmitz (1938m), with the British
species covered by Disney (1983a). Borgmeier (1967a) redescribes a species
from Java and Beyer (1966) describes a species from the Solomon Islands.

Pseudohypocera Malloch
The single Neotropical species, P. kerteszi, has been redescribed several times
under a number of synonyms. Malloch (1912b) provides a brief description in
English. Borgmeier (1930b) provides the most detailed description.

Pseudotertnitoxenia Shiraki
The single Oriental species, known only in the female sex, is too poorly
described to be adequately distinguished from other Termitoxeniinae
(Borgmeier, 1964a). A fresh description is required when new material is
available (the type material being seemingly lost).

Psyllomyia Loew
The females of the four Afrotropical species are keyed by Schmitz (1951 d).
The only male known is described by Beyer (1965b).

Ptochomyia Silvestri
Three Afrotropical species are known. The genus is currently being revised.
For the time being reference should be made to Disney (1993a).

Pulicimyia Borgmeier
The three Neotropical species, known only as females, are keyed by
Borgmeier (1960b).

Puliciphora Dahl
About 70 species are known in this cosmopolitan genus, or more if the
Neotropical Pachyneurella is merely a subgenus ofPuliciphora (see above). The
boundaries of the genus require clarification. The females of the Old World
species are keyed by Disney (1988b), along with an updated key to the
Australasian and Oriental species (Disney, 1990f) and additional information
on Afrotropical species (Disney and Kistner, 1989c; Disney, 1991a). Nearctic
species are keyed by Borgmeier (1963d), who also keys the Neotropical
species (Borgmeier, 1960b). The latter needs supplementing by Borgmeier
(1969b), Borgmeier and Prado (1975) and Disney and Kistner (1989a). The
genera Pachyneurella and Myrmomyia should be taken into consideration as
well. There is need for further revision and keys to males. Descriptions of
new species should only be based on both sexes or females. Descriptions
based on males only will merely add confusion.

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 GUIDE TO LITERATURE FOR SPECIES IDENTIFICATION

Razorfemora Disney
A single Palaearctic species is known. The male is described by Disney
(1990b). The undescribed female is described below (based on a specimen
from the Zaragoza Region of Spain, collected in 1990 by J. Blasco-Zumeta).

RazorJemora nussbaumi Disney

Female: Frons similar to male but the bristles longer and stronger (including
the outer pair on the vertex). No supra-antennal bristles, as in male.
Proboscis, palps and antennae as in male.
Thorax as male, that is with a humeral, two notopleurals, a pre-alar, an
intra-alar and a posterior dorsocentral bristle each side. Scutellum with four
bristles as in male.
Abdomen with brown tergites and brownish grey venter. The latter with
short hairs along median band on segments 2-6 and longer bristles
postero-Iaterally on segments 3-6 and also on flanks of 5 and 6. Tergites 1-6
as Fig. 8.79(a). The terminalia form a heavily sclerotized dark brown capsule
with a yellow, sharply-pointed, ovipositor (Fig. 8.79c). Internally without a
Dufour's crop mechanism and with only two rectal papillae. Beneath tergites
4 and 5 lie a pair of glands joined together posteriorly, where they appear to
discharge at rear of tergite 5 (Fig. 8.79a).
Legs as male except hind femur lacks the basal projection of the male and
the last tarsal segments tend to be more elongate and a little tapered,
especially on mid and hind legs (e.g. Fig. 8.79b). In the case of the front leg
the whole tarsus is longer than in male.
Wing 1.73mm long. Costal index 0.45-0.46. Costal ratios 2.09:0.64:1.
Costal cilia 0.08 mm long. Otherwise it and halt ere similar to male.

Rhabdomyia Borgmeier
The validity of this genus, known only in the female sex, has been called into
question by Disney and Kistner (1989b), who described an Oriental species.
Two species from Mexico were described by Brues (1925a) (under Ecitomyia
and Ecituncula). The procurement of males should resolve the taxonomic
problems posed by this genus.

Rhopica Schmitz
Only two Australasian species have been described. They are both character-
ized by Goto (1985b). However, I have a further four Australasian and an
Oriental species awaiting description, and I hope to produce a key in the near
future.

Rhynchomicropteron Annandale
A dozen species are known from the Oriental Region. The females are keyed
by Disney (1990c), supplemented by Disney (1991g, 1992d). Brown (1992a)
has shown that the, subsequently described, genus 'Gymnoselia' represents
the hitherto missing males. At present males can only be named by
association with their females, unless covered by the 1992d paper cited above
or Schmitz (1929g). Eventually it is likely that two Rhynchomicropteron species
will be found to be the females of the two described species of ' Gym nos eli a'.

289
 IDENTIFICA TION

RhyncophorOlllyia Malloch
The eight Neotropical species and two Nearctic species are keyed by Prado
(1976), and the two Nearctic species are also covered by Borgmeier (1963d).

Sphinctomyia Borgmeier
The single Neotropical species, known only in the female sex, is covered by
Borgmeier (1954).

Spiniphora Malloch
Nineteen species have been described, from the Nearctic, Palaearctic and
Oriental Regions. However, S. bergenstammi is cosmopolitan, probably
through the agency of man. The Nearctic species are keyed by Borgmeier
(1963d). The Palaearctic species are keyed by Schmitz (1941 c) supplemented
by Beyer (1958b), Borgmeier (1962c), Colyer (1969) and Disney (1983a,
19901). The Oriental species are covered by Brues (1936a), Schmitz (1940e),
Colyer (1955a) and Borgmeier (1962c).

Stenodicrania Borgmeier
A single species is known from Malaya (Borgmeier, 1967a).

Stenoneurellys Borgmeier
Only two species are known, both from Brazil. They are covered by
Borgmeier (1924a, 1927d) and Borgmeier and Prado (1975).

Stenophorina Borgmeier
The single Nearctic species, known only in the female sex, is described by
Borgmeier (1963d) and also figured by Peterson (1987).

Stethopathusa Schmitz
The single species from Sumatra, known only in the female sex, is described
by Schmitz (1921b).

