A Comparative Study of the Alimentary Canal in Butterflies, with Special Reference to Their Systematic Relationships Title (With 60 Text-figures) Author(s) HOMMA, Toshihiro Citation 北海道大學理學部紀要, 12(1-2), 40-60 Issue Date 1954-12 Doc URL http://hdl.handle.net/2115/27138 Type bulletin (article) File Information 12(1_2)_P40-60.pdf Instructions for use Hokkaido University Collection of Scholarly and Academic Papers : HUSCAP A Comparative Study ot the Alimentary Canal in Butterflies, with Special Reference to Their Systematic Relationshipsll By Toshihiro Homma (Zoological Institute, Faculty of Science, Hokkaido University) (With 60 Text-figures) I. Introduction So far as the writer is aware, the morphological works on the alimentary canal in butterflies have been published rather scantily, compared with those in other groups of insects. In butterflies, Bordas (1920), Dauberschmidt (1933), Dobkiewicz (1933) and others studied on the comparative morphology of the organ, and above all the work by Dauberschmidt has been valuable to his followers. The comparative morphology of the internal structures of butterflies may possibly give some suggestions on the taxonomy of this group, based mainly on the external characters; wing veins, colour patterns, palpi, antennae, legs and genitalia. From this viewpoint the present writer undertook the comparative study of the alimentary canal of butterflies, taking their taxonomic relationships into his consideration. In the present study the writer took up the following characters; mesenteron, anterior intestine, rectum and Malpighian tubules, and among them a special attention has been paid to the characters of the mesenteron, which were observed by him to be valuable among different groups. Before going further, the writer must express his cordial thanks to Professor Tohru Uchida for his kind guidance a.nd encouragement. The writer is also indebted to Messrs. ShOichi F. Sakagami, ShOzQ Ehara and other gentlemen for their continued interest and many useful suggestions upon the study. I I. Material and method The specimens used were collected by the writer throughout Hokkaido, except those of Erynnis montanus, Choaspes benjaminii, Papilio protenor nymphis, 1) Contribution No. 322 from the Zoological Institute, Faculty of Science, Hokkaido University. Sapporo, Japan. Jour. Fac. Sci., Hokkaido Univ., Ser. VI, Zool., 12, 1954. 40 A limentary Canals of Butterflies 41 P. memnon thunbergii, P. helenus niconiccolens, Eurema hecabe mariesi, E. laeta bethesba, Curetis acuta paracuta, Arhopala japonica and Niphanda fusca shijima which were collected in Honshu by his friends. The species examined include the following fifty-seven species belonging to six familiesll : Fam. Hesperidae Motschul~ky Subfam. Pyrginae Trib. Pierini 1. Erynnis montanus (Bremer) 21. Pieris rapae crucivora Subfam. Rhopalocamptinae Boisduval 2. Choaspes benjaminii (Guerin) 22. P. me!ete aglaope Motschulsky 3. Bibasis aquilina (Speyer) 23. A poria crataegi adherhal Subfam. Hesperiinae Fruhstorfer 4. Ochlodes venata (Bremer et . Fam. Lycaenidae Grey,... Subfam. Curetillae 5. O. ochracea rikuchina (Bremer) 24. Curetis acutol paraClda de 6. Ha!pe varia obscura Nakahara Niceville 7. Parnara guttata (Bremer et Subfam. Amblypodinae Grey) 25. Arhopara japonica (Murray) Fam. Papilionidae Subfam. Lycaeninae Subfam. Zerynthiinae Trib. ThecEni 8. Luhdorfia puziloi yessoensis 26. rtopoetes pryeri yezoensis Rothschild Nakahara Subfam. Parnassinae Trib. Strymonini 9. Parna.,sius stubbendorfii hoenei 27. Strymon w-album jenton£ Schweitzer (Butler) 10. P. glacialis glacialis Butler Trib. Lycaenini 11. P. eversmanni daisetsuzana 28. Lycaena phlaeas daimio (Seitz) Matsumura 5n fam. Plebejinae Subfam. Papilioninae Trib. Castaliini ,,> l~. Papilio machaon hippocratides 29. Tamka hamada hamada (Druce) Verity Trib. Lampidini 13. P. protenor nymphi$ 30. Niphanda Jusca shijima Fruh~torfer Fruhstorfer 14. P. memnon thunbergii von Trib. Glaucopsycbini Siebold 31. Glaucopsyche lycormas lywrmas 15. P. helen us niconiccolens Butler (Butler) 16. P. bianor .1·aponicus Butler Fam. Nymphalidae 17. P. maackii jezoensis Matsumura Subfam. Argynninae Fam. Picridae 32. Boloria thore jezoensis Subfam. Pierinae Matsumura Trib. Coliadini 33. B. Jreija asahidakeana 18. Eurema hecabe mariesi (Butler' (Matsnmura) 19 E. laeta bethesba (Janson) 34. Brenthis ino mashuensis (Kono) 20. Colias hyale poliographns 35. Argynni; paphia geisha 1) The classification applieo in this paper is based on the system of Esaki and Shirozu (1951). 