Transactions of the Society for British entomology

QL

461

S69

ENT

PART 6.

iBIM

f/

T RAN S ACTIO

OF THE

1938

• nli: 1¥

Society for British
Entomology

editor :

F. J. KlLLINGTON, B.Sc., A.L.S., F.R.E.S.,

F.S.B.E.

ASSISTANT EDITOR :

B. M. Hobby, m.a., d.phil.. f.r.e.s.

Date of Publication : 9th November, 1938.

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Courtship and Allied Problems in Insects.
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Wegener’s Theory of the Movement of
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COLLEMBOLA.

ODONATA.

HEMIPTERA.

TRANSACTIONS OF THE SOCIETY FOR
BRITISH ENTOMOLOGY

Vol. 5. 9th NOVEMBER, 1938. Part 6.

THE DESCRIPTION OF THE PUPARIA OF
FOURTEEN BRITISH SPECIES OF
SPHAEROCERIDAE (BORBORIDAE, DIPTERA).

By W. H. Goddard, D.I.C.

(East Anglian Institute of Agriculture, Chelmsford, Essex).

Few descriptions have been recorded of the eggs and puparia
of the British Sphaeroceridae. Richards (1930', p. 319) gives
references to descriptions of the larvae and puparia of four
species, and the identification of two of these species appears to
be very uncertain.

In this paper I have attempted to describe the eggs and
puparia of the species which I have been able to breed, and also
record the localities and habitats of these and other species
whose puparia I was unable to obtain.

The work was carried out at the Imperial College Biological
Field Station, Slough, Bucks, during the winter months of 1933
to 1935. I wish to thank Prof. J. W. Munro for the facilities
afforded to me there, and Dr. O. W. Richards for the identi¬
fication of the material. My thanks are also due to Mr. G. E. J.
Nixon for the identification of Phaenopria cameroni Kieff., which
emerged from one of the puparia.

Methods.

The flies were bred on several different materials, including
decaying lawn-mowings, garden refuse, and the dung of cows,
deer, sheep and mice.

Sterilisation of the materials used for food for the larvae
was found necessary, to kill all other eggs and larvae, so as to
be sure that the puparia described from the flies bred really
belonged to the same species and were not already in the
material.

236

[November,

Flies collected in copula were placed in 3 in. glass tubes
with an inch of moist material at the bottom. These tubes were
covered with muslin and kept at laboratory temperature, and
given a few drops of water whenever they showed any signs of
becoming dry.

Several species were treated in this manner, but the only
three that were successfully bred from the eggs to the adults
were Leptocera silvatica (Meigen), L. heteroneura (Hal.) and
Sphaerocera subsultans (L.). Another method of obtaining the
puparia was then tried.

Cow-dung, deer-dung, mouse-dung and other materials were
collected and examined for larvae and puparia. The puparia
were removed and placed separately in 3 in. glass tubes on
damp cotton wool, and the larvae were returned to their various
foods. By examining the material daily, considerable numbers
of puparia were obtained. When the flies emerged they were
identified ; this eliminated any possibility of describing puparia
which did not belong to the species bred.

This method proved more successful, and the puparia of the
following species were obtained: — Leptocera palmata (Richards),
L. crassimana (Hal.), L. t alp arum (Richards), L. manicata
(Richards), L. vagans (Hal.), L. bequaerti (Villen.), L. melanin
(Hal.), L. fungicola (Hal.), Copromyza stercoraria (Meigen)
and C. glacialis (Meigen).

Difficulty was experienced in keeping the adults alive in an
attempt to obtain the eggs of these species. Many died before
they became mature specimens. This was partly due to the
fungi which appeared in some of the tubes, and also to very
large numbers of nematodes which appeared in the tubes and
destroyed larvae, pupae and occasionally the adults. When the
nematodes attacked the pupae, the adult flies failed to emerge,
leaving the puparia unidentifiable.

Descriptions of the Puparia : Introductory.

The anterior spiracular processes afford one of the most
obvious differences in the puparia of all the species examined,
but this character alone is insufficient to separate the closely
allied species with any certainty. Smaller differences such as
the length and width of the puparia, the numbers and size of
various foveae, and the extent to which the abdominal seg¬
mentation is indicated have also been used in describing the
puparia.

With all these characters it has been extremely difficult to
separate some species, and in the keys which follow allowance
must be made for slight variation in the puparia of some species.

The mouth-parts have been examined, and these appear to
be practically identical in all species.

237

1938.]

To avoid confusion, the segments of the puparia have been
numbered one to eleven, segment one being the prothoracic
segment and segment eleven being the last abdominal segment.

The puparia which have been described in this paper have
been divided into groups according to the different types of
anterior spiracular processes. Each group has a key by which
it should be possible to determine the species. The species are
numbered and more detailed descriptions of the puparia follow
after the keys.

The descriptions of the eggs of the three species Leptocera
silvatica (Meigen), L. heteroneura (Haliday) and Sphaerocera
subsultans (L.) follow the descriptions of their respective

puparia.

Puparia. Key to Groups.

1. Anterior spiracular process spine-like . Group i.

— Anterior spiracular process not spine-like . . 2.

2. Anterior spiracular process palmate . . . Group 2.

— Anterior spiracular process not palmate . 3.

3. Anterior spiracular process consisting of three small tubercles

arising from a common base . Group 3.

— Anterior spiracular process consisting of eight small tubercles

arising from a common base . . . . . Group 4.