Stichillus Enderlein
Twenty-five species are known in this cosmopolitan genus. An Afrotropical
species is covered by Beyer (1960b). Australasian species are covered by
Beyer (1966). Nearctic species are keyed by Borgmeier (1963d). Neotropical
species are keyed by Borgmeier (1959), supplemented by Borgmeier and
Prado (1975). Oriental species are keyed by Beyer (1958a), supplemented by
Kaneko et al. (1961). Palaearctic species are covered by Schmitz (1951c,
1953a), Beyer (1958a) and Kaneko et al. (1961). It should be noted that S.
matsumurai Schmitz is a synonym of S. japonicus (Matsumura) and the S.
matsumurai of Kaneko et al. is a misidentification of S. brunneicornis Beyer.

Styletta Borgmeier
The female of the single Neotropical species is described by Borgmeier
(1961a) and the male by Brown et al. (1991). Other species assigned to this
genus have been transferred to Trucidophora.

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 GUIDE TO LITERATURE FOR SPECIES IDENTIFICATION

Synaptophora Brown
A single Afrotropical species, known only in the female sex, is covered by
Brown (1992b).

Synclinusa Borgmeier
Two species known from Brazil are covered by Borgmeier (1971a) and
Borgmeier and Prado (1975).

Syneura Brues
There are nine Neotropical species, one of which occurs in the Nearctic also.
The species are keyed by Prado (1976).

Syneurina Borgmeier
The single Australian species, known only in the male sex, is covered by
Borgmeier (1963b).

Syntermitoxenia Schmitz
This Afrotropical genus is currently undergoing revision. The females of
three species are referred to by Borgmeier (1964a). Two species assigned to
Perissa, including a male, are covered by Disney (1990j), but are herewith
transferred to this genus:
Syntermitoxenia latiptera (Disney, 1990j:313). Comb. nov.
Syntermitoxenia lewisi (Disney, 1979g: 143). Comb. nov.

Syntermophora Seevers
The single Neotropical species, known only in the female sex, is described by
Seevers (1941).

Tarsocrates Schmitz
A single species is known from New Zealand (Schmitz, 1939a).

Termitomyia Wasmann
The single Afrotropical species, known only in the female sex, is described by
Wasmann (1900a) supplemented by Schmitz (1955f).

Termitophilomyia Schmitz
Three Afrotropical species are known. Borgmeier (1964a) refers to the
females. The male, along with additional figures of the female, of one species
is covered by Disney and Cumming (1992). The genus is currently
undergoing revision.

Termitophorides Borgmeier
The two Neotropical species, known only in the female sex, are covered by
Borgmeier (1967b) supplemented by Borgmeier (1923c, 1924a) and Silvestri
(1947a).

Termitophorina Borgmeier
The single Afrotropical species, known only in the female sex, is covered by
Borgmeier (1967b).

291
 IDENTIFICATION

Tertnitosagtna Reichensperger
Two Afrotropical species, known only as females, have been described.
Borgmeier (1959) redescribed the 'two' species, but his distinctions have
proved unreliable. Fresh material (collected by David Kistner) indicates that
T. ephippium Schmitz (1951a: 77) is a synonym of T. henningsi (Reichen-
sperger 1931: 303). Syn. nov.

Tertnitoscrofa Schmitz
Two Afrotropical species, known only in the female sex, are covered by
Schmitz (1939d) and Delachambre (1966).

Tertnitosphaera Wasmann
The two Oriental species, known only as females, are covered by Borgmeier
(1964a).

Tertnitostrotna Reichensperger
Four Afrotropical species, known only as females, have been described. A
key is provided by Delachambre· (1965). Alamira peckorum, covered by
Disney (1990j), is almost certainly the flying stage of one of these species. The
genus is currently being revised.

J'ertnitoxenia Was mann

A single Afrotropical species, known only from females, is included in this
genus (Borgmeier, 1964a), which is currently being revised. However, with
the abolition of the Alamirinae, a species of Perissa (known in both sexes) is
transferred to this genus herewith:
Termitoxenia oligoseta (Disney, 1990j: 316). Cotnb. nov.

Thalloptera Borgmeier and Schmitz

Nine Neotropical species, known only in the female sex, are keyed by
Borgmeier (1936), supplemented by Borgmeier (1971a).

Thautnatoxena Breddin and Borner

The eight Afrotropical species are keyed by Disney and Kistner (1992).

Tranopeltoxenos Brues
The single Neotropical species, known only in the female sex, is described by
Brues (1923a).

Travassophora Borgmeier
A single species from Brazil is known only in the male sex (Borgmeier,
1938a).

Trichocerina Borgmeier and Prado

This genus is known from a single male from Brazil (Borgmeier and Prado,
1975).

Trineurocephala Schmitz
The three N eo tropical species are keyed by Borgmeier and Prado (1975).

292
 GUIDE TO LITERATURE FOR SPECIES IDENTIFICATION

Triphleba Rondani
About 90 species (including those formerly placed in Citrago) are known in
this genus, which is recorded from all regions except the Afrotropical.
However, the genus is primarily Holarctic. The Nearctic species are keyed by
Borgmeier (1963d). The Palaearctic species are keyed by Schmitz (1943)
supplemented by Schmitz (1949c, 1955b), Delage and Lauraire (1970),
Disney (1983a, 1987j, 1994c), GotE> and Takeno (1983b) and Michailovskaya
(1986). Oriental species are covered by Beyer (1958a) and Australasian
species by Schmitz (1939a).

Trophithauma Schmitz
This genus in the male sex 'cannot be distinguished from Megaselia by any
remarkable character' (Borgmeier, 1963d). The females are indeed distinc-
tive, but so are numerous Megaselia females. It must remain doubtful that this
is a valid genus, rather than some Megaselia species with similarly striking
females. There are five reported species. The Nearctic species is covered by
Borgmeier (1963d). A Neotropical and Oriental species are covered by
Schmitz (1929g) (with T. splendidum being a synonym of T. portentum) and
GotE> (1984c) covers two Japanese species.

Trophodeinus Borgmeier
This genus is known from the Nearctic and Neotropical Regions. The males
have long been known under the name Bactropalpus, until correctly associated
with the females by Brown (1986). The males of seven of the nine named
species are keyed by Borgmeier (1963c), with a synonym amendment by
Brown (1986). The three known females are keyed by Borgmeier (1963d),
but two of these are only known as females. Their males are probably
described under another name.

