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B 2
S tudies on the Mouth Parts and Sucking Apparatus
of the Blood-Sucking Diptera.
N o. 4.
T he Comparative Anatomy of the Proboscis in the
Blood-Sucking Muscidæ.
I N two previous studies I have described the structure and mechanism of the
proboscis in Philœmatomyia insignis, Ansten, and Lyperosia minuta, Bezzi,
and in doing so adhered as far as possible to the terms already in use for the
various parts of the Muscid proboscis, thereby committing myself to the view
that a close homology exists between them. So far as the gross anatomical
features, such as the division of the proboscis into the three parts found in
Musca, and the general shape and relations of the labrum-epipharynx and
hypopharynx, are concerned, the homology is evident at a glance, but those
structures which are immediately concerned in the act of biting, in accord-
ance with the descriptions of the mechanism given in the papers mentioned
above, are modified in remarkable manner, and to so great an extent that the
relationship between them and the parts designated by the same name in Musca
can only be properly understood when the intermediate forms are examined.
In the present paper I propose to discuss the changes which have been produced
in the Muscid type of proboscis in correlation with the adoption of a blood-
sucking habit, and to show that the blood-sucking flies of this group can be
arranged in a series, commencing with those flies which are blood-suckers by
habit but have no biting mouth parts, and passing on through those flies which,
while still retaining the Musca facies, are provided with more or less efficient
biting organs, to those forms, such as Stomoxys and Glossina, which are typi-
cally blood-suckers, and have entirely lost the characteristic structure on the
labella by means of which the non-biting flies absorb nourishment. Arranged
in this manner those flies which I have been able to study form a most inter-
esting series, in which the gradual progress of a change in structure along a
definite path, correlated to a change in the habit of life, can be traced step by
step from its origin to a stage in which the structural modification is extreme.
The pointed and rigid proboscis of Glossina, typically a piercing organ, bears
little or no superficial resemblance to the blunt and retractile organ of Musca,
and yet there are few structures in the one which cannot be traced in the
other, and the mechanism by which food is obtained is closely related in
the two cases.
2
The flies which I have examined are as follows:—Three species of the
genus Philœmatomyia, viz. lineata, Brunetti, insignis, Austen, and gurnei,
Patton and Cragg; Hœmatobia* irrituns, Linn. Stomoxys calcitrans, Linn.,
Lyperosia minuta, Bezzi, and exigua Meij; Glossina palpalis, R. D., and. sub-
morsitans, Newstead. The order in which they are placed in the above list
represents approximately the degree of specialisation which is found in their
mouth parts, and this corresponds very closely with the positions assigned to
the various genera by systematic dipterologists. The genus Philœmatomyia
closely resembles the genus Musca, being distinguished from it mainly by the
presence of a ring of biting teeth, the degree of development of which differs
considerably in the three species so far known. Hœ matobia is remarkable in
that, although the general shape and structure of the proboscis shows little
difference from that of Stomoxys, traces of a pseudotracheal membrane still
persist. The genus Glossina is placed by Austen in association with Stomoxys
and Lyperosia; the condition of the mouth parts shows that it is more
highly specialised than either, and the method of reproduction supports this
view. Stomoxys and Lyperosia closely resemble one another, and the struc-
ture of the proboscis indicates that these flies have entirely abandoned food
other than blood.
Before going further it will be well to discuss the general aspects of the
question and to substantiate the hypothesis indicated in the opening paragraph,
that the blood-sucking Muscids are derived from non-biting forms. Special
attention is due to this point, for certain workers, notably Minchin and Prowa-
sek, have assumed the contrary to be the case, and the former author has used
this assumption as a support for the theory that the Hæmoflagellates are the
descendants of organisms which originally inhabited the intestinal tract of
vertebrates, and that the flagellates found so commonly in the digestive tract
of insects are derived from Hæmoflagellates, those insects in which they occur
being either blood-suckers or the descendants of blood-suckers. It would be
out of place to discuss the purely parasitological aspect of this question here,
but the subject is of sufficient importance to necessitate a consideration of the
evidence there is in favour of or against the hypothesis from the entomological
point of view.
In the first place, it must be remembered that the Diptera are simply a
highly specialised division of the class Insecta, that the Insecta are Arthropods,
and that the sucking mouth which is one of the main characteristics of the
Diptera is a modification of a biting mouth. The remote ancestors of all
Diptera were undoubtedly biting creatures, either animal or vegetable feeders,
or predaceous. Even in the Diptera the sucking mouth shows varying degrees
of development, and it is most improbable that there are any instances of
* And an undetermined Indian species.