Puparia. Key to Species.

Group i.

Anterior spiracular process spine-like.

Including Leptocera vagans (Haliday), L. heteroneura (Hali¬
day), L,. palmata (Richards), L. talparum (Richards), L. crassi-
mana (Haliday) and L. manicata (Richards).

1. Tubercles branching from, and resting close to, main spine¬

like structure ..... . . . . . . . . . . . . palmata.

— Tubercles branching from, but not resting close to, main

spine-like structure . . 2.

2. Spine-like process with eleven or twelve tubercles branching

from it . . . . . . vagans.

— Spine-like process with ten or less tubercles branching

from it . . 3.

3. Length of spine-like processes equal to distance between

them — . . crassimana.

— Length of spine-like processes less than distance between

them . . 4.

4. Longitudinal foveae absent from the mid-point near the

lateral margins of the segments . . . . heteroneura.

— Longitudinal foveae present at the mid-point near the lateral

margins of some segments . . . 5.

238

[November,

5. Longitudinal foveae situated at the mid-point near the
lateral margins of segments two to eleven inclusive (dorsal
view) distinct ............ . . . . talparum.

— • Longitudinal foveae situated at the mid-point near the lateral
margins indistinct ........................... ....... manicata.

Group 2.

Anterior spiracular process palmate.

Including Copromyza glacialis (Meigen), C. equina (Fallen),
C. stercoraria (Meigen) , Leptocera silvatica (Meigen) and
L. bequaerti (Villeneuve).

1 . Longitudinal foveae situated at the mid-point near the lateral

margins, not continuous on segments two and three (dorsal
and ventral view) .................................... C. glacialis.

— Longitudinal foveae situated at the mid-point near the lateral

margins, continuous ............ L. silvatica, L. bequaerti.

2. The lobes forming the posterior spiracular process only partly

visible from the ventral surface ... C. stercoraria, C. equina.

Group 3.

Anterior spiracular process consisting of three small tubercles
arising from a common base.

Including Leptocera fungicola. (Haliday) and L. melania (Haliday).

1. Ventral surface with distinct transverse wrinkles on seg¬
ments four, five and six ........................... L. melania.

— Ventral surface without transverse wrinkles ... L. fungicola.

Group 4.

Anterior spiracular process consisting of eight small tubercles
radiating from a common base ...... Sphaerocera sub sultans .

Detailed descriptions of Puparia and Eggs.
Leptocera silvatica (Meigen) (Fig. 1).

The puparium is brown. In the five specimens measured the
length varied from 3-42 to 3-82 mm. and the width at segment
seven from i*o to i-n mm.

The puparium tapers evenly both posteriorly and anteriorly.
The two anterior spiracular processes each consists of a dark
brown palmate structure, similar to L. bequaerti Villen., with
seven short tubercles branching from it, which are lighter in
colour towards the apex. They are situated anteriorly on the
sides of segment one.

239

1938.]

Fig.' i.

Leptocera silvatica (Meigen). A, Dorsal view of puparium ; B, Ventral view
of puparium ; C, Mouth-parts of the larva ; D, Segment one showing anterior
spiracular processes ; E, End view showing posterior spiracular processes |

F, Egg.

Dorsal view. The segmentation of the puparium is indis¬
tinct except for the last three segments. The dorsal surface is
covered with fine transverse lines, which become larger and
more uneven on the last segment.

Segments one to ten have a longitudinal fovea situated at
the mid-point near the lateral margin of each. These vary in
size, and extend approximately for £, £, J, -J-, ^ the

length of the segments respectively. On segments two and
three the foveae join to make one long furrow.

The posterior spiracular processes are situated terminally
and consist of two small tubercles with dark flattened ends
which slope towards the centre of the last segment. The dis¬
tance between the apices of the two tubercles is slightly less
than half the width of the posterior margin of segment eleven.

Ventral view \ The ventral surface is covered with fine trans¬
verse lines which are more distinct than on the dorsal surface.
They are larger on segment eleven, especially near the anal
opening, which is situated centrally on this segment. There
are longitudinal foveae situated at the mid-point near the lateral
margins of segments two to ten inclusive. On segments two
and three the foveae join and form a long furrow. On seg¬
ments four, five, six and seven they are smaller, and on seg¬
ments four and five can only be seen with difficulty. A row of
pseudopods is visible on segments five to eleven inclusive.

240

[November,

The ten eggs measured varied in length from 0-63 to o*68 mm.
and in width from 0*2 to 0*23 mm. The egg is oval with the
dorsal surface flattened except for a slightly raised area in the
centre, which falls away towards the anterior end to form a
slight hollow. The egg tapers anteriorly to form a slight lip.
There is a row of small spine-like processes round the edge of
the dorsal surface. The entire surface of the egg is covered with
minute indentations which are smaller and more numerous on
the dorsal surface and can only be seen in certain lights.

Leptocera bequaerti (Villeneuve) (Fig. 2).

The puparium is brown. The length of the specimen drawn
is 3 mm. and the width at segment six is 0-89 mm. Anteriorly
the puparium tapers gradually to the posterior margin of seg¬
ment three, which measures 0-72 mm., and then more abruptly
to the anterior margin of segment one, which measures 0-28 mm.
Posteriorly the puparium tapers evenly to the last segment, the
base of which measures 0-56 mm. and the apex 0-33 mm. The
two anterior spiracular processes each consists of a palmate
structure with six small tubercles. The width of a process is
o-o6mm. and the length 0-07 mm. They are situated anteriorly
on the sides of segment one.

