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Abstract
The spiracular and tracheal systems of the lemon-butterfly, Papilio demoleus are described in detail. The
insect has 9 pairs of spiracles of which 2 are located in the thorax and 7 in the abdomen. Only the second
thoracic spiracle has the external closing mechanism, the rest having the internal closing mechanism. The
tracheae of the preabdominal region are largely thin walled, without taenidia and easily collapsible while
those of the abdomen have taenidia and are not easily collapsible. The tracheal air-sacs are largely present in
the preabdominal region. Some tracheae that d o not tracheate any organ but open into another trachea or
air-sac have been referred to as the tracheal shunts. Their possible function is to feed additional air to $he
branches or air-sacs into which they open. The spiracular tracheae maintain a fairly uniform (generalised)
branching pattern throughout different segments.
Introduction
Literature on the adult lepidopterous respiratory system, by and large, is
restricted only to the structure and function of the spiracles (Snodgrass 1935; Beckel
1956; Beckel and Schneiderman 1957; Tonapi 1959; Van der Kloot 1963; Mathur
1967),the tracheal system having been completely neglected. The present studies which
include both the spiracular and tracheal systems of an adult butterfly, Papilio demoleus
may, therefore, be of some interest.
abdominal trachea; TAS, air-sac trachea; TCB, cerebral trachea; TCD, dorsal tracheal
commissure; TCV, ventral tracheal commissure; TD1, TD2, first and second dorsal
trachea; TGM, ganglionic mass traceha; TG,, TGs, first and eighth (abdominal) terga;
TI, inner trachea; TMT, metathoracic trachea; TO, outer trachea; TOPS, optic-stalk
trachea; TPD, postero-dorsal trachea; TS1, TS2, positions of the first and second
thoracic spiracles; TT, tergal trachea; TTA, anterior tracheal trunk; TTD, dorsal
tracheal trunk; TTL, longitudinal tracheal trunk; TTPL, pleural tracheal trunk; TTV,
ventral tracheal trunk; TTVS, visceral tracheal trunk; V, valva; VPR, ventral process.
Observationsand discussion
1. The Spiracles
The butterfly has 9 pairs of spiracles: 2 in the thorax and 7 in the abdomen. Of these only the second
thoracic spiracle has the external closing mechanism, the rest having the internal closing mechanism.
Thoracic spiracles
The first thoracic spiracle lies in the pleural membrane of the prothorax close to the mesonotal
acrotergite (Fig. 1). The rather shallow atrium of the spiracle communicates with the exterior through the
spiracular (atrial) orifice and internally with the spiracular trachea through the tracheal orifice. The
spiracular orifice is confluent with a narrow annular sclerite, the peritreme, surrounding the orifice (Fig. 2).
The inner margin of the peritreme is prolonged into a number of cuticular and much branched processes
forming a sort of a thatched covering on the spiracular orifice and constituting a filter apparatus which
prevents entry of foreign particles into the spiracle. Beckel (1956) finds such processes to arise from the
bottom of the atrial chamber, Tonapi (1959) from the peritreme and Mathur (1967) from the inner wall of the
atrium. In the present insect, the filter apparatus is distinctly the product of the peritreme and, therefore, is in
agreement with Tonapi’s description. Mathur’s ‘inner wall’ does not make it clear whether the processes arise
from the outer region or the bottom of the atrium. The atrial wall of the present insect has become
differentially sclerotised into an internal closing apparatus. This apparatus has been likened to a bow
(Landois and Thelen 1867)having 3 distinguishable parts: a rigid arm, a flexible arm and an extension of the
latter, the lever or manubrium. The rigidarm (Fig. 3) forms the posterior half of the apparatus and is closely
approximated to the inner surface of the corresponding half of the peritreme. The approximation and
relatively heavier sclerotisation render this arm rigid or immovable. Theflexible arm constitutes the anterior
half of the apparatus and is connected to the corresponding half of the peritreme through a narrow band of
intervening membrane which together form the closing valve of the spiracle. Poorer sclerotisation of the
flexible arm as compared to the rigid arm results in a dorsal hinge which makes the arm flexible or movable.
