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The Respiratory System and the Mode of Respiration of the Water-Bug, <Emphasis Type="Italic">SphæRode

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The Respiratory System and the Mode of Respiration of the Water-Bug, <Emphasis Type= THE RESPIRATORY SYSTEM AND THE MODE OF RESPIRATION OF THE WATER-BUG, SPH.~RODT~'MA /~US7"ZCU~PI FABR., WITH REMARKS ON THOSE OF 2VEPA, LA CCO}'REP}IES AND RANA TRA. BY M- J. Pt~SSW&LLA AND 0. J. G~o~GF.. (From the Depart me~t of Biology, ~,ViIso~z Cotlcge, Bo,~7:bay.) Received August 19, 1935. (Communicated by Prof. P, R. Awati, I,~;.s.) I. Introd~ction ~nd Historical Survey. ALl, adult insects are air breathers, and whether they live in air or water they depend upon atalospheric air for their respiration. The immature stages of a few insects have become completely independent of atmospheric air by developing blood gills or tracheal gills for respiring dissoh'ed oxygen from the. water, while most are like the adult air breathers. This is so be- cause insects are primarily terrestrial creatures and have, like the aquatic mammals, taken to water secondarily, possibly on account of keen competi- tion on land. Amongst the adult insects a few of the Heteroptera and Coleoptera alone are perfectly adapted to live in water. These water-bugs and water-beetles carry a supply of air under their wing-covers and swim about freely in water, their hind legs acting as et~cient oars. While sub- merged they breathe this air by means of the dorsal th oracic spiracles. Occa- sionally they come up to the surface to renew this supply. When on the surface, they breathe directly from the atmosphere by means of the last pair of abdominal spiracles which is specially adapted for the purpose. Of the Keteroptera, the Cryptocerata are aquatic or semi-aquatic. The aquatic families of the Cryptocerata are: Naucorid~e, Be]ostomatida, Nepidm, Notollectidm, Pleid~e and Corixid~e,~all of these have peculiar adaptations which have solved the problem of breathing the atmospheric air under water. The study of these insects has occupied the attention of various workers, but except for Neifa and l~anatra of the family IXTepidm, detailed work has not so far been done on these insects, Of the earlier investigators, Dufour made a wide study of the aquatic I-lemiptera. He wasthe first (1821) to note the "tracheo-parenchylnatous" organs peculiar to the non-flying aquatic bugs, in NelOa and 7~ana~m. Martin (1896) pointed 280 Respimlio~z o/l/ze Wc~le~'-BuL Spheerodema rusficum Pa3r. 28t out certain respiratory adaptations in the nymphs of the Belostomatidm, Naucoridm and Nepid~. Dogs (1908), lV[aulik (19!6) and Brocher (t91.6) studied the respiratory system of Ne2bcz cin,ere~ in detail. Ferri~re (1914) made a comparative study of the "tracheo-parenchyrnatons" organs of Nef)a, Rar~atra and Naucoris. l~{uttkowski (1920) gave a collective review of the respiration in the aquatic insects. In 1921 l~{uttkowski made a valuable contribution on the r61e of insect blood in carrying oxygen. This function of insect Mood has thrown a new light on the study of the respiration of aquatic insects. Poisson (1924.) in his study of the aquatic IIenliptera dealt with the "tracheo-parenchymatous" organs, especially of Nep~. Recently, Lee (1929b) and Wigglesworth (1931) reviewed the subject of the respiration of insects. Hamilton (19.31) in her monograph on Nefia ci~,erea added to our knowledge of the respiratory system of this insect. The above records show that except for Nep~ detailed studies on the respiration of other aquatic insects i~ave not been undertaken, and there is a valuable field awaiting investigation. Spha~rodema rusticum I?abr. which forms the subject of this paper belongs to the family Belostomatid~, Apparently no work has been done on this insect, and while studying its respiratory system, the writers under- took also to work out its morphology. This has however been incorporated in a separate paper. The only record found on this water-bug is by Hoffmann (1926) who studied the life-history of this species, and described the five nymphal instars. The material for the present sttxdy was collected from a fresh-water tank in a suburb of Bombay. Together with Sphcerodema were also collected specimens of Laccotr@hes elongatus, L. grisea and species of Ra,~atra (Nepidse), Plea (Pleidse), Notonecta (Notonectidse) and Microvelia (ttydro- metrid,'e) and comparative observations made on them. Specimens of Nefba cir~erea were obtained for comparison of certain structures from the British lViuseum (Nat. I-Iist.). Sphcerodema r~sticum--The tzvo forms, Sphcerodema rusticum manifests a remarkable case of alary dimorphism. One form possesses normal wings while the other has degenerate wings. In the latter the hemelytra have lost