&-

3

3

Fig. 2.

Leptocera bequaerti (Villeneuve). A, Dorsal view of puparium ; B, Ventral
view of puparium ; C, Part of segment one showing anterior spiracular
processes ; D, Posterior spiracular plate.

241

1938.]

Dorsal view. The segmentation of the puparium is distinct
especially on segments one, two, three, nine, ten and eleven.
The anterior margin of segment one is wrinkled between the
anterior spiracular processes. There is also a raised and rounded
area on segments one, two and three, at the sides of which is a
furrow which becomes less deep at the junction of segments one
and two, and two and three. Segments four to ten have a
shallow fovea on the lateral margins which decreases in size to
segment seven and on this segment is difficult to see. It then
increases to segment ten, where it is situated centrally on the
lateral margins and extends to half the length of this segment.
Two lobes situated terminally on segment eleven form the pos¬
terior spiracular processes. The spiracular openings are situated
on the apex of each lobe. The distance between the lobes is
approximately one-third of the width of the apex of the segment.

The dorsal surface is covered with fine transverse lines. On
segment two and three these lines are more pronounced, and on
segment eleven they become shorter and deeper.

Ventral view. Segments two and three have two longitudinal
furrows, one each side of the mouth-parts, which are situated
centrally and can be clearly seen inside the puparium. Seg¬
ments four to ten inclusive have a longitudinal fovea situated
at the mid-point near the lateral margins of each ; these extend
J, i> h 2y I? i length of the segments respectively. A row
of pseudopods is visible on segments five to ten inclusive. The
ventral surface is covered with transverse lines which become
coarser on segments one, two, three and ten, and shorter and
thicker on the last segment, especially near the anal opening,
which is situated centrally on this segment.

Leptocera fungicola (Haliday) (Fig. 3).

The puparium is yellow-brown. The length of the specimen
drawn is 1 -8 mm. and the width at segment six is 0*59 mm.
Anteriorly the puparium tapers gradually from segment six to
segment four and then more abruptly to the anterior margin of
segment one, which measures 0-19 mm. Posteriorly it tapers
evenly from segment six to the posterior margin of the last
segment, which measures 0-17 mm.

The two anterior spiracular processes each consists of three
very short finger-like processes arising from a common base.
They are similar in shape to the anterior spiracles of L. melania
Hal., but are shorter and thicker. The width of one complete
process is 0-03 mm. and the length 0-02 mm. They are situated
anteriorly on the sides of segment one.

Dorsal view. The segmentation of the puparium is indis¬
tinct except for segments one and eleven. The surface is smooth
except for a few scattered transverse lines. The anterior margin

242

[November,

Leptocera fungicola (Haliday). A, Dorsal view of puparium ; B, Ventral
view of puparium; C, Part of segment one showing anterior spiracular
processes ; D, Posterior spiracular plate.

of segment one is raised centrally to form two small vertical
ridges between the anterior spiracular processes. The lateral
margins of segments one and two are also slightly corrugated.

Segment eleven is covered with numerous irregular wrinkles
which become closer towards the apex. The posterior third of
segment eleven is constricted centrally and forms two lobes, on
the ends of which are the posterior spiracles. These lobes pro¬
ject beyond the end of the puparium. The posterior spiracular
processes are similar to those of L. melania Hah, but L. melania
has no lobes. The distance between the spiracles of L. fungi¬
cola Hal. is half the basal width of segment eleven.

Ventral view. Segments two and three have two shallow
longitudinal furrows, one on each side of the mouth-parts, which
can be clearly seen inside the puparium. Segments four to ten
inclusive are smooth except for a few scattered transverse lines.
The pseudopods are extremely difficult to see. The anal open¬
ing is situated centrally on segment eleven, which is wrinkled
as on its dorsal surface.

'Leptocera heteroneura (Haliday) (Fig. 4).

The puparia are light yellow-brown. They vary in length
from 2-o to 2-i mm. and in width from o-y to o-8mm. ; eight
were measured. The puparium tapers evenly both anteriorly
and posteriorly. The two anterior spiracular processes each

1938.]

243

consists of a black spine-like process similar to L. manicata
(Richards), with seven or eight short pale tubercles branching
from it. The spine-like processes are shorter than those of L.
manicata and measure approximately half the distance between
the two processes. They are situated anteriorly on the sides of
segment one.

Fig. 4.

Leptocera heteroneura (Haliday). A, Dorsal view of puparium ; B, Ventral
view of puparium; C, Mouth-parts of larva ; D, Segment one showing
anterior spiracular processes ; E, End view showing posterior spiracular
processes ; F, Egg.

Dorsal view. For its size the segmentation of the puparium
is distinct. The surface is covered with fine transverse lines.
On the last segment there are two oval depressions situated
directly above the base of the two tubercles which form the
posterior spiracular processes. These tubercles are situated
laterally on the posterior margin of segment eleven. The
spiracular openings are on the flattened ends of the tubercles,
which slope slightly towards the centre of the segments.

Ventral view. A row of pseudopods is visible on segments
five to eleven inclusive. The surface is covered with fine trans¬
verse lines which become much coarser on segment eleven,
especially near the anal opening, which is at the centre of this
segment.