42 T. Homma Hemming 48. Vanfssa indica (Herbst) 36. A. ch'lYlolta basalis Matsumura Subfam. Apaturillae 37. A. laodice japonica Menetries 49. Apatu.ra ilia substituta Butler 38. A. yuslana Motschulsky 50. Sasakia charonda charonda Subfam, Limenitinae (Hewitson) 39. Limenitis camilla japonica Fam. Satyridae Menetri~s 51. Ypthima argus argus Butler 40. Neptis coenobita ainu ShirClzu 52. Erebia niphonica doii Nakahara Subfam. Nymphalinae 53. E. ligea rishirizana Matsumura 41. A raschnia levana lev(lna (Linne) fi4. Oeneis daiselsu;mna Matsumura 42. A. burejana strigosa Butler 5;';. I.elhe callipteris diluta Esaki et 43. Polygonia c-aureum c-aureum Nakahara Linne 56. Aranda schrenckii me-nalcas 44. P. c-album hamigera (Butler) Fruhstofer 45. P I-album samurai Fruhstorfer 57. NeolJe gosthkevitschii 46. Nymphalis io geisha (Stichell goschkevitschii (MellCtrii's) 47. Aglais urticae connexa (Butler) The entire body, without dissected, was fixed and preserved in about 75% alcohol. In the present study, males were employed in all cases, with a few exceptions in which females were used. III. Observations The alimentar~r canal of butterflies is of a tubular form, being straight in some species or looped in others. The canal is distinctly divided into a stomodaeum, a mesenteron and a proctodaeum. The wall of the mesenteron is relatively tl).ick and the walls of the other two sections are thin and generally transparent. Two circular valve-like folds separate the three sections, one between the stomodaeum and mesenteron is stomodaeal valve, the other between the mesenteron and procto­ daeum is proctodaeal valve. In some species, the anterior part of the mesenteron surrounding the stomodaeal valve is distinguished as the cardia (Figs. 1, 10, 11, etc.). The stomodaeum is divided into the three regions; the pharynx, the oesophagus and the crop. The proventriculus is reduced to a slightly dilated tube. The crop is a large food reservior, which is connected with the stomodaeum by a short narrow duct. The mesenteron, which is called sometimes stomach, is of a somewhat flat tube or elongate sac of relatively small capacity and nearly uniform diameter, and in some species differenciated into two regions. The mesenteron is generally provided with many circular folds on its surface, and in some species, furnished with numerous processes on the surface of the anterior part, though variable in shape among species. The mesenteron seems to be of a remarkable specific characters, though sometimes slightly variable partially among individuals in its externals. The procotodaeum is divided into an anterior intestine, which is looped upon itself in most cases, and a rectum. The two regions are separeted externally by a constriction. The rectum is dilated anteriorly into a rectal sac, A limentary Canals of Butterflies 43 and narrows posteriorly to form a rectal proper which is represented by a straight tube and reaches directly the anus. Generally the anterior intestine opens into the rectum on the side of rectal sac, the anterior region of which forms a diverticulum, a rectal caecum, sometimes tinted yellow for its contents. The epithelium of rectal sac, and in a few cases, also that of the rectal caecum, forms numerous conspicuous structures known as Phy·········· ....SIGI the rectal papillae. The chief outgrowth of the alimentary canal forms a pair of Mal­ pighian tubules. On either side, the tubule Storn is generally divided into three branches at a common point, or, in other species, succesively at two points of a distance, as shown in Fig. 8. A type of the alimentary canal of bu~­ terflies is shown in Fig. 1. In this study, the mesenteron att­ Ment racted the writer's attention, as already mentioned. The following characters were observed in the alimentary canal of males, but, so far as the writer observed, in most species, the sexual difference in the externals of the canal is not ascertained. The technical terms used in this paper are mainly based on Aint ..... 0 •••• those of Snodgrass (1935). Fam. HESPERIIDAE Proe 1) Erynnis montanus (Bremer) (Fig. 2) Specimens examined: 5 males. The mesenteron is somewhat flat and straight tube, on the surface of which many circular folds appear, and the folds are bor­ dered with narrow grooves. The anterior part of the mesenteron is slightly wider than the 1 posterior part of that. The ant€rior intestine is approximately 2-3 times- as long as the Fig. 1. Parnassius stubbendorfii mesenteron. The rectal caecum is moderately hoenei Schweitzer. Stom, stomo­ developed, without papillae. The Malpighian daeum ; Ment, mesenteron; Proc, proctodaeum; Phy, pharynx; Oe, tubule is divided into three branches succes­ oesophagus; Cr, crop; Car, cardia; sively at two close points. SIGl, salivary gland; Mal, Mal­ 2) Choaspes benjaminii (Guerin) (Fig. 3) pighian tubules; Alllt, anterior Specimens examined: 5 males. intestine; An, anus; Rect, rectum The mesenteron bearing many circular (rect, rectum proper; rsc, rectal sac; rca; rectal caecum). folds all over