Tropophleba Schmitz
Schmitz (1929i) assigned two N eo tropical species, known only in the female
sex, to separate subgenera because of considerable differences. The latter are
clearly more significant than the resemblances and so I have raised
Dicranodeina to the generic rank (see above).

Trucidophora Brown
Brown et al. (1991) erected a new genus for a Nearctic and a Palaearctic
species of Styletta. The former is described by Brown (1988) and the latter by
Disney (1990b).

Tubicera Schmitz
There is supposedly one Neotropical and one Palaearctic species in this
genus, but the former almost certainly belongs to a different genus (Disney,
1990b). It is covered by Borgmeier (1928c). The European species is covered
by Schmitz (1920c).

Tubiceroides Borgmeier
The two Neotropical species, known only in the female sex, are keyed by
Borgmeier and Prado (1975).

293
 IDENTIFICATION

Veruanus Schmitz
This genus is of doubtful validity, and now only includes the original
European species (Disney, 1990b). The latter is covered by Schmitz (1929g).

Wandolleckia Cook
The single Afrotropical species is covered by Baer (1953, which figures the
male anal tube as the penis in error). The female is also figured by Schmitz
(1916b) (under a synonym).

Woodiphora Schmitz
Forty species are known in this cosmopolitan genus. The Afrotropical species
are keyed by Disney (1985d), supplemented by Disney (1991h). The
Australasian and Oriental species are keyed by Disney (1989£). The single
European species is covered by Disney (1983a). The Nearctic and Neo-
tropical species are keyed by Borgmeier (1967e).

Woodiphorides Beyer
A single species, known only in the male sex, is described from New Guinea
(Beyer, 1961).

Xanionotum Brues
Thirteen species are known from the Nearctic and Neotropical Regions. The
Nearctic species are keyed by Borgmeier (1963d) and the females of the 12
species known in this sex are keyed by Borgmeier (1968a). The Brazilian
species known only in the male sex is described (as Metopina photophila, the
female being a different species) by Borgmeier (1959).

Zikania Borgmeier
The two Neotropical species, known only in the female sex, are covered by
Borgmeier (1925a, 1936).

8.4 FOSSIL PHORIDAE

The fossil Phoridae that have been studied are preserved in amber. They are
sometimes found in amber beads. Fossils in amber are reviewed by Poinar
(1992). Work on fossil scuttle flies is patchy and lacks any critical evaluation.
The described species are catalogued by Borgmeier (1968b) and updated by
Spahr (1985), the major addition being by Hong (1981). Further additions are
made by Disney (1987n), Grimaldi (1989) and Grimaldi et al. (1989). Some of
the fossil Sciadoceridae (McAlpine and Martin, 1966; Brown and Pike, 1990)
have been treated as Phoridae by some authors. Whether these interesting
flies should be included in the Phoridae is still in dispute (see Disney, 1985a,
1991m). Whereas the modern Sciadoceridae and Phoridae are readily
distinguished, the placement of some of the fossil genera is not so easy. Only
when a consensus is achieved with regard to the groundplan of the Phoridae
(8.5) can these disputes be resolved.
The genus Eophora (Handschin, 1944) is known only from puparia, and is

294
 FOSSIL PHORIDAE

therefore essentially unrecognizable in our present state of knowledge. In

addition no type species was designated, so that the name has been declared
invalid by Brown (1990c). However, if at any time a specimen were to be
designated the holotype and be given a specific epithet, the name could be
reinstated.
Grimaldi (1989) reported the genus 'Metopina' from the late Cretaceous and
lower Miocene. However, these fossils clearly have the arista dorsal in
position. This will immediately distinguish them from all modern Metopina
(Borgmeier, 1963d). Furthermore Wada (1991) has indicated that an apical
arista is the ancestral (plesiomorphic) state within the Aschiza, so that the
dorsal position of the arista in Grimaldi's fossils can be regarded as a
diagnostic feature. Therefore it seems wiser to assign them to a separate
genus, than leave standing the less probable hypothesis that the genus
Metopina has been in existence for at least 80 million years. I propose,
therefore, to assign Grimaldi's fossils to a new genus Pseudometopina. I
accordingly rename the Cretaceous species
Pseudometopina goeleti (Grimaldi, 1989: 68) comb. nov. (the type species of
the genus Pseudometopina)
Only further studies can evaluate the relationship between Metopina and
Pseudometopina. The similarities could be largely due to convergence. (Note:
Grimaldi referred to the 'ptilinal suture', in his description of these interesting
fossils. This, however, is an error. It is clear, from his excellent photographs
of the modern Metopina oligoneura, that he is referring to the incipient
transverse ridge, which is the precursor of the post-ptilinal ridge - see
Disney, 1988g, 1991 k.)

KEY TO EXTINCT GENERA OF PHORIDAE

Most of the Tertiary fossils belong to genera still found today. I give below a
key to genera known only as fossils. This key is designed only to be used for
specimens that will not run out to any known genus of modern Phoridae in
the keys above.
1. Veins M1 and M2 present as separate branches of vein M1 +2, even
though they may not be joined at the actual forking (as in Fig.
8.96b) Sciadoceridae
Vein M1 +2 (vein 4, the first thin vein) not forked (as in Fig. 8.96c)
Phoridae2
2. Vein 3 (Rs) with hairs along the length of dorsal face. Hind tibia with
longitudinal hair palisades but no isolated bristles above apical quarter 3
Vein 3 bare beyond any hair at base. Hind tibia with or without isolated
bristles above apical quarter, but without any longitudinal hair pali-
sades 4
3. Scutellum with 4 bristles Protoplatyphora Brues
Scutellum with 8 bristles on margin Protophorites Brues
4. With no isolated bristles above apical quarter of hind tibia. Vein 6 with

295
 IDENTIFICATION

sudden bend near middle, opposite and opposed to curve of vein 5 (as in
Fig. 8. 56a). (Arista dorsal in position.) Pseudometopina Disney
Hind tibia with isolated bristles above apical quarter. Vein 6 without such
a bend in the middle 5
5. Frontal bristles include a transverse row of six bristles between the four
upturned supra-antennal bristles and the four bristles of the median row
(the medio-Iaterals plus pre-ocellars) Electrophora Brues
Frons with only eight bristles between the supra-antennals and posterior
o~li 6
6. Supra-antennal bristles downturned. Third antennal segment globose,
with pre-apical arista Hypoceridites Brues
Supra-antennal bristles upturned. Third antennal segment greatly elon-
gated, with dorsal arista well before apex Rhoptrocera Hong