The eggs are oval and vary in length from 0-43, to 0-45 mm.
and in width from o-i to 016 mm. ; ten were measured. The
dorsal surface is flattened and has four longitudinal ridges.
The remainder is rounded and is covered with minute depres¬
sions, which are visible only in certain lights.

244

[November,

Leptocera crassimana (Haliday) (Fig. 5).

The puparium is brown and varies considerably in size. Two
were measured. These varied in length from 2-31 to 2-55 mm.
and in width at segment seven from 0*75 to 0*78 mm. The two
anterior spiracular processes each consists of a black spine-like
process, similar to L. manicata (Richards), with eight or nine
short pale tubercles branching from it. The length of each pro¬
cess is approximately the same as the distance between them.
They are situated anteriorly on the sides of segment one and
are often curved inwards towards the centre of the segment.

Dorsal view. The segmentation is distinct. The surface is
covered with fine transverse lines which become coarser on
segments one and eleven. There is a longitudinal fovea situated
at the mid-point near the sides of segments two to ten inclu¬
sive ; the foveae vary in size, and on segments two to five
inclusive extend for approximately f, J, -J, -J the length of the
segments respectively. From segments six to ten inclusive the
foveae become very small and do not extend for more than one-
sixth of the length of the segments. Two terminal tubercles

Leptocera crassimana (Haliday). A, Dorsal view of puparium; B, Ventral
view of puparium ; C, Segment one showing anterior spiracular processes ;
D, End view showing posterior spiracular processes.

1938.]

245

form the posterior spiracular processes. These tubercles have
flattened ends which bear the spiracular opening's, which slope
slightly towards the centre of the last segment.

The three spiracular openings are situated on the outer
margins of the spiracular plate, and on the outer margins of
the openings there is a row of extremely fine branched hair¬
like processes, which, however, can only be seen with high
magnification.

Ventral view. The ventral surface is covered with fine trans¬
verse lines which are more distinct than on the dorsal surface.
They become coarser on segments one and eleven. There is a
longitudinal fovea situated at the mid-point near the sides of
segments two to ten inclusive, the fovea extending for one-third
of the length of the segments except on segments two, three
and four, where it extends f, | the length of the segments
respectively. A row of pseudopods is visible on segments five
to eleven inclusive. The anal opening is situated centrally on
the last segment.

Leptocera palmata (Richards) (Fig. 6).

The puparium is pale brown. The length of the specimen
drawn is 2-66 mm. and the width at segment six is 098 mm.
From segment six the puparium tapers evenly both anteriorly
and posteriorly, the width of the anterior margin of segment

Fig. 6.

Leptocera palmata (Richards). A, Dorsal view of puparium ; B, Ventral
view of puparium ; C, Part of segment one showing anterior spiracular
process ; D, Posterior spiracular plate.

246

[November,

one and the posterior margin of segment eleven being- 0*29 mm.
The two anterior spiracular processes each consists of a black
spine-like process similar to L. manicata (Richards), with eleven
or twelve short pale tubercles branching from but resting close
to the main structure. The length of the process is approxi¬
mately two-thirds of the distance between them. They are
situated anteriorly on the lateral margins of segment one.

Dorsal view. The segmentation is very indistinct. The sur¬
face is covered with fine transverse lines which become coarser
on segments one and eleven. The two tubercles situated ter¬
minally on segment eleven form the posterior spiracular pro¬
cesses. The spiracular openings are situated at the ends of the
lobes, which are paler than at their base. The distance between
them is half the width of the posterior margin of segment eleven.

Ventral view. The surface is covered with fine transverse
lines which become coarser on segment eleven, especially pos¬
teriorly to the anal opening. A row of pseudopods is visible on
segments five to ten inclusive. I have been unable to ascertain
whether pseudopods are present on segment eleven. The anal
opening is situated centrally on segment eleven, in the centre
of a flat diamond-shaped area. There is a small black spot
situated each side of the anal opening on this area.

Leptocera manicata (Richards) (Fig. 7).

The puparium is pale brown. The length of the specimen
drawn is 2*i6mm. and the width at segment seven is 072 mm.
The puparium tapers evenly anteriorly from segment seven to
the centre of segment four, and then tapers more abruptly to
segment one, where it is 0^3 mm. wide. Posteriorly, the
puparium tapers evenly to the posterior margin of segment
eleven, which is 0*28 mm. wide.

The two anterior spiracular processes each consists of a
long black spine-like process with eight or nine short pale
tubercles branching from it. The length of each process is
slightly less than half the width of the anterior margin of seg¬
ment one. They are situated anteriorly on the sides of seg¬
ment one.

Dorsal view. The segmentation is distinct except between
segments four and five, five and six, six and seven, and seven
and eight.

There is a longitudinal fovea situated at the mid-point near
the sides of segments two to ten inclusive. These vary in size
and extend for approximately J, J, J, J, J, J, J the length
of the segments respectively. The dorsal surface is covered
with fine transverse lines which become coarser on segments
one and eleven.

1938.]

247

Fig. 7.

Leptocera manicata (Richards). A, Dorsal view of puparium ; ; B, Ventral
view of puparium ; C, Part of segment one showing anterior spiracular
process ; D, Posterior spiracular plate.

Two tubercles situated terminally on segment eleven form
the posterior spiracular processes. The spiracular openings are
situated on the flattened ends of the lobes. The distance between
the two lobes is half the width of the posterior margin of the
segment.

