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The Structure of H
æmatopota Pluvialis
*
(Meigen)
BY
CAPTAIN F. W. CRAGG, M.D., I.M.S.
INTRODUCTION.
T
HE rapid advance which has taken place in recent years in our knowledge of
the hœmatozoa has given a special interest to the study of the blood-sucking
diptera, which, in so many cases, have been found to act as transmitting agents.
A knowledge of the structure of the fly is essential in order to understand the
various phases in the development of the parasite, and, as regards the pathogenic
forms, the life history and habits of the transmitter have proved to be the
dominant factor in any attempts to combat the disease in a rational manner.
Hœmatopota pluvialis is a fairly typical member of the Tabanidæ, a large and
widely distributed family of flies, in which the blood-sucking habit is almost uni
- versal in the female sex. Up to the present, no member of the group has been
proved to act as the host, in the strict biological sense, of a blood parasite, though
there is little doubt that certain species can act as the transmitting agents of
Surra and the allied diseases. Apart from this, the study of the group from the
purely zoological point of view is of considerable interest. Hœmatopota shows,
perhaps, the minimum amount of differentiation from the mandibulate type which
is consistent with a blood-sucking habit, and thus affords an excellent opportunity
for the study of the process of the adaptation of structure to function. As will
appear later, this fly has many points of resemblance to both Anopheles (7) and
to Musca, (6) and is, generally speaking, simpler in structure than either of these.
Its relatively large size enables one to demonstrate such complicated processes as
the mechanism of feeding or the articulation of the wing much more satisfactorily
than could be done with smaller and less primitive insects.
A word is necessary with regard to the name Hæmatopota. Until recently
this
nam
e was accepted without dispute, but recently an old paper by Meigen, of
which only two copies are known to exist, has been unearthed, and from this it
appears that, previous to the adoption of the name Hœmatopota, this genus was
named by Meigen Chrysozona (8). A strict adherence to the established rules of
zoological nomenclature would therefore necessitate the substitution of the name
* The substance of this paper was presented as a thesis for the degree of M.D. Edin.
B
2
Chrysozona for the well known one of Hœmatopota. The matter is still in dispute,
and it remains to be se
en whether the change of name will be universally accepted.
The old name is used in this paper pending a settlement, since Mr. Austen is kind
enough to inform me that he has not adopted the name Chrysozona for official use
in the British Museum.
FIELD NOTES.
Under various local names, such as " cleg," " gadfly," " horse fly," the female
Hœmatopota pluvialis is well known in most parts of the British Isles, and is
much the
commonest member of the Tabanidæ found in this country. It is on
the wing from early June to the end of August or even later, and frequents damp
pastures and country lanes, being seldom found far from water, or near houses.
Its appearance depends to a remarkable extent on favourable conditions, viz.,
abundance of sunshine and an absence of wind, and even in suitable weather it
varies greatly in numbers in different years, from no ascertained cause.* Its
flight is at times swift, but usually, when about to feed, it is rather deliberate and
hovering, and is accompanied by a faint hum. So far as is definitely known, the
fly is exclusively a blood-sucker, and attacks indifferently horses, cattle, or man,
choosing those which are stationary or moving quietly about, and not selecting
any particular part of the victim, the piercing organs being able to penetrate even
the thickest parts of the skin.
The method of feeding, as observed on one's own hand, is as follows. After
a short preliminary investigation, in which the labium appears to be used as a
tactile organ, the piercing styles are inserted by a series of short sharp forward
thrusts of the thorax, the labium being retracted and the labellæ everted behind
the rest of the proboscis, and the maxilliary palps extended in front. As the
wound is deepened the fly elevates itself on its hind legs, till the abdomen is tilted
up at an angle of 45° or so, the forelegs being meanwhile extended in front of the
proboscis, with their tibiæ held parallel to the surface. While in this position the
abdomen gradually distends, and slight peristaltic movements can be seen in it.
In a short time a clear fluid begins to exude at the anus, and this goes on as long as
the insect feeds, the fluid becoming tinged with blood towards the end of the
meal, and even, in some cases, appearing to consist of unaltered blood. Several
times during the process the mouth parts are withdrawn a little and again thrust
in; finally they are withdrawn by a few sharp jerks, and the fly, after resting a
moment to clean its proboscis with its forelegs, flies away. A small drop of
blood usually oozes from the puncture, and a faint ring of hyperæmia, with
usually some slight irritation mark the site for a day or so. The pain caused by
* The larvæ of this fly live in damp mud along the banks of streams and ponds. A flood occurring at the
breeding season would therefore materially reduce the numbers of the imagines in the following year.