3
reversion from the advanced condition to the earlier one. As I have shown
in a previous paper, the biting flies in the Orthorrapha have a relatively simple
form of apparatus, in which the primitive arthropod structures can be easily
traced, for piercing the skin of the host. The sucking portion of the mouth is
in fact formed by the simple apposition of the epipharynx and hypopharynx,
while the true segmental appendages retain a form not at all unlike that found
in quite primitive arthropods. Tabanus, for example, is a true "biting" insect,
in the sense that it makes a wound by means of its mandibles and first maxillæ;
the wound having been made, it sucks up the blood by means of a sucking
pump in the head, through the canal formed by the apposition of the epipharynx
and the hypopharynx, exactly as is done by non-biting flies which feed on
vegetable or animal juices,. The flies which possess functional mandibles and
maxillæ are undoubtedly nearer the primitive arthropod type than those which
have neither, and it is reasonable to suppose that, among the Orthorraphic
flies, such forms as Tabanus are more primitive than the flower feeders, and
that in those cases in which the female has a complete biting and sucking
mouth and the male only a sucking mouth, the female is the less removed from
the immediate ancestor of tho species. The loss of the appendages, in this case,
is an indication of specialisation. The mandibles and first maxillæ of the
biting flies of this group are clearly primitive structures retained, and their
existence is good evidence that the flies are more ancient than those which have
lost them, due regard being given of course to other characters. In this case
and in this limited sense the hypothesis of Minchin is supported by entomolo-
gical considerations, and it is quite possible that the flagellates present in
non-biting Orthorraphic flies have for their immediate ancestors similar
parasites which inhabited the intestinal tract of blood-sucking flies.
The case is quite different with regard to the Cyclorraphic flies. In no
case do these possess mandibles or first maxillæ, the only recognisable remnant
of these being the palpi, which are believed to be remainders of the first
maxillae. The biting apparatus is developed on the labella, which are derived
from the second maxillæ, appendages which take no part in the act of biting
in the Orthorraphic flies. Not only is the apparatus formed out of different
appendages, but it is of a different nature, and consists, as I have shown in the
case of Philœmatomyia and Lyperosia, of an arrangement of teeth which are
drawn through the wound by means of muscles situated in the labium. As
will be shown in the course of this paper, these teeth are represented in the non-
blood-sucking Muscids, and their development is the key to all the modifica
Lions found in the different forms
The labella, at the end of the fused second maxillæ, are modified in most of
the Diptera which do not suck blood into an apparatus of the of an absor-
4
bent pad, for the collection of fluid nourishment, which is sucked up the canal
in the proboscis by means of the pharyngeal pump, The commonest form of
this is the well known pseudotracheal membrane found in the house fly.
This form of apparatus is not peculiar to the Muscid flies, but occurs in the
Orthorraphic flies as well, and is absent only in those forms which obtain
their nourishment by actually piercing some resistant layer to obtain fluid
from underneath, and it is evident that its development is generally
coincident with the disappearance of the biting appendages. In some
instances, as for example the Tabanidæ,, it exists side by side with functional
and powerful mandibles and maxillæ, the fly being then able to absorb
fluid either from a moist surface or through a resistant dry layer which
has to be pierced by the stylets. In no case, however, does one find any
form of biting apparatus on the labella existing together with functional
mandibles or first maxillæ. The development of labellar teeth is a secondary
phenomenon, which takes place when a non-biting fly, the mouth of which is
adapted only for sucking fluids from moist surfaces, assumes the blood-sucking
habit. How this occurs in nature will be discussed in more detail when deal-
ing with those flies which are blood-suckers but are without biting mouth
parts.
We may sum up the evolutionary history of the blood-sucking Diptera as
follows. The early insect mouth parts were formed for biting and furnished
with simple arthropod jaws: the cockroach represents such a condition. At a
remote period the second pair of maxillæ became fused, a condition found in
all Diptera. The sucking mouth characteristic of the Diptera is formed by the
elongation of the epipharynx and hypopharynx, which together form a tube up
which the fluid nourishment is sucked. In the most primitive dipterous types the
typical arthropod appendages are retained, co-existent with the sucking tube,
and the insect is able to use its appendages to pierce a hole in skin or other
substance to obtain fluid food from underneath. Such a condition is found in
the Orthorraphic blood-sucking flies. At a later period the appendages of the
fourth and the fifth segments of the head, that is, the mandibles and maxillæ,
disappear, and the function of obtaining food is carried out by the second max-
illæ, the distal ends of which are modified to form a pad suitable for the absorp-
tion of fluid from moist surfaces. This condition is found in many families,
both in the Orthorrapha and the Cyclorrapha. It reaches its highest degree of
development in the complex labella of Musca. Still more recently some flies
have developed from this stage a most complex arrangement consisting of a
ring of teeth on the end of the labium, and have adapted themselves to a blood-
sucking habit. Throughout the evolution there is to be noticed a constant dimi-
nution in the importance of the anterior appendages, and the gradual transfer-
5
ence of the entire function of obtaining food to the second maxillæ. These
act in the more primitive forms merely as a sheath for the stylets, in the next
stage as an absorbent pad, and in the last stage, that of the blood-sucking
muscids, as cutting organs.