Ventral view. Segments two and three form a slightly
raised and rounded area, and have a shallow longitudinal furrow
near the sides. Segments four, seven, eight and nine have a
longitudinal fovea situated near the sides. On segment four it
is at the mid-point, on segments seven, eight and nine on the
anterior half of the segments. The foveae extend for approxi¬
mately J, if, if, J the length of the segments respectively. There
are two small black spots, situated one on each side of the
mouth-parts on segment one. The whole surface of the puparium
is covered with fine transverse lines which become coarser on
segments one, ten and eleven. A row of pseudopods is visible
on segments five to ten inclusive.

Leptocera t alp arum (Richards) (Fig. 8).

The puparium is pale brown. The length of the specimen
drawn is 2-35 mm. and the width at segment seven is 077 mm
From segment seven the puparium tapers evenly both anteriorly

248

[November,

and posteriorly. The width of the anterior margin of segment
one and the posterior margin of segment eleven is 0*24 mm.
The two anterior spiracular processes each consists of a long
black spine-like process with eight or nine short pale tubercles
branching from it, similar to L. manicata (Richards). The
length of each process is slightly less than the distance between
them. They are situated anteriorly on the sides of segment one.

B

A

Fig. 8.

Leptocera talparum (Richards). A, Dorsal view of puparium ; B, Ventral
view of puparium ; C, Part of segment one showing anterior spiracular
process ; D, Posterior spiracular plate.

Dorsal view. The segmentation is indistinct except between
segments two and three, and ten and eleven. There is a longi¬
tudinal fovea situated at the mid-point near the sides of seg¬
ments two to eleven inclusive. The foveae vary in size and
extend approximately f, J, f, f, f, f, $ the length of

the segments respectively. The dorsal surface is covered with
fine transverse lines which become coarser on segments one and
eleven. Two tubercles situated terminally on segment eleven
form the posterior spiracular processes. The spiracular open¬
ings are situated on the ends of the lobes. The distance between
the lobes is half the width of the posterior margin of segment
eleven.

Ventral view. Longitudinal foveae are situated at the mid¬
point near the sides of segments two to ten inclusive. These
vary in size and extend approximately f, f, f, f, J, J, J, i

1938.]

249

the length of the segments respectively. The entire surface of
the puparium is covered with fine transverse lines, which be¬
come coarser on segment eleven. The pseudopods are extremely
difficult to see. On each side of the anal opening, which is
situated centrally, there is a minute black spot.

Leptocera rnelania (Haliday) (Fig. 9).

The puparium is yellow-brown. The length of the specimen
drawn is i-6mm. and the width 0-5 mm. The anterior spiracular
processes each consists of three short thick finger-like processes
arising from a common base. They are black-brown in colour.
The measurements of one complete process are 0-026 mm. long
and 0-02 mm. wide. They are situated anteriorly on the sides
of segment one.

Dorsal view. There is a longitudinal fovea situated at the
mid-point near the sides of segments two to ten inclusive. They
vary in size, on segment two being very shallow and extending
not more than one-third of the length of the segment, while on
segments three and four they are deeper and extend to half the
length of the segments, and on segments five and six they are
still deeper and arise from the anterior edge of the segment,
curving towards the centre. Their length is approximately one-
quarter the width of the segments. On segments seven to ten
inclusive the foveae are reduced considerably. They are more
parallel to the long axis of the puparium and extend J, J, J
the length of the segments respectively. On the posterior third
of the last segment there is a W-shaped depression. The pos-

Fig. 9.

Leptocera melania (Haliday). A, Dorsal view of puparium ; B, Ventral view
of puparium ; C, Segment one showing anterior spiracular processes ; D,
Posterior spiracular plate.

250

[November,

terior spiracular processes are situated terminally and consist
of two small brown papilliform processes, the distance between
them being- one-third of the anterior width of the last segment.

Ventral view. Segments two and three form a slightly
raised area and have two shallow longitudinal furrows, one
each side of the centrally placed mouth-parts, which can be
clearly seen inside the puparium. Segments four, five and six
have a central transverse furrow extending for about three-
quarters the width of each segment and deepening at their
extremities. On the lateral parts of segments seven to ten
inclusive there is a longitudinal depression one-third of the
length of the segments. The depressions on segment ten are
very shallow and can only be seen with difficulty. The anal
opening is situated centrally on the last segment.

Leptocera vagans (Haliday) (Fig. io).

The length of the puparium varies from 2-4 mm. to 2-62 mm.
and the width from 0-84 mm. to 0*98 mm. ; four were measured.
The two anterior spiracular processes each consists of a black
spine-like process similar to L. manicata (Richards), with eleven
or twelve short pale tubercles branching from it. The spine-like
processes are shorter than L. manicata, and measure approxi¬
mately half the distance between the two spiracular processes.

Fig. io.

Leptocera vagans (Haliday). A, Dorsal view of puparium ; B, Ventral view
of puparium ; C, Mouth-parts of larva ; D, Segment one showing anterior
spiracular processes; E, End view showing posterior processes.

1938.]

251

Dorsal view. The puparium tapers evenly both anteriorly
and posteriorly. The anterior margin of segment one is con¬
stricted centrally. The surface of the puparium is covered with
fine transverse lines. The segmentation is distinct, especially
between segments ten and eleven. Two short tubercles situated
laterally on the posterior margin of segment eleven form the
posterior spiracular processes. The spiracular openings are
situated on the flattened ends, the distance between the ends
being the length of the posterior margin.