3
the bite is usually trifling, less than that of a mosquito bite. Some individuals,
however, appear to be remarkably sensitive, and suffer for days, even a week or
longer, from local irritation and inflammation.
The fly takes about three minutes to obtain a full meal, and while feeding it
is not readily disturbed; and the process can be watched on one's hand
through a pocket lens.
The habit of defæcating while feeding, usual in blood-sucking insects, has an
important bearing on the question of the normal parasites of the fly, for it is obvious
that any encysted phases of such parasites which may be free in the hind gut
at the time of feeding will be deposited on the skin of the host, and will be in a
position to be taken up by another fly. In the case of the Tabanidœ it is easy
to see how such parasites might be caught in the everted labellæ, and so find their
way into the pharynx as the insect feeds. This species is not gregarious, except
in the sense that large numbers may be found feeding in the same host, and they
do not appear to collect in particular localities in the evening, as described by
Hine (9) for certain American Tabanidæ. In view of their habit of defæcating on
the skin of the host, it does not appear to be necessary to assume that parasites
are transmitted from one fly to another, as described by Patton (10), by the un
- infected fly inserting its proboscis into the fæces of an infected fly, in their
evening resting-places.
The male of this species is extremely seldom met with,* even in localities in
which the female is common enough to be a nuisance. During two months in a
district where the females could be seen almost every day, only two males were
caught, and both of these were obtained early in the season. The male is a flower
feeder, and does not suck blood. A suggestion as to the cause of its remarkable
scarcity will be made when considering the reproductive organs of the female.
METHODS.
The parts were studied by the dissection of fresh specimens, and by the pre
- paration of serial sections. Dissections of the various organs, after fixation in
sublimate alcohol, were stained in various ways, chiefly with borax carmine and
Delafield's Hæmatoxylim, and mounted as permanent preparations. The chitin
- ous parts were dissected after treatment with caustic soda solution for varying
periods, with or without decolorization with chlorine gas. The proboscis, head,
thorax, and abdomen were separately embedded and cut in serial sections by the
combined paraffin and celloidin method, and the sections, though showing a good
deal of shrinkage as a result of the long time required for complete penetration of
the fluids, were very useful in checking the result of dissection.
* Vide Bruce, Report of Sleeping Sickness Commission. Proc. Roy. Series. B, Vol. 33, No. B 565.
B2
4
For the study of the mouth parts and of the complicated structures at the
base of the skull, it was found convenient to embed the whole head, after prolonged
treatment with a weak solution of caustic soda, in paraffin, and then to cut away
by hand the portions not required. The paraffin was then dissolved off in clove
oil, and the preparation mounted, with or without further dissection, in a hollow
slide. Similar preparations were made of the sides and lower part of the thorax,
and were found to be much more satisfactory than simple dissections.
The separate organs were fixed and preserved, after dissection, in Bles' fluid,*
and subsequently embedded and cut in sections. This is a most convenient
method of fixation when one is away from a laboratory, and the tissues, even after
preservation for several months in the fluid, stain quite well.
In the description of the parts, the terms superior, anterior, etc., are used in
their strict anatomical sense, with reference to the fly in a resting position on a
horizontal surface. It is important to note this in the description of the head,
since the mouth parts are almost perpendicular to the long axis of the body. The
term proboscis is used to include all the protruding mouth parts.
EXTERNAL ANATOMY.
I.—The Head.
( a) External Appearance. (Plate III, Fig. 10.)
The head has the shape of a somewhat flattened dome ; it is rather more than
twice as broad as it is long, and is a little broader than the thorax. The antennæ
project forwards from the apex of the dome, and the proboscis hangs downwards
from the lower surface. The base of the dome, forming the posterior surface of
the head, is concave to correspond with the convexity of the prothorax. The
outer thirds of the head, as seen from the front, are occupied by the large com
-pound eyes, the space between them being filled in by a square-shaped piece of the
epicranium. This piece is light fawn coloured, is covered with fine hairs, and has
on it three pigment spots, two large and anterior, the third, much smaller, mesial
and behind the other two. There are no ocelli. The posterior border of this
epicranial plate, constituting the vertex of the head, is hollowed out into a shallow
groove, which is continued on to the posterior surface of the head. Bounding this
groove there are two ill-defined sutures, which pass from the posterior and internal
angles of the eyes to the posterior surface of the head, converging towards the
occipital foramen.
* Formalin
.
.
.
7 parts.
Alcohol, 70 per cent
.
.
.
90 „
Glacial Acetic Acid
. 3 „
5
In front of the above piece, at the apex of the dome, there is a shining black
transverse band, elevated somewhat from the surface, which bears the antennæ.