8.5 THE CURRENT DEBATE ON PHORID

CLASSIFICA TION
The affinities of the Phoridae have been a matter of dispute in the past, but the
current consensus places them in the section of the Aschiza which includes the
Lonchopteridae plus Phoroidea but excludes the Syrphoidea (1.3). The
Empidoidea is regarded as the sister group of the Cyclorrhapha (i.e. the
Aschiza plus Schizophora). Which of these two groups should include the
Opetiidae is unresolved (Disney, 1987d). Recently reviewed evidence, from
the study of the thoracic musculature (Ulrich, 1991) and of the larval
mandibles (Sinclair, 1992), tends to support the current consensus. The latter
view, summarized by McAlpine (1989), needs to be considered along with
more specific data relating to the inferred groundplan of the Phoridae (e. g.
Disney, 1986c,d, 1988g, 1989f, 1991m; Brown, 1992b). Rival interpretations
of these data form the basis of the current debate relating to the supra-generic
classification of the Phoridae. There is currently only a partial consensus.
The unresolved disputes mainly relate to the inferred groundplan of the
Phoridae. In particular the paper by Brown (1992b) offers views differing
considerably from my own. His paper includes many interesting new data,
which he has made the basis for some proposals for radical changes to the
current supra-generic classification. While his new data are greatly to be
welcomed, I regard most of his proposals as being premature. He himself
remarks of his hypotheses that 'the large number of character state reversals
and convergences is distressing'. His proposals, however, are worthy of
consideration. One can never prove a phylogenetic hypothesis. They serve to
direct attention to interesting data and gaps in knowledge. New data may
undermine or support hypotheses. In the case of Brown's proposals, I would
caution against acceptance of his radical revisions of the classification of
phorid genera until a consensus is reached. My disagreements are both
general and specific.
My general disagreement is with regard to the application of the outgroup
comparison method for the resolution of the polarities of postulated
transformation series of homologous character states. This issue has been

296
 THE CURRENT DEBATE ON PHORID CLASSIFICATION

discussed elsewhere (Disney, 1993e). The current consensus with regard to

the Phoroidea (e.g. McAlpine and Martin, 1966, Brown, 1992b) places the
Sciadoceridae as the sister group of the Phoridae, the Ironomyiidae as the
sister group of the Sciadoceridae, and the Platypezidae as the sister group of
the lronomyiidae. Oldroyd (1964) remarks that Bezzi called Sciadocera
ruJomaculata (Fig. 8.95) 'the most wonderful fly in the world' 'because it
combines features of Empids, Dolichopodids, Phorids, and Platypezids, at
one of the most obscure points in the evolutionary plan of the flies'. It is
precisely because it is such a mosaic of ancestral (plesiomorphic) and derived
(apomorphic) character states that Sciadocera is such an intriguing insect.
Which are the derived states and which the ancestral?
The basic problem, in relation to two postulated homologous states, A and
B, which both occur in the Phoridae, is whether A evolved from B or B from
A. The outgroup method examines the sister group for the presence of A and
B. If A is restricted to the Phoridae but B is not, it is argued that B is the
groundplan state for the Phoridae and A is the derived state. In applying this
method Brown tends to compare the Phoridae with the Sciadoceridae, then
the latter with the lronomyiidae, and then this family with the Platypezidae,
and finally the latter with the Empidoidea. By contrast I tend to use the
Sciadoceridae plus the lronomyiidae plus the Platypezidae (and, if necessary,
the Empidoidea and other Brachycera as well) as the outgroup for the
purposes of comparison. This difference can give rise to very different
interpretations. For example, tergite 8 of the female abdomen is present in
some Phoridae and absent in others. It is absent in both the Sciadoceridae and
Ironomyiidae, but present in Platypezidae and Empidoidea. Brown con-
cludes that the groundplan of the Phoridae has no tergite 8. He then
postulates that it evolved afresh (possibly as an 'underlying synapomorphy'
or 'atavism' which re-emerged after being dormant for millions of years) in
many Phoridae, as a 'new', derived, state. By contrast I conclude that tergite
8 is part of the groundplan of the Phoridae, and has been independently lost in
Ironomyiidae, Sciadoceridae and some Phoridae.
I have defended my approach (Disney, 1993e) with reference to the
prevalence of mosaic evolution (without which all phylogenetic reconstruc-
tion would be impossible - as emphasized by Ax, 1987) and by pointing to
the long separation of the Sciadoceridae and Phoridae. They have been
separate for at least 80 million years (McAlpine and Martin, 1966; Grimaldi,
1989). The re-emergence of a lost character is, of course, a possibility.
However, I maintain that it should never be assumed and this assumption
then be employed in the construction of a cladogram. The latter should be
produced without the aid of the disputed transformation series, which is then
subsequently mapped onto the cladogram to test the hypotheses of dis-
appearance followed by re-emergence of one of the character states of the
senes.
Some of Brown's specific proposals I consider to be insufficiently justified.
An example concerns the hypandrial bristles (Fig. 8.14a). Brown postulates
that these are a novelty, and then proposes the transfer of Triphleba from the
Phorinae to the Metopininae. It is true that Sciadocera lacks such bristles.
However, several hypandrial bristles are the norm in Ironomyiidae and