Ventral i new-. The surface is covered with fine transverse
lines. A row of pseudopods is visible on segments five to eleven
inclusive. The anal opening is situated centrally on segment
eleven.

Copromyza glacialis (Meigen) (Fig. n).

The puparium is brown. The length of the specimen drawn
is 3-44 mm. and the width at segment six is 1-03 mm. The
puparium tapers evenly both anteriorly and posteriorly to the
posterior margin of segment ten. Segment eleven is only three-
quarters the width of segment ten. The two anterior spiracular
processes each consists of a palmate structure similar to C.
stercoraria (Meigen) with eight short tubercles. They are
situated on the anterior half of the sides of segment one.

Fig. 11.

Copromyza glacialis (Meigen). A, Dorsal view of puparium ; B, Ventral
view of puparium ; C, Mouth-parts of larva ; D, Segment one showing
anterior spiracular processes ; E, End view showing posterior spiracular

processes.

252

[November,

Dorsal view. The segmentation of the puparium is distinct.
The surface of the puparium is covered with fine transverse
lines which become coarser on segments one, two, three and
eleven. Segments one, two, three and five to ten inclusive have
a longitudinal fovea situated at the mid-point near the sides of
each. These vary in size and extend approximately J, J and
i> 3> ¥ ^e length of the segments respectively. The

button-like processes situated terminally on segment eleven
form the posterior spiracular processes. In the specimen drawn
these could not be seen from the ventral surface. The distance
between the processes is slightly less than the width of one of
the buttons.

Ventral view. The ventral surface is covered with fine trans¬
verse lines, becoming coarser on segments one, two, three and
eleven, especially near the anal opening, which is situated cen¬
trally on this segment. A row of pseudopods is visible on seg¬
ments five to ten inclusive. Pseudopod-like structures are also
present on segment four, but as yet I have been unable to ascer¬
tain whether they are true pseudopods. Segments two, three
and five to ten inclusive have a longitudinal fovea situated at
the mid-point near the sides of each. These vary in length and
extend for the length of segments two and three, and for J,
£, J and J the length of segments five to ten respectively.

Copromyza equina (Fallen).

I have compared the puparia of C. equina and C. stercoraria
and have been unable to separate them with any certainty. The
only difference I have found between the two species is in the
size. Four of the six specimens of Copromyza equina are larger
than the one specimen of C. stercoraria, which was measured.
The remaining two are slightly smaller. The measurements
are: — Length, 3-5 — 3*8 mm.; width at segment seven, i*i6 —
1 -31 mm.

Copromyza stercoraria (Meigen) (Fig. 12).

The puparium is brown. The length of the specimen drawn
is 3-64 mm. and the width at segment seven is 1-24 mm. The
puparium tapers evenly both posteriorly and anteriorly. The
anterior margin of segment one is 0*42 mm. and the posterior
margin of segment eleven is 0-48 mm. wide. The two anterior
spiracular processes each consists of a palmate structure similar
to Leptocera bequaerti with eight short tubercles. They are
situated anteriorly on the sides of segment one.

Dorsal view. The segmentation of the puparium is distinct,
especially between segments two and three, three and four,
eight and nine, nine and ten, and ten and eleven. The sides of
segment one and anteriorly of segment two are wrinkled. There

1938. J

253

Fig. 12.

Copromyza stercoraria (Meigen). A, Dorsal view of puparium ; B, Ventral
view of puparium ; C, Anterior spiracular process ; D, Posterior spiracular

process.

is a raised area situated centrally on segments two and three.
Longitudinal foveae situated at the mid-point near the sides are
present on segments two to ten inclusive. They extend for § , §,
3> 4> 3 the length of the segments respectively. The
dorsal surface is covered with fine transverse lines. On seg¬
ments one, two and three the lines become coarser, and on the
last segment they develop into short irregular wrinkles. There
is a row of minute spots in the sutures between the segments.
They continue across the dorsal surface and end just short of
the row of pseudopods on the ventral surface. Two lobes
situated terminally on the dorsal surface of segment eleven
form the posterior spiracular processes. These lobes are only
partly visible from the ventral surface of the puparium. The
spiracular openings are situated on the ends of the lobes. The
distance between the lobes is one-third of the width of the pos¬
terior margin of the last segment.

Ventral view. There is a slightly raised and rounded area
situated centrally on segment two. Segments two to ten inclu¬
sive have a longitudinal fovea situated at the mid-point near

254

[November,

the lateral margins of each. These vary in size and extend
approximately f, -J the length of the segments

respectively. There is also a shallow transverse fovea situated
centrally on the anterior margin of segment eleven. The pseudo-
pods can be seen clearly, and arise from a point near the anterior
ends of the longitudinal foveae of segments five to ten inclusive.
The ventral surface is covered with fine transverse lines which
become coarser on segment one and the lateral parts of seg¬
ment two. Posteriorly segment eleven is coarsely wrinkled,
especially near the anal opening, which is situated centrally on
this segment.

Sphaerocera subsultans (Linnaeus) (Fig. 13).

The puparium is black-brown. The length varies from 3*36
to 3-85 mm. and the width at segment seven from 1-16 to
1 -28 mm.; eight were measured. The puparium tapers evenly
both anteriorly and posteriorly. The anterior spiracular pro¬
cesses each consists of eight short thin tubercles radiating from
a common base. They are situated anteriorly on the sides of
segment one.