the membrane portion and the inner wings their elltire anal region while the remaining portion of the inner wing 9is very much reduced in size. Intermediate forms with degenerate inner wings .but with the hemelytra having the membrane portion partly or fully developed .were also found. The differences in the wings have an important bearing on the structure of the thorax. The indirect wing muscles together with the phragma to which they are attached are found missing in the degenerate 282 M. J. Presswalla and C~ J. George winged ~orm, Wtlile in the fll~l winged fOrlll these parts are present, lint the wing muscles 1lave been Illodified tO fornl the peculiar si:rilctures called the " tracheo-parenchymatous" organs, Ass~ciated with. t'hese organs are a few air sacs ill the mesothorax which are also absent it.~ the degenerate winged form. The siguiticance of these differences ill l:he two forms is not quite clear. "1'he degenerate winged t!orm devoid of these sl;rnct~lres does not seem to be adversely affected in any way. Th.e general morphological characters a~ld those of the genitalia of the two agree and establish that th.ey are two forms of the same species. The presence of intermediate forms and a common habitat also sxlggest that they are conspeeitic. Leccotrephes and Ra.~aatm, collected along with Sflha'rodema, in- variably showed normal winged forms only, although these have also lost the power of flight and have their wing muscles modified in.to the "tracheo- parenchymatous" organs. ti. The Respiratory System of the Adult. 1, The Tracheal ,~vstem (Fig. 1). "i'here are two nlaia loilgitudinal tracheal trtmks (A.T.) x-trailing along the dorsal region of the abdomen. Posteriorly they ope~ on tSe ventral wall of two strap-shaped retractile organs (R.O.), which represent the modi- fied eighth abdominal segment, by a pair of spiracles (8th~Abd. Sp.). An- teriorly Each trtmk terminates by a very large sl)iraele (lst Abd. S17..) situated dorsally on the intersegmentaI membrane between the metanotum and the first abdominal segment, on the margin of.' the body. A very stout trachea, the thoracic spiracuIar trm~k (S.T.), connects this large spiracle wit11 a spiracle (Mt. Sp.) o~a the dorsal anterior side of the metathorax, below the base of the wing. From this the spiracular trunk is continued into the prothorax obliquely inwards, as the main thoracic trunk (T.T.). The trunks of the two sides converge into the head. In tlle an.terior part of the pro- thorax each of these trtm.ks bifareates into a dorsal, and a ventral branctl, the former (E.T.) supplying tracheoles to the head and. the brain, and th.e Iatter (R.T.) leadilag into the rostrum. ][immediately on entering the pro- thorax, on each o[ these obli(lt~e thoracic trunks opens a large oval spiracle (Ms. Sp.). This spiracle is situated on the ventral wall, attached to the intersegmental membrane between the pro- and meso-sterna. From each of the two abdominal trtmks (A.T.) are given off to the out- side six lateral abdominal branches (S.B.) in segments first, third, fourth, fifth, sixth and seventk, each terminating in a spiracle on th.e sternal wall. The first abdominal spiracular branch is very short and lies in the reduced first abdominal segment. This branch snpt)lies a duster of tracheae to the Respirafio,a o~ fhe l~Vate~'-]Jug, Sph~erodema rusticum Yaar. 283 Ms.Sp. .r e> ..... T 9 +~ Ms,T.c. ,-,~.+,,. /o.+.I/ f~ '+ "T. '8. (a) --J._ / / - ~' L.T. 3~dAbd (s) S.B. 4th J,~.~p.., (6) ~t~Abd.Sp. (~) 6~Abd.Sp, (8) (s) S tsA bd.sp. r FIG. I+ The general arrangement of the trachem in the adult. V R.O. 284 M. J. Presswa1Ja and C. J. George reprodut'tive organs in both the male and the female. 'rhe second abdominal segment heing very l1l\l{'h n·duced and complc.!tely ()v(.~r1apped by the 1irst it possesses uo spiracular trudleal hrandl und no spiradc. 'the remaining five lateral abdominal hranches lie ill segments three to seven and end in spiracles on the ventral wall of these Ht.~gmellts. 'the spiracular branches of the ahdomillul segments, thn'e to six, ('adl givt.~s out three secondary trachea!. One (A.C.'I'.) arising anteriorly ll<.'ar the inllcr eud supplies the digcl4tiVt' organs. A sc(~oncl (1'.13.) originutt's posteriorly front the middle of the spirucuhLr branch (\11(1 supplies the fut body and the body \vall of the sllIrotUldillg region. '1'he third bnl.1wh (V.'l\) is again anterior and emerges nearer the spimc1e. It immediately rUllS ventrally inwards al1d jOillS with the c.'orrespom1illg tm.l'l1t.~n. of the opposite side. It sends out tra<,'heoles to the ventral body wall of its segment. At its origin, all outer anterior branch (A.B.) supplies the outer (:onnexival n'giotl between the spiracle of its own segment and that of the preceding one.
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