Mathur’s ‘completely sclerotic semicircles’, obviously the equivalents of the above 2 arms in the present
insect, do not leave any room for a hinge and, therefore, the hinge lines shown by him are rather vague. The
lever is a slightly arched process projecting anteriorly and giving insertion to an occlusor muscle that
originates from the ventral extension of the rigid arm. Mathur describes the origin of this muscle from the
ventral end of the ‘spiracular rim’ which again is vague. The contraction of the occlusor muscle pulls the lever
down which in turn pushes the flexible arm backwards (towards the rigid arm) stretching the intervening
membrane across the tracheal orifice and screening it off from the atrium. A dilator muscle present in the
caterpillars of some insects (Snodgrass 1935)is absent in the adult of this insect so that the spiracle opens by
the relaxation of the occlusor muscle and/or elasticity of the lever.
c------i t----------r
2 0.2 m m
3 0.2 mm
- 0.2 mm
6
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FIGS.2-6.+2) First thoracic spiracle (outer view) after removing the scales from the pleural membrane
(stippled): (3) First thoracic spiracle (inner view) showing the internal closing apparatus: (4) Second thoracic
spiracle (outer view) showing external closing mechanism. Dotted lines indicate internal structures: (5) First
abdominal spiracle (outer view) showing modified body scales surrounding the spiracular orifice: (6)An
abdominal spiracle behind the first (outer view) showing the filter apparatus in the form of scaly-plug.
366 K. P. SRIVASTAVA
The second thoracic spiracle lies in the upper region of the segmental membrane between the meso- and
meta-pleuron but closer to the former (Fig. 1). The spiracular orifice ofthis spiracle is narrow and elongated
with its anterior margin overgrowing into a slightly hardened narrow plate, the anterior lip (Fig. 4). The
lower end ofthe anterior lip is drawn into a small ventral process which gives insertion to the occlusor muscle
originating from the outer surface of the inner end of the mesofurca. On contraction the muscle pulls the
ventral process downwards and forwards which draws the anterior lip backwards over the spiracular orifice
to cover it, thus constituting an external closing mechanism.
Abdominal spiracles
Of the 7 pairs of abdominal spiracles present in this insect, the first pair is located in the pleural
membrane near the base of the laterosternite (Fig. 1) with the remaining ones nearer the tergal margins of
their respective segments. The eighth (free) segment in the male is devoid o f a spiracle while the female which
has only 7 free segments has a spiracle for each segment. The first abdominal spiracle is not provided with a
filter apparatus though modified body scales with long pedicels and deeply notched margins densely
surround the spiracular orifice and seemingly prevent entry of foreign particles into the spiracle (Fig. 5).
Each of the remaining abdominal spiracles is provided with a filter apparatus formed by ordinary body
scales. The scales are inserted in overlapping rows (Fig. 6) on the anterior slightly sunken margin of the
spiracular orifice (Fig. 7) forming a sort of scaly-plug. The structure of the abdominal spiracles is similar to
that of the first thoracic spiracle in every respect except the following: (1) a peritreme is lacking in the
abdominal spiracles. This may be because the filter apparatus in these spiracles is formed by the body scales
and not by the cuticular processes of the peritreme, (2) their closing apparatus is a reversed replica of that of
the first thoracic spiracle i.e., the rigid arm is anterior and the flexible arm posterior so that the lever points
backwards, and (3) their closing apparatus extends much beyond the spiracular orifice at both ends. The
occlusor muscle is present in its normal position.
FIGS. 7, 8 . 4 7 ) An abdominal spiracle after removing the scaly-plug. Anterior sunken margin of the
spiracular orifice is shown heavily stippled and the internal structures partly dotted: (8) Dorsal mesothoracic
air-sac tracheating the dorsal longitudinal muscles.