Dorsal view. The segmentation of the puparium is distinct.
The surface is covered with transverse lines which become

Fig. 13.

Sphaerocera subsultans (Linnaeus). A, Dorsal view of puparium ; B, Ventral
view of puparium ; C, Mouth-parts of larva ; D, Anterior end of puparium
showing anterior spiracular processes; E, Dorsal view of segment eleven
showing posterior spiracular plates; F, Egg.

1938.]

255

coarser and more numerous on the anterior two-thirds of the
last segment. Segments three, four and five have a shallow
fovea situated at the mid-point near the sides of each ; the
foveae extend approximately -J*, J, J the length of the segments
respectively. The foveae become elongate and more pronounced
on segments six to ten inclusive, and extend J, J, §, f, f the
length of the segments respectively. There is a small deep
cavity situated in the centre of segment eleven.

The posterior spiracular processes are situated on the pos¬
terior quarter of the last segment, and therefore cannot be seen
from the ventral surface. They consist each of a small black
oval opening in the surface of the puparium. The space between
them is half the width of the posterior margin of segment eleven.

Ventral view. The ventral surface is covered with trans¬
verse lines which become coarser on segments nine, ten and
eleven, especially near the anal opening, which is situated cen¬
trally on the last segment. The anal opening is situated in the
centre of a pale, depressed, square area. Segments two, three
and four have a shallow fovea situated at the mid-point near
the sides of each ; the foveae extend approximately J, \ the
length of the segments respectively. These foveae become
elongate and more pronounced on segments five to ten inclu¬
sive and extend J, f, f, f, j, J the length of the segments
respectively.

Description of egg. The length of the egg varies from
o -72 to 0-78 mm. and the width from 0-24 to 0-27 mm. ; seven
were measured. The egg is white, and the surface is covered
with thirteen or sometimes fourteen ridges, which extend for
the whole length and end anteriorly at a flat disc through which
the larva emerges.

Locality and Habitat Records.

As the eggs, larvae, pupae and adults were collected, the
following data were noted : —

It will be seen that the term ‘ mouse-run ’ is used ; by this
is meant the tunnels made by mice at the base of long grass.

A few of the species mentioned in the paper have not been
recorded for Buckingham by Richards (1930). These have been
marked with an asterisk (*).

*1. Leptocera fontinalis (Fallen).

1 cf, Bucks, Slough, decaying lawn-mowings, 16 Oct., ’33.

*2. Leptocera curvinervis (Stenhammar).

2 c? cf , 4 $ $ ) Bucks, Slough, garden refuse, 16 Oct.,’ 33.

3. Leptocera nivalis (Haliday).

1 d*, 1 $ , Bucks, Slough, mouse-run, 17 Oct., ’33.

256

[November,

4. Leptocera silvatica (Meigen).

2 d* dS 9 9 .9 ) 9 pupae, Bucks, Slough ; Berks, Windsor
Great Park, in cow-dung, decaying lawn-mowings,
7-8 Feb., 31 Oct., ’34.

Three females collected on 7 Feb., ’34, laid one hundred
eggs the net day. One adult emerged on 23 March.
When the eggs are laid they are buried in the decay¬
ing grass, except for the dorsal surface.

*5. Leptocera bequaerti (Villeneuve).

1 d* , 3 9 9> collected as larvae, Bucks, Slough, mouse-
run, 14 Feb., ’35.

The four larvae pupated three days after being collected,
and the adults emerged from sixteen to eighteen days
later.

*6. Leptocera fungicola (Haliday).

1 cf) collected as pupa, Bucks, Slough, mouse-run, 23

Jan-, ’35-

The adult fly emerged eight days after the pupa was
collected.

*7. Leptocera minutissima (Zetterstedt).

4 cfcf, Bucks, Slough, rabbit-hutch, 13 June, ’35.

*8. Leptocera heteroneura (Haliday).

1 cf, 3 9 9) Bucks, Slough, decaying lawn-mowings,
16 Oct., 1 Nov., ’33.

One female collected on decayed lawn-mowings on 16
Oct*) ’33, was placed in a tube with half an inch of
sterilised decayed grass cuttings at the bottom. From
this tube sixty-two flies emerged from thirty-one to
thirty-eight days later. Four pairs of these flies were
placed in four other tubes with decayed lawn-mowings.

In the first tube eggs were laid five days after copula¬
tion had taken place. Four eggs were laid and one
larva was seen two days later. In the second tube
eggs were laid six days after copulation ; seventy eggs
were laid in ten days. The parent fly died four days
.later. The maximum number of eggs, twelve, was
laid on the fifth day. In the third tube egg-laying
commenced nineteen days after copulation. Sixty-five
eggs were laid in sixteen days. The parent died four
days later. The maximum number of eggs, fifteen,
was laid on the eleventh day. In the fourth tube eggs
were laid twelve days after copulation. Sixty-one eggs
were laid in thirty-two days, the maximum number,
sixteen, being laid on the eighth day. The eggs were
scattered on the surface of the grass. Some were de¬
posited on the glass of the tubes.

9. Leptocera longisetosa (Dahl).

1 9 ) Berks, Windsor Great Park, deer-dung, 14 Oct., ’33.

257

1938.]

10. Leptocera crassimana (Haliday).

2 cfdb 2 9 $, 3' pupae, Bucks, Slough, decaying grass,
31 Oct., 12 Dec., ’33, 9 Feb., ’34.

