Sc I. Rep. Kall

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. The iam i1 y Muscidae are divided into into two subfamili 田 Calliphorinae and Muscinae. In the subfamily Muscinae the g巴nus Mesemhrina is in c1 uded together with typical Muscids such ;;l S Musca ， Stomoxys ，Morellia ， etc. Phylogeny Phylogeny 01 Caty ρtratae 219

Townsend's classification

Order Order Diptera Family Tachinidae Suborder Suborder Orthorrhapha Family Muscidae Suborder Suborder Cyclorrhapha Subfamily Calliphorinae Series Series Aschiza Ge nus Calliphora Series Series Schizophora Genus Eucalliphora Section Section Myodaria Ge nus Lucilia Subsection Subsection Schizometopa Ge nus Proto ρ hormia Superfamily Superfamily Anthomyioidea Subfamily Muscinae Ge nus Myiospila Genus Musca Genus Genus M ω cina Family Phasiidae Superfamily Superfamily Muscoidea Section Pupipara Family Family Macronychiidae

In In 1911 Schnabl and Dziedzicki published their voluminous work ，Die Anthomyiden ， in in which the Anthomyidae an~ divided into three groups. The genus Liste is included ぶ in in the Mydaeidae. The genera Grathomyia ，Muscina ，Morellia ，Musca ，Sto 仰 oxys ， Lyterosia ，etc. ， are placed in the Muscinae Ordo 1，and in the Muscinae Ordo II the genera Mesemorina ，DasYl うhora ，Pyrellia ，etc. ， are included. The classification adopted by Schnabl and Dziedzicki in this work is based largely on the male genitalic characters. characters.

Schnabl and Dziedzicki's classification

Anthomyidae Anthomyidae A. Mydaeidae 1. 1. Gruppe: Coenosinae B. Muscinae Ordo 1 2. 2. Gruppe: Anthomyinae 4. Formenreihe: 1. 1. Formenreihe: Hylemyidae A. Aricinae 2. 2. Formenreihe: Homalomyidae B. Muscinae Ordo II 3. 3. F ormenreihe: 3. Gruppe: Gastrophilinae

Stein Stein (1 918) divided the Muscidae Calyptratae into two families; Anthomyidae and Tachinidae ，on the bareness or plumosity of the hypopleuron. Seguy (1923) divided the Anthomyidae into two subfam i1i es; Anthomyinae and Gasterophilinae. Gasterophilinae. In his Anthomyinae ， eight groups ， that is ，subfam i1i es ，such as Fanninae ，Stomoxydinae ，Muscinae ，etc. ， are recognized.

Seguy's classification

Anthomyidae Anthomyidae 66 Groupe Ariciinae An thomyinae (s. 1.) Muscinae (s. 1.) 16r Groupe Co enosiinae Muscinae (1 6r groupe) ou Muscinae 26 Groupe (a). Hylemyinae limnophoraeformes Schnabl (b). (b). Pegomyinae 76 Groupe Stomoxydinae 36 Groupe Pegomyinae transitoriae 86 Groupe Muscinae (s. s.) Eginiinae Eginiinae Muscinae (2 6 groupe) ou Muscinae 46 Groupe Mydaeinae polietesformes Schnabl 56 56 Groupe Fanniinae Gasterophilinae

Shannon (1 923) endeavoured to maintain the Calliphorids as a distinct family. 220 220 Katsushige HORI

Shannon's classification

Family Family Calliphoridae Subfamily Caliiphorinae

Subfamily Subfamily Phormina 日 Tribe Lucili 且i Trib 色 Chrysomyini TrihεCalliphorini Tribe Tribe Phorrnini

Senio ト White ( divided the mto two Muscidae and Tachinidae ，and in the former three subfamilies are the latter flve subfamilies are listed.

Senior- White's classification

Acalyptrata Acalyptrata Family Tachinidae CaJyptrat 呂 Subfamily Calliphorinae Fa l11 ily IVlusddae Subfamily Rhiniinae Subfam i! y Anthomyiinae Subfam i! y Sarcophaginae

Subfamily Subfamily Muscinae Subfamily D 己xiinae Subfamily Subfamily Stomoxydinae SubfamHy Tachininae

Imms (1 925) divided the section or of Diptera into the two undennentioned The are inck;ded in the 正¥calypterae ，and the are subdivided into three families. Both the IViuscids and Glossinids ar 己 includεd in his The Tachinidae are again subdi 可rided into three subfamilies 。

Imms's classification

Superfamily Superfamily Acalypterae Subfm 百 ily Sarcophaginae Family Family Cordyluridae Subfamily Calliphorinae Superfamily Superfamily Calypterae Subfamily Tachinina 巴

F、amily Anthomyidae Family 0 己steridae Family Family Tachinidae

仁omstock (1 924) dividεd the section into two were assigned to to the Acalyptrat 田 In the Anthomyioidea and Muscoidea ， are recognized ， The Muscoidea are subdivided into eight families. families. The areεach treated as a distinct family in the 1'¥、 Auscoidea.

