Comparative Anatomy of the Internal Organs of the Calyptrate Muscoid Flies V. Consideration on the Phylogeny
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Sc I. Rep. Kall <t za 押 a Ulliy. ， Vo 1. 12 ，No 圃 2 ，pp. 215~254 December December 1967 Co 聞 parative Anatom y of the Internal O lt' gans of the Calyp tl" ate Muscoid FHes V. Consideration on the Phylogeny of the Calyptratae I(atsushige I(atsushige HOIU Detarlment Detarlment of Biolo lJ j九 Faculty of Science ，I{a 押 azawa University (Rec 日ived 20 September 1967) Introdudion The Calyptr 同 ata 色* are considered as a rather well- c1 efined group which is fairly large large and originatecl monophyletically from the common acaJyptrate stock according to to somεrecent Dipterists. Owing to the vastness of thεnumb 巴r of closely 町田mbling species ， the cl 呂ssification of the Calyptratae has givεn rise to many clisputes for several several decades 岨 Practically no two authors of th 邑 Diptera have agreecl either on the subject subject of major groupings or the Calyptratae or the principle of the cJ assification to be be used. In most of these works ，with the exception of Townsen c! 's paper (1 935-42) and few others ， the taxonomic criteria for generic and supergeneric classification have mainly been the external morphological characters of the adult flies ， ancl those for for specific c1 assification the characteristics of the genitalia of both sexes as well as the the adult external morphology. The larval charactεrs ，with the exception of Townsencl (1 935-42) ，HaU (1948) ，Roback (1 951) ，etc. ，have not been usecl to any great extent in in the classificatio l1 of the Calyptrate Diptera. Most of the systems proposecl until now have been mainly Eounded upon the characters of the vεnation ， the antennae ， chaetotaxy ，etc ，、 togεther with the genitalia. 1n the past ，th 日refore ， it is not too much to say that the family Iin lI ts within the Calyptratae as well as the subfamily or or tribal limits within one family of the same have been subject to much shifting. It is ， therefore ， clear that the co-orclinate comparison of the external and intemal morphological morphological characters at a11 the stages is to be more reliable than the results obtained obtained from a comparison only in the extemal m01 ・phology of the adult stage. キ The terms Schizometopa ， Thecostomata ，Muscomorpha ，Calyptrata ，Calypterata ，Calypteratae ， Calypterae ，Calyptrates ，Calypterates ，etc" are also used by Dipterists 個 - 215 日 216 216 Katsushig 昨日ORI works this line have not yet been carried out in this field of study 。 The pr 巴sent unde 1' took the comparative anatomy of the internal organs organs of the adult flies which had been igno 1' ed many p1' evlous Dipterists (1 960. 印刷; 1962a; 1962b). The results obtained from these works ，which a1' e based on 83 species ，were summarized here and the data of every trait were arranged in ascending ascending or descending orders to clarify ， if pos ヨible ， the relationships amcng the different different groups of the Calyptratae. Some impo 1' tant external charactε 1' s adopted by the form 色l' Di pterists such as the the the fourth the squamae ， the notopleurals ， the hypopleu 1' als ， though most of these we 1' e used for the separation of of families ， genera or for the past 100 years or more ， are here also shortly treated treated in discussin 史 this f1' om thεevidences of external as as welL Since the materials in th1S serIes of papers a1' e very limited ， the conclusions conclusions should be as purely tentative ，and ， the 1' efore ， a great great deal of additional detailed works on many more mate 1' ia1s ， especially those bεto the Scatophagidae ， Tachinidae (s. lat.) and sevεra1 other small small groups will certainly shed on this inte 1' esting but difficult Such detail 己d a1' e beyond the of my present work. As far as the present author is aware ， the sciεntific names of seve 1' al of the treated treated in th i. s series of the author's work are recently revised by some Dipterists Dipterists as follows: ← Orthellia Orthellia lati) う Zimin → O 姐 caerulea Wiedeman l1 (see (see Hennig (1963): Die Fliegen.... ， pp. 928-9. J → D. Zetterstedt (see (see Hennig (1963): Die Fliegen.... ， p. 952. J Morellia 必oriorum (Fallen) ー→ M. Zimin [see [see H 邑nnig (1963): Die Fli 句 en.. ..， p. 976. J Bigot → senomera (Seguy) (see (see Hori (1964): Kontyu ， vo l. 32 ，p. 511. J Stomorhina discolor → St. obsoleta (Wiedemann) [see [see Kurahashi (1967): ScL Rep. Kanazawa Univ 吋 12(2): p岨 282J Onesia sp. → Melinda [see [see Hori (1961): Konty 日， vo l. 29 ，p. 194.J vomitoria vomitoria (Linné) ー~ c司 lata Coquillett [see [see Hori and Kurahashi (1963): Konty 日， vo l. 31 ，p. 284. J kinoshitai kinoshitai Hori → Sarcoj うhaga gravelyi (Senio 1' -White) 〔呂田 Kano et al.