The adults emerged from the pupae three days after
being collected.

11. Leptocera palmata (Richards).

2 d* db 1 9 , Bucks, Slough, mouse-run, 1 Jan., ’35.

3 C? cf > 4 9 $ > Surrey, Hindhead, dead rabbit, 8 Mar., ’36.

12. Leptocera manicata (Richards).

10 cfdb 6 9 9 , collected as larvae and pupae, Bucks,
Slough, mouse-run, 8 Mar., ’34, 28 and 29 Jan., 12
Feb., 8 Apr., ’35.

The adults emerged from six to fourteen days after
pupation.

13. Leptocera talparum (Richards).

13 cfdb 11 9 9 , collected as larvae and pupae, Bucks,
Slough, mouse-run, 18 Mar., ’34, 12 Feb., 8 Apr., ’35.

The adults emerged from five to eighteen days after
pupating.

*14. Leptocera rwfilabris (Stenhammar).

1 9 5 Bucks, Slough, decaying grass, 31 Oct., ’33.

*15. Leptocera luteilabris (Rondani).

3 cf db 3 9 9, Bucks, Slough, rabbit-hutch, 13 June, ’36.

16. Leptocera spinipennis (Haliday).

1 db 1 .9 > Bucks, Slough, mouse-run, decaying grass,
17 and 31 Oct., ’33.

*17. Leptocera aterrima (Haliday).

1 9 , Bucks, Slough, lawn-mowings, 31 Oct., ’33.

*18. Leptocera melania (Haliday).

1 9 collected as pupa, Bucks, Slough, mouse-run,
1 Jan., ’35.

The fly emerged sixteen days after the pupa was col¬
lected.

*19. Leptocera vagans (Haliday).

1 db 4 9 9 , Bucks, Slough, mouse-run, decaying grass,
16 and 17 Oct., ’3.3.

One of these females was placed in a tube with some
sterilised decayed grass. On 29 Nov., ’33, twelve
adults (10 9 9 , 2 d* cf) emerged and on 30 Nov., ’33,
ten more (9 9 .9 > 1 cT).

20. Leptocera lugubris (Haliday).

2 9 9 , Berks, Windsor Great Park, cow-dung, 24
Oct., ’33.

*21. Leptocera ferruginata (Stenhammar).

1 db Bucks, Slough, decaying grass, 3 Oct., ’33.

22. Copromyza nitida (Rondani).

2 9 9> Hants, Alton, sheep-dung, 14 Nov., ’3,3.

258

[November,

23. Copromyza roserii (Rondani).

4 0*0*, i 9> Surrey, Hindhead, dead rook, dead rabbit,
18 Mar., ’36.

*24. Copromyza glacialis (Meigen).

1 cf, i collected as larvae, Bucks, Slough, Farnham
Royal, nest of field mouse, mouse-run, 24 Apr., ’31,

12 Feb., ’35.

One fly emerged seventeen days after pupating.

25. Copromyza equina (Fallen).

1 cf, 2 9 9 , Hants, Alton; Bucks, Slough; Berks,
Windsor Great Park. In decaying lawn-mowings,
cow-dung, sheep-dung, 13 and 24 Oct., 14 Nov., ’33.

26. Copromyza similis (Collin).

4 9 9 , Berks, Windsor Great Park, cow-dung, deer-
dung, 24 Oct., ’33.

27. Copromyza stercoraria (Meigen).

25 , 27 9 9 , collected as larvae, Bucks, Slough,

mouse-run, 16 Mar., ’34, 28 and 29 Jan., 11 and 14
Feb., 8 Apr., ’35.

The flies emerged seven to ten days after pupating.

28. Sphaerocera sub sultans (L.).

4 cfcf, 2 9 9) Bucks, Slough, decaying grass, 16 Oct.,

13 and 21 Nov., ’33.

A pair of flies was placed in a tube with some sterilised
decayed grass. Thirty-six eggs were laid two days
later, and one adult emerged thirty-three days after
the eggs were laid.

The eggs were scattered on the surface of the grass and
not buried as in Leptocera silvatica.

*29. Sphaerocera nitida (D'uda).

3 9 9 , Bucks, Slough ; Hants, Alton ; decaying grass,
sheep-dung, 2 and 11 Nov., ’35.

30. Sphaerocera pusilla (Fallen).

1 cf, 3 9 9 , Bucks, Slough, decaying grass, 10 Oct.,

14 Nov., ’33.

Parasites.

Two parasites were reared from the Sphaerocerid pupae.
Three specimens of Phaenopria cameroni Kieff. emerged from
three puparia of one of the smaller species of Leptocera (prob¬
ably L. vagans) ; one specimen of Opius sp. emerged from the
pupa of Copromyza sp.

Many of the flies when collected were found to have several
large red mites attached to various parts of their abdomens.
These mites have not yet been identified.

Reference.

Richards, O. W. 1930. Proc. Zool. Soc., 18: 261-345.

HOLYWELL PRESS, ALFRED STREET, OXFORD

LEPIDOPTERA (contd.).

Lithocolletis Concomitella Bankes and
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DIPTERA.

CONTENTS.

Goddard, W. H., D.I.C. : The Description of the Puparia of
fourteen British Species of Sphaeroceridae (Borboridae,
Diptera).

Communications for the Transactions should be sent to the
Editor at: —

68, Whitworth Crescent, Bitterne Park, Southampton.