297
 IDENTIFICATION

Platypezidae. An alternative homology is that hypandrial bristles in Phoridae

represent the last remnants of the gonopods (Fig. 8.15). Either interpretation
leads me to the conclusion that hypandrial bristles are a groundplan feature of
the Phoridae (Disney, 1994c). There are other examples of unresolved
polarities of postulated transformation series. For example, a two-segmented
palp (e.g. Fig. 8.41b,c) was proposed as a groundplan feature of the Phoridae
(Disney, 1988g). Brown (1992b) counter-proposes that a one-segmentd palp
is the groundplan state, because it is so in Sciadocera. However, evidence from
the subfamily Thaumatoxeninae (cf. Figs 8.66b and 8.89b) indicates that a
two-segmented palp is a groundplan feature of this subfamily at least (Disney
and Kistner, 1992).
Apart from polarity problems, there are differences of opinion regarding
the homology, or otherwise, of critical character states. For example, Brown
(1992b) proposes that the dorso-lateral sclerotized patches on the seventh
abdominal segment of Dohrniphora females (Fig. 8.40c) and those of
Aenigmatias females (Fig. 8.21b,c) are a synapomorphy (a shared, uniquely-
derived, character state). I have counter proposed (Disney, 1993c) that in
Dohrniphora these patches are the lateral portions of tergite 7, and that the
essential novelty is the loss of the sclerotisation of the median part of the
tergite. By contrast in Aenigmatias I consider tergite 7 to be absent and the
postero-lateral sclerotized patches to be a novel development, associated with
the musculature operating the undisputed novelty of the two series of hooks
at the rear of the segment. In short, I reject the postulated homology between
the patches in the two genera. Likewise, the homologies of the thin veins of
the wing are in dispute (Disney, 1991c, 1991m; Brown, 1992b). The
homologies I believe to be best supported by the available evidence are shown
in Fig. 8.96. The principal difference from Brown's preference is that he
regards vein 4 (the first thin vein) as being Ml and vein 5 as M2. This
difference of interpretation affects the proposed family assignments of certain
Cretaceous genera. According to my interpretation the mutant shown in Fig.
1.1 b represents a throw-back mutation, restoring the lost outer cross-vein of
the phorid ancestors. This cross-vein is retained in many other families of
Diptera (e.g. Fig. 8.96a).

298
 THE CURRENT DEBATE ON PHORID CLASSIFICATION

Figure 8.16 AdelopteromYla, male (a) RIght wmg (b) Frontal vIew of head

299
 IDENTIFICATION

Figure 8.17 (a) Adelopteromyta male hypopygmm (b) A parvlpenms female, nght
wmg

300
 IDENTIFICATION

(b)

Figure 8.18 (a) Adenophora pulchella female, abdominal tergites 4-6. (b) Auxanom-
matidia medleri female, left front leg.

301
 IDENTIFICATION

Figure 8.19 Aenictacantha setifera female, anterior faces oflegs. (a) Hind femur, tibia
and metatarsus. (b) Front leg (omitting coxa and trochanter).

302
 IDENTIFICATION

(a)

Figure 8.20 Aenictacantha setiJera female. (a) Frontal view of head. (b) Dorsal view of
thorax and abdominal tergites.

303
 IDE NT IFICAT ION

(a)

(c)
I
Figure 8.21 Aemgmatzas (a) A lubbock! male, fight wmg (b), (c) Termmal
abdommal segments from above m A lubbockt (b) and A franzt (c)

304
 IDENTIFICATION

Figure 8.22 Aemgmatzas lubbockl (a) Female from above (termmal segments
withdrawn) (b), (c) left and nght faces of male hypopygmm

305
 rDENTIFICATION

......'

(a)

(b) -

(e)
. •• , ,..;-0:.
~.....

Figure 8.23 Aemgmatopoeus kohlt female, rIght palp from below. (b), (c) Aemgma-
tlstes latlfrons female, head from below (b) and head and thorax from above (c).

306
 IDENTIFICATION

Figure 8.24 Aemgmatlstes herero female, head facmg left, vIewed from above (a) and
below (b)

307
 IDENTIFICATION

... ',: "

, .

, "
, '
... ,
"..'..
,',
.'

:",': "
, "
', ' ..
:' . '
',:." .··i·'·.

(a)

(b)

Figure 8.25 Apocephalus male hypopygla, left faces (a) A borealts (b) A lacey I

308
 IDENTIFICATION

(a)

Figure 8.26 Assrnuthenurn rostra turn female (a) Head and front of thorax from
above (b) Head, thorax and front of abdomen from side S = positIOn of antenor
thoracIc spiracle on side, beneath overhang of thoraCic notum

309
 IDENTIFICATION

(a)

Figure 8.27 Auxanommatldia medlen male. (a) Left face ofhypopygmm. (b) Frontal
view of head.

310
 IDENTIFICATION

Figure 8.28 (a) Auxanommatidia medleri female, frontal view of head. (b) Beckerina
umbrimargo, left humerus and notopleuron of male (H = humerus, M = mesopleuron,
N = opening of notopleural gland, S = spiracle, T = tegula).

311
 IDENTIFICATION

(a)

.. ~

(b)

Figure 8.29 (a), (b) Beyermyia spinulicosta: (a) axillary bristle; (b) tips of costa and
vein plus all of vein 3. (c) Chonocephalus heymonsi, right wing.

312
 IDENTIFICATION

Figure 8.30 Chonocephalus heymonsl, male hypopyglUm from below (a) and from
side (b)