， (1967): Fauna Japonica ，Sarcophagidae (Insecta: Diptera) ，p. 91. J Many mordem Dipterists have attempted to divide the Sarcophagidae into many genera genera mainly 011 the 01 the male genitalia. This divisicn ，however ， is not here ， PhyloCJ{ フ幻:y of Calyttratae 217 The author wishes to offer his sincere thanks to Pro f. M. Kato of Tδhoku Univ 色rsity fm 、 his kind advicεan こJ criticisrn. The author' .s cordial thanks are also due to to Pro f. T. Okada of and D r. M. Tsukamoto of Nagasaki University for thε10an of tl1 eir 岨 own 1i teratm ・εancl also for their valuahle sllggestions 島 H :i storicals The history of the classific 旦tion of th 令 Calyptratae as well as that of other groups groups of insec .t s begins with Linnaeus (1 758) ，'1 /ho described three genera ，β([usca ， Oestrus ，and Co n. ops ， in which many extant 子 flies w 芭re contained. The systεmatic charact 色rs of the Diptera adopted by him 不Tere mainly the morphological clifferences clifferences of th 巴 The important systematic characters of the Calyptratae Calyptratae such as the size of the squamae ， habits ， immatllre stages ， ora1 ptilinu l11， head sclerit 巴s， transverse mesonotal suture ，cha 己totaxy ，etc. ，were clder Dipterists such as Harris (1 776) ，Dumeril (1 801 ， (1803 ， 1830 ， 1838) ， Latereille (1 805) ，Fall 白1 (1 814-20) ， Rondani (1 830 ， 186.3) ，Osten-Sacken (1881 ， 1884) ，Brau 巴r and Bergenstaη1m (1 and many others. The systematic values of of thεse characters 疋 d th 色se pione 色rs in this field are of great imp のrtance and of general use even at the IJr 色sent time. The erection of the Calyptratae is accrεdited to Robineau- Desγoidy. He ，accorcling to 0 ョtenωSacken (1 896) ， divided the Myodair 巴s into nine gWclpS ，which he called families 日ntes) ，and the first of these ， on the basis of the laγge size of their squamae ，he called Calypteratae. This division ， however ，was not of later authors ， because it did not not in c1 ude the The first major work on the Calyptratae was done by Girsclmer (1 893) ，who oIf ered a new system of classification on the basis of many characters of of the adult fl i. es. In this system he fi. rst divided the Muscidae Calypterae into two families ， and Tachinidae ，on the basis of the presence or absence of the hypopleural hypopleural hris t1 es ロ This Girsclmer's system is not so vviclely different from the rεcent ones. Girschner's Girschner's classification 1. 1. A. bteilung Musciclae A. calypterae Gattung Coenosia 1. 11. A. bteilung Muscidae Calypterae Gattung Dialyta I. I. Farnilie Anthomyidae 2. Grnppe Muscinae 1. 1. Gruppe Coenosiina 巴 Gattung 正lydrotaea Gattllng Gattllng Cordylu Ta Gattung 品わ IOstila Gattung Gattung ScatothaCJa Gattung Musca Gattung Gattung Fucellia G 旦ttung Stomoxys (ユ attung Lis 戸e Gattung Hae 討zatobia 218 218 Katsushig 邑 HORI GGGGaaaatttt!pttttdCLHUUUnnnnhafLggggdb GPMDK;YFBMyaa，必 rumzhm .mm10 。ム白白目白E cvmsbdMHumeGpM 国 e uu-wPPVEhPPOPE 広明化ゆ ・ QUA 品sqAVJ 門 V-rMdEI--Mmoe6nf 邑 由陶HbL加国叫mDm初 wnvEE- ttiJ4 M t- 凶 '庄 明巴出 た FjMW 仙 GeE5 官 dG mm ・沼ぽ 古 U yHAγi F 山町 市 dd T ・ mu.m 制白 児 ii 出 1 明「 旬 B .づ 山 凶 ， i UL 出 色 由L Modifying the systems of older 羽Torkers ，CoquilleU ， in 1901 ， proposed another syst 邑m. He believed that this arrangement would indic 計 e the natural relationships of of the various fam i1i es in a dearer manner than any of those that had been thus far proposed. proposed. Coquillett's Coquillett's classificatio l1 Suborder Suborder Proboscid 色呂 Family Sarcophagidae Section Section Orthorhaph 呂 Family Muscidae Section Section Cyclorhapha Family Anthomyidae Superfamily Superfamily Muscoidea Group Acalypterae Group Group Calypteratae Family Scatophagidae Family Family Oestridae Suborder Eproboscidea Fam i1 y Tachinidae Family Hippoboscidae Family Family Dexiidae Family Nycteribidae Williston Williston (1 908) offerεd a system on thεbasis of examination of the North Americanεns. In this system he divided the Calyptratae into six families. The are are induded in the The genera Limnothora ， l¥l lydaea ， Hydroj う Anthomyia and Liste are in c1 uded in the Anthomyidae. In the Muscidae most of the Calliphorid genera such as Callil 幼 ora ， Phormia and lYlesembrina are included in additio l1 to the typical Muscids. The are included in the Tachinidae. W iU iston's classification Acalypterae Acalypterae Oestridae Cordyluridae Cordyluridae Sarcophagidae Calypterae Calypterae Dexiidae Anthomyidae Anthomyidae Tachinidae 乱1uscida 記 To 干京w氾y s剖up 肝ε町rfami 江lieεs 丸 An此1社thomy loidea 勺 and Mu出1路sc ∞oidea 丸 bγ the 伊pr古巴 S巴nce or absεncε of 出the hypopleural hypopleural bristles ， pteropleural course patteγn of fourth longitudinal vein ， number and arrangement of stemopleural bristles ，etc. The genεra Myiostila ，Muscina ， etc 吋 are in c1 uded in the Anthomyioidεa. In the superfamily Muscoidea ， three famili 田; Macronychiidae ，Tachinidae ，Muscidae ， are recorded.