RESPIRATORY SYSTEM OF PAPILIO DEMOLEUS 361
cause of formation of most of the larger air-sacs. A few tracheae that open into other tracheae or air-sacs
instead of tracheating any organ have been designated as the tracheal shunts whose function appears to be
to feed additional air to the branches or air-sacs into which they open. Though there is a fairly uniform
branching pattern of the spiracular tracheae* in different segments, the readiness of the tracheae to branch
off and supply adjacent organs results in occasional overlapping in tracheation. The following paragraphs
carry the description of the tracheation in different segments without considering the homologies of the
tracheae.
*Spiracular trachea refers to the short trachea that arises directly from the spiracle and is generally
inconspicuous.
?For descriptive convenience the immediate divisions of the spiracular tracheae have been
designated as trunks, their subdivisions as tracheae and further divisions as branches.
368 K. P. SRIVASTAVA
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FIG.9.-Tracheal system of the preabdominal (head and thoracic) region in dorsal view. Saccular tracheae
and air-sacs are shown without taenidial striations.
RESPIRATORY SYSTEM OF PAPILIO DEMOLEUS 369
T
T
TTV
FIG.10.--Tracheal system of the abdominal segments in dorsal view with portions of internal organs in a
male butterfly.
370 K. P. SRIVASTAVA
The upper branch ascends the dorsal tergal muscles and joins its counterpart of the other side to form the
dorsal abdominal airsac which tracheates the tergal muscles from the dorsal aspect. The anterior tracheal
trunk is conspicuous on account of its broad cylindrical shape and more strongly cuticularised taenidia. The
trunk curves inwards and then runs forwards joining the second thoracic spiracle. On its way it gives out
lateral tracheae which are saccular and coalesce with each other to give rise to the lateral metathoracic air-sac
tracheating the inner surface of the dorso-ventrally disposed muscles of the segment. The pleural tracheal
rrunk takes a lateral course and tracheates the outer surface of the dorso-ventral and other metapleural
muscles. The ventral tracheal trunk (not shown in the Fig.) is a delicate saccular trunk which descends
ventrad and joins its counterpart to form the ventral tracheal commissure like the ventral tracheal trunks in
other abdominal segments (vide infra). The commissure tracheates the overlying ventral nerve cord and
digestive organs.
Acknowledgments
The author wishes to thank Mr A. N. Singh, the artist of the Department, for his
help in the preparation of the illustrations.
References
BBCKEL,W. E. (1956).-The morphology. histology and physiology of the spiracular regulatory mechanism
of Hyalophora cecropia (Lepidoptera). Proc. Tenth Internat. Congr. Ei7t. Montreal. 2:85-115.
BECKEL,W. E. and SCHNEIDERMAN, H. A. (1957).-Spiracular closing mechanism in Hyalopliora. Sc,ience.
126~352-353.
EPHRUSSI,B. and BEADLE, G. W. (1936).-A technique of transplantation for Drosophila. Amer. Naturalist.
70:218-225.
IMMS.A. D. (1963).-"A general textbook of entomology" (Asia Publishing House: Bombay).
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Dytiscus larven. Flugers Arch. ges. Physiol. 179:113-120.
*LANDOIS. H. and THELEN, W. (1867).-Der Tracheenverschluss bei den Insecten. Zeitschr. wiss. Zool.
17:187-214.
MATHUR, R. B. (1967).-The spiracular mechanism in the Indian Sann hemp moth, Utetheisa pulchella
L h n e (Lepidoptera: Arctiidde). Mushi. 40(11): 139-146.
SNODGRASS, R. E. (1935).-"Principles of insect morphology" (McGraw Hill Book Co.: New York and
London).
TONAPI,G. T. (1959).-On the structure and mechanism of the spiracular regulatory apparatus in the adult
Corcyra cephalonica Staint. Czrrr. Sci. 28(11):457-458.
VANDER KLOOT,W. G. (1963).-The electrophysiology of the nervous control of the spiracular muscle of the
pupae of the giant silkmoth. Comp. Biochem. Ph.vsio1. 9:317-333.
*Not seen in original.
[Manuscript received April 11, 19751