This band occupies almost the entire space between the eyes, and is separated
from the epicranial plate behind and the clypeus below by definite sutures.
The Clypeus is an oblong plate, rather narrower above than below, which
stretches from the antennal plate to the labrum. It is light fawn in colour, and is
speckled with fine pigment spots, and covered with fine hairs. There are two
pigment patches, resembling those on the superior epicranial plate, in its upper
half. The anterior openings of the intracranial tunnels, to be subsequently des
- cribed, lie in the lower third of the sutures bounding the clypeus in each side.
The Genœ, as seen from the front, fill in the triangular areas between the eyes and
the clypeus; there is no suture intervening between them and the posterior
surface of the head. They resemble the clypeus in colouring, and are dotted over,
especially in the upper parts, with coarse pigment spots, and have a dense coating
of long fine hairs, which project inwards to merge with those on the maxillary
palps.
The posterior surface of the head is vertical, and slightly concave. The
occipital foramen is situated near its lower border; in front of this opening, there
is on each side a small stirrup-shaped aperture, the posterior end of the intracra
- nial tunnel.
The Eyes are conspicuous for their large size and beautiful colouring. The
ground colour is a bright iridescent greenish gold, which, unfortunately, fades
rapidly on the death of the insect. Running transversely across the eye there are
five irregular bands of reddish brown ; these markings are retained, though some
- what faintly, in specimens preserved in alcohol. The facets of the eye are hex
- agonal with rounded angles and are of approximately equal size and shape through
- out. There are a few minute hairs scattered over the surface.
The Antennœ (Plate III Fig. 12) are situated on the antennal plate already
noted. They consist of three joints, the distal end being about as long as the first
and second together. The first joint is cylindrical, the second short and round.
The third joint is ringed, having upon it three constrictions, which divide it into
four segments. The first of these segments resembles the basal joint, but is more
slender ; the second and third segments are equal, and short and round ; the
terminal segment, which is rounded at its apex, is a little longer than the two
preceding ones together. The first joint is rather sparsely covered with large, black,
forwardly directed bristles ; the second joint has a set of similar bristles arranged
in a whorl, and a few similar but smaller hairs are arranged about the constrictions
on the distal j
oints, and in a loose whorl round the middle of its first segment.
Wel
6
( b ) The Mouth Parts.
These consist of two paired organs, the mandibles and maxillœ, three single
mesial styles, the labrum, epipharynx, and hypopharynx, and a large fleshy labium,
in the groove on the anterior surface of which the piercing styles lie when at rest.
The Mandibles (Plate II, Fig. 9) are the most powerful of the piercing organs.
The blade has the shape of a short, broad, pointed and slightly recurved sabre.
The internal edge is armed in its distal half with a regular row of extremely
minute serrations ; the proximal half of the edge is extremely fine and attenuated.
The external edge is sharp only at the distal end. Proximally it broadens out, so
that the blade becomes triangular in cross section, and two stout strands of chitin
are developed along its margin, one of which crosses the surface of the blade,
while the other runs directly upwards to the base of the mandible. These two
strands are produced into two cornua, and the horseshoe-shaped arch between
them is further strengthened by thick fibres. The external horn articulates with
a projecting angle of the epicranium, the joint corresponding to the ginglymus of
mandibulate insects. The internal horn is free, and lies on the membrane which
completes the base of the skull in this region.
The Maxillœ (Plate II, Figs. 1 and 2) are remarkable in that the several parts
of a primitive first maxilla, viz., cardo, stipes, galea, lacinia and palpus, can be
readily recognised.
The Galea, or blade, is a stout slip of chitin, the same length as the mandi
- ble. It is quadrilateral in section at its tip, but becomes flattened proximally,
and at the base merges with the stipes, the yellow chitin characteristic of the
piercing parts giving place to that of the ordinary skeletal type. The tip is armed
with a covering of minute flat teeth or rasps, arranged in a somewhat imbricate
manner, with their points projecting upwards. These teeth are continued along
the edge of the blade as it becomes flattened, for about half its length on the inner
side, not quite so far on the outer side. The blade is strengthened by the develop
- ment of two strands of chitin, one of which, the larger, runs down the middle, and
merges with the stipes, while the other, lying nearer the internal edge, is produced
into the lacinia.
The Maxillary Palp, which rises from the outer side of the stipes some distance
above the base of the galea, is two-jointed and antenniform. The first joint is
short and cylindrical, the second conical and pointed, and much larger than the
first. Both joints have a dense covering of long fine hairs, many of which are
pigmented. The two palpi turn inwards and forwards from their origin, and so
come to lie in contact with one another, in front of the basal piece of the labium,
in such a way as to close in the groove in which the piercing organs lie when at
rest.