313
 IDENTIFICATION

(a)

~~~..: :~
:,..':" ,~.::
":::;~y/~.i/,?r:?:~:~>. :':"~':>~':':::'.'~)-:'.,:': '
'.:.. ~:': :':~':-~.~.~.::'; :: :",

(e)

Figure 8.31 Comcera males (a) C malae, thlrd antennal segment and ansta
(b) C craSSlcosta, outer (left) and mner face (nght) of nght clasper (surstylus) of
hypopygmm (c) C schmttmannl, nght wmg

314
 IDENTIFICATION

I
(a)

Figure 8.32 Cotlicera crassicosta male. (a) Anterior face of hind tibia. (b) Antenna.

315
 IDENTIFICATION

(a)

(c)
(b)

Figure 8.33 (a) Penssa paktstanensls male, left face of hypopygmm (b)-(d)
Clttelloxenta peradentyae (b) base of brIStle on abdomen, (c) postenor face of hmd
femur, (d) left face of head

316
 IDENTIFICATION

" .. ',' I

-=:5L\il i':, -::ii 1

. ..... ' .

.... : ",, ' .

..

. <~ .>.: ..' .: ~:' >:>:".::- .·z ·. ·.

. • ' • • • •. '.0 '", '.' ,
',. '. . ....

. ... ..

(b)

Figure 8.34 Crasilla setifera female. (a) Right wing . (b) Abdomen from above, from
rear oftergite 2 to segment 6.

317
 IDENTIFICATION

, ~\. ' . ~.. "

...... -

"'-.

. , ..........
.....

(a)

(b)

Figure 8.35 Crasilla longiseta female. (a) RIght wing. (b) Abdominal segments 2--6
from above.

318
 IDENTIFIFfCATTON

(a)

." ........,... " ... .

' .... ,. .......

.. .
~,~

......

(b)

Figure 8.36 Dahltphora slgmoldes male (a) Thud antennal segment (b) Right wmg

319
 IDENTIFICATION

(c)

Figure 8.37 Dtcranopteron setlpenntS female (a) Right palp, Inner face (b) Head from
left side (c) Abdominal segments 1 and 2 from above

320
 IDENTIFICATION

Figure 8.38 Dlocophora appretiata (a) Left palp offemale from above (b) OVIpOSitor
from above (c = vestigial cercus) (c) Male hypopyglUm, left face (E = epandnum,
Tl0 = terglte 10, S = surstylus)

321
 p

(e)
(f)

Figure 8.39 Diplonevra males, details oflegs. (a)-(e) Posterior faces of bases of hind
femora and trochanters: (a) D. abbreviata; (b), (c) D. jlorea; (d) D. nitidula; (e)
D. jimebris. (f) D . watsoni, posterior face of hind tibia (P = hair palisades).

322
 IDENTIFICATION

.. '

(b)

(c)

Figure 8.40 Dohrnlphora . (a) D. trud,ae male, posterior face of base ofhmd femur
(P = sensory pIt). (b) D. kzstneri female, aXIllary brIStle from above. (c) D. cornuta
female, rear of abdominal segment 6 and segment 7, from sIde.

323
 (a)

(c)
Figure 8.41 (a) Dundophora fimbriiterga female abdominal tergites 1-5 from right
side. (b), (c) Dohrniphora diminuens female palp, inner face (b) and from below (c).

324
 2T '7~
\ (b)
, I
\

3T

4T

5T
(a)

s
~-
w
A

(d)

Figure 8.42 ECltophora females (a) E brevlptera abdomInal tergItes 2-5 (2T-5T)
(b) E co/leglana nght WIng (c), (d) E brevlptera (c) left two-thirds offrons and left
eye, (d) nght half of thorax from above (H = head, S = scutellar bnstle, W = WIng
rudiment, A = abdomen)

325
 fDENTIFICATJON

'-', .. -';.:\. ...,

(a)

".

Figure 8.43 (a) Ecltoptera humeralls female, nght wmg (b), (c) Eplcnemls females,
antenor faces of mid femora (b) Eplcnemls sp (probably E testIs), (c) E flavldula

326
 IDENTIFICATION

(b)

Figure 8.44 Eplcnemlsfiavldula female. (a) From above. (b) Head and thorax from
below (legs omItted).

327
 IDENTIFICATION

(a)

,- ., .
. ~ .
.::.:..:...
. .... -_
. -:;':,:>~~.'.;::."
'.' . .
"~:t':· . . . "
... .-.
,.: ~

':::. ~ :.:'"

- ---_.. .: ::':'?~~;:>::'
(b)

Figure 8.45 (a) Ep,cnem,s testIs, left face of male hypopyglUm (b) Franssema
malayszae female, abdominal tergltes 2-5

328
 IDENTIFICATION

(c)

Figure 8.46 (a), (b) Eutermlphora abdomina/Is female (a) side view of whole fly
(b) frontal view of head (c) Hypocera mordellarta, antenor face ofhmd tibia

329
 IDENTIFICAT ION

(a)

JAn..:-
...... D
,,":
r.··'\.\ N
...::::.:.::.:....
~
. ",~- 'Y
.~
~) --~========~----

Figure 8.47 (a), (b) Gymnophora males, notopleuron of left sIde (H = humerus,
C = no to pleural cleft, D = duct of notopleural gland, AN = anterior notopleural
bnstle) (a) G quartomollls, (b) G arcuata (c) Gymnoptera longlcostalzs male hypo-
pygmm, left face

330
 IDENTIFICATION

Figure 8.48 (a) Hypocera mordellana male, left face of hypopygmm (b),
(c) Hypocendes nearctlcus male (b) left palp, (c) left face ofhypopygmm

331
 IDENTIFICAT ION

(a)

(b)

(d)

(e)

Figure 8.49 (a) Hypocendes nearctlcus male, basal region offront half of wmg (b)-(e)
Indophora clarkI (b) base ofhmd femur of male, (c) male antenna, (d) tarsal segments 4
and 5 offront leg offemale, (e) left face of male hypopyglUm

332
 IDENTIFICATION

(a)
(b)

(c)

Figure 8.50 Indophora clarki female (a) Antenna (b) Tarsal segments 4 and 5 of
mIddle leg (c) Tarsus ofhmd leg (d) RIght wmg

333
 IDENTIFICATION

(b)

(c)

Figure 8.51 (a) Megasella brokawl OVIposItor, left face (b), (c) Macultphora
aenzctophtla parts ofhmd leg of female (b) posterIor face of ubI a, (c) antenor face of
femur

334
 IDENTIFICATION

(a)

\ r , 1

\ i \ \ I
'I '\
I
(b)