Comstock's classification

Section Section 1. Myodaria Family Oestridae Subsection Subsection 1. Acalyptratae Family Phasiidae Family Family Cordylurida 日 Family i) !!egaprosopidae

Subsection Subsection II. Calyptrataε Fa l11 ily Calliphorida 邑 Superfamily Superfamily Anthomyioidea Famiiy Sarcophagida 巴 Family Family Anthomyiidae Family Tachinida 巴 Superfa l11 uy Muscoidea Family 恥1uscidae

Family Family Gastroph i!i da 芭 Pli ザlageny 0/ Caly β tratae 22j

Lundbeck (1 927) diviclecl the Schizometopa ， or CalyptraLae ， into two large families;

Antho ll1 yidae and Tachinidae ， ancl the laUer are again subcliviclecl into nine subfa ll1 ilies.

Lunclbeck's Lunclbeck's classification

Family Family Anthomyidae Subfamily Calliphorinae Family Family Tachinidae Subfam i! y Sarcophaginae Subfamily Subfamily Gastrophilina 巴 Subfamily Rhinophorinae

Subfamily Subfamily Oesterinae Subfarnily Dexiin 呂e Subfamily Subfamily Hypoderminae Subfamily Tachininae Subfamily Subfamily Phasiinae

Karl (1 928) clividecl th 巴 Muscidae into six sllbfamilies.

Karl's Karl's classification

Fa 、、冶加a Tribus Mydaeini

LTn 川叫1te 釘rf 臼a口mi 江lie Mus 民cinae e Tribus Li mnophorini Tribus Tribus Muscini Unterfam i1i e Mydaeinae Tribus Tribus Stomoxydini Unterfam 日ie Anthomyinae Unterfamilie Unterfamilie Phaoniinae Tribus Anthomyini Tribus Tribus Phaoniini Tribus Fucelliini

Tribus Tribus Hydrotaeini Unterfamilie Co 己nosiinae Tribus Tribus Fanniini Tri bus Chelisiini Unterfamilie Unterfamilie Eginiinae Tribus Coenosiini Tribus Tribus Azeliini

H 巴nclel (1 928) clivided the Mllscoiclea ，or Calyptratae ， into the Haplostomata ancl

Thecostmata ，the for111er correspond to the 恥'!uscicla 巴 acalyptratae ancl the laUer to

the Musciclae calyptratae. The Thecostomata are dividecl into four fa ll1 ilies. The family Corclyluriclae are inclllclecl in the Thecostomata. The Calliphoricls and Sarcophagids are recognizecl each as a subfamily in the Larvaevoridae.

日告 nclel's classification

Superfam. Superfam. Muscoid 巴 a (Schizophora) Fam. Larvaevoridae (Tachinidae)

Haplostomata Haplostomata (Muscidae acalyptratae ， Unterfam. Calliphorina 告 Myodaria Myodaria inferiora) Unterfam. Sarcophaginae Thecostomata Thecostomata (Muscidae calyptratae ， Unterfam. Rhinophorinae Myodaria Myodaria superiora) Unterfam. Phasiinae Fam. Fam. Cordyluridae Unterfam. Dexiinae

Fam. Fam. Oestridae Unterfam. Larvaevorina 巴 (Tachininae) Fam. Fam. Muscidae

Using the hypopleural charact 巴r，viz ， bareness or plumosity ，Wainwright (1 928) cliviclecl cliviclecl the Calyptratae into two large families: Anthomyiclae and Tachiniclae. The Calliphoricls ，Sarcophagicls ，Tachinicls ，etc. ，are recognized respectively as a subfamily of of the Tachinidae. 222 222 Ka i.: sushi 即日ORI

classification classification Calyptratae Calyptratae Subfamily Sarcophaginae Family Family Anthomyidae Subfamily Calliphorinae Family Family Tachinidae Subfamily Phasiinae Subfamily Subfamily Tachininae

Seguy (1 928) divided the Oestridae (auc t.， s. into two families:

Anthomyiidae and Tachinidae. In the Tachinidae thr 巴e subfamilies are recognized. The subfamily Calliphorinae of this family are large and contain seven tribes.