To return for a moment to the hypothesis of Minchin. All Diptera, and
one may add all Hemiptera also, are descended from biting or mandibulate
ancestors, but these ancestors were not necessarily blood-suckers, and conse-
quently flagellates found in non-blood-sucking flies may or may not be
descended from ancestors which were at one time hæmoflagellates. It is not to
be supposed that there has been any direct line of descent, and it appears to me
to be impossible to decide, in the present stage of our knowledge at least,
whether those forms, which in the past gave rise to the non-blood-sucking flies
we know to-day, were blood-suckers or not. Certainly they were provided with
mouth parts which could be adapted for the purpose with comparatively little
change, but there is no evidence of any importance one way or the other.
In the case of the Muscid biting flies Minchin's hypothesis can only be
accepted in a limited sense. Enough has been said already to show that in this
group the evolution has progressed from non-blood-sucker to blood-sucker and
that the flies of the Stomoxys group are more recent than the simple Musca
forms. That being the case, if the flagellates found in Muscid flies are descended
f r o m h æmo f l a g e l l a t e s , t h e n t h e y c a n o n l y h a v e r e a c h e d t h e f l y a t t h e r e mo t e
period when the fly was mandibulate, before the appendages had atrophied from
disuse. To be precise, if the ancestral stock of H erpetomonas muscœ-domesticœ
at any time inhabited the blood of a vertebrate, it must have changed its
habitat at that period in the evolution of the house fly when the mandibles and
maxillæ were present and functional, and after the outgrowth of the labrum-
epipharynx and hypopharynx had taken place; and this change can only have
occurred if one of the mandibulate ancestors of Musca domestica happened to
be also a blood-sucker. Before the formation of the sucking tube the insect was
purely a biter, and probably lived on solid or semi-solid food, and after the
disappearance of the mandibles and maxillæ it was purely a sucker, and could
not make the necessary wound.
The statement made above, that the Muscid biting flies have for their
immediate ancestors non-biting flies of the Musca type, is entirely in accord
with the other features of their structure and life history. A detailed exposition
of the reasons for the systematic position of the blood-sucking genera would be
out of place here, but two points of importance may be noted in passing. In the
first place, the venation of the wing shows characteristic changes. According
NOTE.—I have been unable to consult the paper quoted by Prowasek.—Fr. Brauer, Uber die Verbindungs-
glieder Zwichen den orthorrhaphen and cyclorrhaphen Dipteren, etc.—Verhandl. der zool-hot Gesellschaft,
1890. s . 273.
c2
7
The Proboscis of Musca.
Since all the flies to be discussed in this paper are to be regarded as derived
from a Musca-like ancestor, a brief note on the structure of the proboscis of
this type must be given. Descriptions of the proboscis of Musca and the allied
forms are numerous enough, and I have indeed little to add to what has
already been said, but it will be necessary, in view of what follows, to define
clearly what is meant by certain of the terms employed, and in certain cases
to introduce new ones. Any points which I omit, and regarding which the
reader may wish to refresh his memory, are dealt with in Kraepelin's complete
account. The species which I have specially studied, and to which the descrip-
tion mainly applies, is Musca nebulo, the common bazaar fly of Madras. The
differences between different species are however so small that the description
will apply in all the important points to any species of like habit.
The proboscis of Musca consists of three parts, termed from above down-
wards the rostrum, haustellum, and the labella. Between each two of these,
and between the rostrum and the head, there is a joint which permits of move-
ment in a vertical plane. When the proboscis is in use, the three joints are
ext ended so t hat t he t hree part s are i n t he same st rai ght l i ne, and t he whol e
proboscis hangs downwards from the under surface of the head of the fly,
approximately at a right angle to the long axis of the body. When the pro-
boscis is not in use it is retracted by the flexion of all three joints, in such a
way that the three parts are folded on one another and tucked into an orifice
on the under surface of the head. This orifice is termed by some systematists
the "buccal orifice," an unfortunate name, for it is separated from the true
mouth of the fly by the whole length of the pharynx, and is moreover merely
an interval in the chitinous integument of the head, and has nothing to do
with the buccal cavity. I propose to adopt the term epistomal orifice for this
aperture. Of the three divisions enumerated above, the rostrum is, strictly
speaking, a portion of the head. It contains the pharynx, and the only portion
of the true mouth parts in relation to it is the pair of maxillary palps, which
are attached to its anterior surface; the haustellum is the proboscis proper, and
consists of the surviving head appendages, viz., the second maxillæ, fused to
form the labium, and the labrum-epipharynx and hypopharynx. The labella
are parts of the second maxillæ which have remained separate at the distal end.
The nature of the joints and the range and nature of the movements of the
proboscis will be discussed later, but it should be noted here that the extension
of the rostrum is of the nature of an evagination of a part of the head capsule.
The so-called "buccal orifice" is merely a thickened rim where the rigid chitin
,of the head capsule gives place to the soft and flexible wall of the rostrum.