Figure 8.52 Lepldophoromyw lablda female (a) Right wing (b) Abdominal tergltes
1-5

335
 IDENTIFICATION

(b)

:~l~,~,,
~:.
__ ......

'" "

(c)

....

(d)
Figure 8.53 Macultphora males (a), (b) Posterior faces offront femora of M setifera
(a) and M aemctophtla (b) (c), (d) Right wmgs of M setifera (c) and M aemctophtla (d)

336
 (c)

Figure 8.54 (a) Menozziola obscuripes male, left face of hypopygium. (b), (c)
Mesopathusa modesta male, hypopygium from left (b) and right (c) sides.

337
 IDENTIFICATION

I I
Cd)

Figure 8.55 Mesopathusa females. (a) M. cloveri wing rudiment from above (T =
thoracic notum, A = abdomen, W = wing rudiment). (b) M. cloveri abdominal
tergites 4-6. (c) M. modesta abdominal tergites 4-6. (d) M. modesta head and thorax
from above.

338
 IDENTIFICATION

(a)

(e)

(b)

(e)

Figure 8.56 (a) Metopina crassinervis female, right wing. (b)-(e) Microselia: (b) M.
beaveri female, segments 3-5 of front tarsus. (c)-(e) M. bingana: (c) female segments 4
and 5 offront tarsus; (d) female left antenna; (e) male segments 4 and 5 of front tarsus.

339
 IDENTIFICATION

Figure 8.57 Microselia. (a) M. beaveri female, frontal view of head. (b) M. bingana
male, left face ofhypopygium.

340
 IDENTIFICATION

(a)

--

".'.",

(b)

Figure 8.58 Mlcroselta brngana female, OVIposItor from left sIde (a) and above (b)

341
 IDENTIFICATION

(a)

(b)

(d)

Figure 8.59 (a), (b) Metopina ciceri male, right wmgs of two dlfferent specimens. (c),
(d) Mesopathusa modesta male: (c) front leg, anterior face; (d) right wing.

342
 IDENTIFICA TION

--
Figure 8.60 (a)-(c) Anterior faces of hind tibiae: (a) Multinevra macropygidia male;
(b) Aenigmatias lubbocki female; (c) Psyllomyia braunsi female; (d) Multinevra macro-
pygidta male, right wing.

343
 IDENTIFICATION

(a)

." ...--

(b)

Figure 8.61 Multtnevra macropygldla male (a) Frontal vIew of head (b) Left face of
hypopygmm

344
 (a)

(b)

Figure 8.62 MyoplOmylQ harman! female (a) Right wing (b) Frontal view of head

345
 fDENTIFICA TlON

(a)
o
(b)

'~ijf?;~'~\;f:;,:{:',';,,: > . ' .

.: ...•. : "

(e)

,: : . .: ' .
.' :' .'." .' ~ ,
o.
-.; ',' .:.:', ~ , ~"::::':: "
000

. ",',.
. '.'
. . " ,. , - ' "
.'

:. o·:,'• 0• •

':':'" .
. ,'.
" " '.'

(d)

Figure 8.63 NecpeYlSsa tmgle/. (a) Left antenna of male. (b) Front leg of male.
(c) Right wmg of male. (d) Right wmg offemale.

346
 IDENTIFICATION

(a)

.........

-...., ",

~\-~../' ..
\\,
-')

(d)

Figure 8.64 Females of Necpenssa and Ptochomyza (a) P cummlngae left antenna
(b) N tlnglel left antenna (c) P cummlngae tarsal segments 1 and 2 of front leg
(d) P aJra, left face offront part of thorax (e) N tlnglel proboscIs and fight palp

347
Figure 8.65 Obscurtphora sheppard! (a) Female frons (b) Male, anterIor face ofhmd
leg (c) Male, rIght wmg (d), (e) Left and rIght faces ofhypopygmm

348
 IDENTIFICATION

(a)

~-:. i • ~

(b) .. .; /
I •

(d)

Figure 8.66 Palp,davma k,sfnen male (a) AnterIor face of mId tibIa (b) RIght palp
(c), (d) RIght and left faces ofhypopyglUm

349
 IDENTIFICATION

I
(a)

I
(b)
(el

Figure 8.67 Palplclavma klstnen (a) Male, frontal view of head (b) Female, left
palp (c) Female, left face of thorax

350
 IDENTIFICATION

Figure 8.68 Palplcalvma females (a) P tonkmenm, rIght wmg (b) P ktstnen frontal
vIew of head

351
 IDENTIFICATION

(. (a)

(b)

Figure 8.69 (a) Necperissa tinglei male hypopygium. (b)-(d) Perittophora couturieri
female: (b) left palp from above, (c) front leg, (d) rIght wmg.

352
 IDENTIFICATION

(a)

(c)

(b)

"'"
' " ' '''

(d)

Figure 8.70 (a) Phalacrotophora berolinensis female, abdominal tergite 3. (b) Plecta-
nocnema nudipes male hypopygium, left face. (c), (d) Postoptica platypezoidea male: (c)
dorso-lateral view of head, (d) right wing.

353
 IDENTIFICATION

[
(b)
....

Figure 8.71 Piastophondes setuilClncta male (a) Hypopygmm, left face. (b) AbdomI-
nal segment 6 from below

354
 IDENTIFICATION

(c)

(d)
Figure 8.72 PlastophoYldes setullclncta tarsal segments (a) Female front leg (b) Male
front leg (c) Female hmd leg, segments 4 and 5 (d) Female mIddle leg, segments 4 and
5

355
 IDENTIFICA TION

(b)

Figure 8.73 (a) Plectanocnema nudipes male. anterior face of hind tibia. (b)Postoptica
platypezoidea male. frontal view of he ad.

356
 IDENTIFICATION

': :';~'::: ::
.. . '.. ,. .
. ., ' , '

(al

Jt=<
...:-~
(b) >
, - - - - -_ _ ----., (e)

:'

Figure 8.74 (a) Pseudacteon spatulatus male, left face of hypopyglUm (b)-(d)
Pseudohypocera kerteszl (b) hmd tarsal claw of male, (c) base of thIrd segment offemale
ansta, (d) postenor face ofhmd femur of male

357
 IDENTIFI ATION

(a)

Figure 8.75 (a) Psyllomyta braunsl female, nght wmg (b) Pultclphora rufipes male,
left face ofhypopyglUm

358
 IDENTIFICATION

..;~.;i.;;?~;~:~lt~~:~;IJ.t>
Yo ""_. - - -_ _ _ _- --

........
-

(a)

(b)
-D
Figure 8.76 Ptochomyia cummingae male. (a) Right wing. (b) Left antenna. (c) RIght
front leg.

359
 IDENTIFICA TlON

Q" ..
:.)

(a)

(c)
--" ~

Figure 8.77 Ptochomyta cummmgae female (a) Right wmg stump (b) Haltere
(c) Front end ofleft face of thorax (d) ProboscIs and antenna, left face (e = eye)

360
 IDENTIFICATION

(a)
I I

.....

(c)

Figure 8.78 (a), (b) Pultctphora females, abdommal tergltes 4 and 5: (a) P. knzghtl,
(b) P. rosei. (c)-(e) RazorJemora nussbauml male: (c) posterIor face ofhmd femur and
tIbIa; (d) left face ofhypopygmm; (e) left antenna.

361
 IDENTIFICATION

-~
(b) "

(c)

I~ (d)
(a)

Figure 8.79 (a)-(c) Razoljemora nussbaumt female (a) abdommal tergltes 1-6,
(b) antenor faces of mId-tarsal segments 4 and 5, (c) OVIposItor from above,
(d) Rhabdomyta klStnerorum female, left wmg

362
 IDENTIFICATION

)11 1 ~\ t
~/: i
(a)
\ '

: :
(b)

Figure 8.80 (a) RhabdomYla k,stnerorum female, abdommal tergItes 2-6 (b) Rhyncho-
mlcropteron beaven female, proboscIs

363
 IDENTIFICATION

:.

(b)

(a)

(d)

Figure 8.81 Rhychomicropteron necaphidiforme female. (a) Proboscis. (b) RIght side of
frons. (c) Anterior face ofhmd femur. (d) Right wing.

364
 IDENTIFICATION

,'. ' .

,-. -',' -:-,:-,

.'
(a)

(b)
(c)

.' ... .
. . . : .':.: " .' . ' ..
. :: : :: .: . . . ' ,', ':. '.- .'
. -.' . . . . . .' ...... -'
.. -:',' - .. :. :.'
.
',

. .
. . . : :. ',' .. ... . ..
.
:: . ..
'

: .. ::.. . ."
"

, , -',. , :"
(d)

----..!

f~~J-----------------
(e)

Figure 8.82 Syntermaoxenta (a)-(d) Males (a) S [atlptera, rIght WIng, (b) S lewlSl,
left face ofhypopygmm, (c) S latlptera, left face ofhypopygmm, (d) S lewIsI, rIght
WIng, (e) S aplOgaster, abdomInal brIstle offemale

365
 ID ENTlFICATrON

........
"7)
.

. ,
.-
'.
,

- -
.'

.- . .
' .

. ' . . ',' . .

, ,- " . .
.... -
(a)

,.',

...~ . . . . .......... .
~ ..... ~. . .
. .

(b)

Figure 8.83 Syntermltoxenra females, rIght wmgs (a) S latlptera, base of wmg from
below (b) S latlptera (c) S leW1Sl

366
 IDENTIFICATION

(d)

(e)

Figure 8.84 (a), (b) TermltophtiomYla braunsl antennae: (a) female, (b) male.
(c) Termltoxema ollgoseta male, left antenna (at a larger scale than (a) and (b)). (d) T
ollgoseta male, front leg. (e) Termttosagma hennmgsl female, right wmg stump

367
 IDENTIFICATION

(a)

(b)

(d) -

(c)

Figure 8.85 Termltophtlomyta braunst (a) Female, rIght wmg (b) Male, rIght wmg
(c) Male, rIght front leg (d) Male, anterIor face ofhmd femur

368
 IDENTIFICATION

(a)

.•.. ·<.:i".;,.· ~;:;6:':,~t~·~lfii~l~~

(d) ·--:-------~--::.tj> ··.- '.,.
Figure 8.86 Termltostroma sp ,female (a) Bnstles at base of abdommal segment 2
(b) Front leg (c) RIght wmg (d) Base of nght wmg from below

369
 fDENTIFICATION

(b)

(d)

Figure 8.87 Termlfoxema ollgoseta (a) Female, rIght wmg (b) Female, brIstles at
sIde of abdommal segment 3 (c) Male, left face ofhypopyglUm (d) Male, fight wmg

370
 IDENTIFICATION

(b)

:. " -:".:-
..:. . ,',. , '-
. ,~.' ., - .. '. - .. ' . '-
:.
'

. . ..
,.,-; . . .:- :: ...
. .'. -'.',: : : ~. ' . ' '. :" .
';' ,.' ::. . . -' : - ..

(d)

Figure 8.88 Javanoxenta george!. (a) RIght wmg of male (b) left face of male
hypopygmm, (c) left antenna of male. (d) RIght wmg offemale.

371
 IDENTIFICA TION

(b)
Figure 8.89 Thaumatoxena andremll male (a) Frontal vIew of head (b) RIght palp

372
 IDENTIFICATION

Figure 8.90 Thaumatoxena andretnll female, from above

373
 IDENTIFICATION

.. ., ' .. :. . ..
... . .
"
.. . .
: , '. ,'

:"::'.:.:. . :. . '>~ ~ .::... >.':--" .

: ":. ' '. '.

'"
. . . :, . , '
, ',. " "
','
.
;
,

(b)

Figure 8.91 (a) Thaumatoxena grasset female, antenor face of hmd femur.
(b) Tructdophora ewardurskae male hypopygmm, left face.

374
 IDENTIFICATION

(d)

Figure 8.92 Woodiphora. (a) W. distans male, front tarsus. (b) W. dormanae male front
tarsus. (c) W. multinevra female, anterior face of hind tibia. (d) W. parvula female, hind
tibia.

375
 IDENTIFICATION

.,. P

ca .::::
Ca) 1------/

(b)

···x
l ....
.....
......

..'
\..,

....-. - - -
~-.-

(e)

Figure 8.93 WOodlphora (a) W pallldmervls female, abdomInal tergltes 5-7 (b)
W santoshl male, left notopleuron (c) W ajYotroplca male, left notopleuron
(C = notopleural cleft, D = duct from notopleuron gland, AN = antenor notopleural
bnstle, S = sac-lIke notopleuron gland plus duct, T = tegula at base of WIng)

376
 iOENTIFlCATION

(a) (b)

(d)

(c)

Figure 8.94 (a)-(c) Spzmphora bergenstammz female (a)-(b) dorsal and ventral views
of abdommal segment 7, (c) last tarsal segment of hmd leg (d) Stzchzl/us sp
(undescnbed) male, frons

377
 IDENTIFICATION

Figure 8.95 Sciadocera ruJomaculata male. (After Oldroyd, 1964.)

378
 IDENTIFICATION

R1

- -____------------~1~1 + 2

M3 + 4

1A

Figure 8.96 Proposed homologies of the wing veins. (a) Dolichopus pennatus
(Dolichopodidae). (b) Sciadocera ruJomaculata (Sciadoceridae). (c) Megaselia rufipes
(Phoridae).

379