Seguy's classification

Oestridae Oestridae auct. ，s. 1. Oestrocalliphorini Anthomyiidae Anthomyiidae Rhyncho 巴strini Anthomyiinae Anthomyiinae Microcerellini Gasterophilinae Gasterophilinae Sarcocalliphorini Tachinidae Tachinidae Eucalliphorini Oestrinae Oestrinae Polleniini

Hypoderma Hypoderma tinae Rhi 且iini Calliphorinae Calliphorinae

Malloch divided the family Muscidae ioto seven subfamilies.

Malloch's Malloch's classification

Family Family Muscidae Subfamily Li spinae

Subfamily Subfamily Anthomyiina 巴 Subfamily Phaoniinae Subfamily Subfamily Fanniinae Subfamily Stomoxydin 初 Subfam i1 y Co 邑nosiinae Subf 呂mily Muscinae

Curran (1 934) recognized six subfamilies in the In his Muscidae ， the Scatophagids ，Anthomyiids and some members of Muscids which lack the hypopleural bristles bristles are in c1 udedo The Metopidae are newly erected for the combination of the the Sarcophagids ，Calliphorids ，some members of Muscid8 which have hypopleural bristles ，and a few Tachinids.

Curran's Curran's cla8sification Family Family Muscidae Family Tachinidae

Family Family Gasteroph i1i dae Family Oestrida 色

Family Family M 己topiidae Family Cuterebridae

Townsend (1 935-42) made an extensive contribution to the study of the

Calyptrate Calyptrate Di pt 日ra of the entire world. In his work the Rhiniida 巴 are proposed for the the combin 且tion of the Mesembrinellini ，Benga1iini ， Rhiniini and Polleniin i. The Stephanostomatidae are a180 recognized as a distinct family for the combination of the the Agrini ，Miltogramini ，Metopini and their alli 問。 But his work ，though excellent ， 18 18 far from the currεnt systems now used students. Enderlein Enderlein ( divided the iuto three “ Protomuscaria ， Phyloge1 ょ:y 01 Calyttrat αe 223

Acalyptrata ，and Calyptrata. The family Cordyluridae ancl Scatophagidae are assigned to th 巴 Protomuscaria. 1n this system he recognized three superfamilies in the Calyptrata; Muscoiclea ，Protachinoidea ， ancl Tachinoiclea. 1n the superfamily MllSCoidea the Stmoxidae are entered as a separate family together with the Anthomyiidae ， Mllscidae. The Gastrophilidae are also in c1 11decl in this sllperfamily. Fannai and its allies allies are listed as a tribe Fanniini in the subfamily Phaoniinae. The members of Liste Liste are assigned to a separate subfarnily in the Anthomyiidae. The superfamily

Tachinoicl 巴a are dividecl into seven families; Phasiiclae ，Hypodermidae ，Calliphoridae ， Sarcophagidae ，Rhinophoriclae ，Tachiniclae ancl Dexiidae.

Enderlein's c1 assification

Untergruppe Untergruppe Protomuscaria Familie Bypodermidae Familie Familie Cordyluridae Familie Calliphoridae Untergruppe Untergruppe Acalyptrata Unterfamilie Phor ・ffilnae Untergruppe Untergruppe Calyptrata Tribus Chrysomyiini Superfamilie Superfamilie Muscoiclea Tribus Phormiini Familie Familie Anthomyiidae Unterfamilie Ca l1i phorinae Unterfamilie Unterfamilie Phaoniinae Tribus Calliphorini Tribus Tribus Phaoniini Tribus Luciliini

Tribus Tribus Hydrotaeini Unterfamili 巴 Poll 巴niinae Tribus Tribus Fanniini Unterfamilie Engyzopinae Unterfamilie Unterfamilie Li spinae Unterfamilie Rhynchomyiinae Unterfamilie Unterfamilie Mydaeinae Tribus Rhynchomyiini Tribus Tribus Azeliini Unterf 品milie Rhiniinae Tribus Tribus Mydaeini Familie Sarcophagidae Unterfamilie Unterfamilie Fucelliinae Unterfamilie Moriniinae Unterfamilie Unterfamilie Anthomyiinac Unterfamili 巴 Sarcophagina 巴 Unterfamilie Unterfamilie Coenosiinae Tribus Sarcophagini Familie Familie Muscidae Tribus Bellieriini Unterfamilie Unterfamilie Morelliinae Unterfamilie Miltogrammatinae Unterfamilie Unterfamilie Pyrelliinae Tribus M 巴topiini Unterfamilie Unterfamilie Muscinae Tribus Miltogrammatini Familie Familie Stomoxidae Unterfamilie Agriinae Familie Familie Gastrophilidae Tribus Agriini Superfamilie Superfamilie Protachinoidea Tribus Nyctiini Familie Familie Eginiidae Familie Rhinophoridae

Superfamilie Superfamilie Tachi 日 oidea Familie Tachinidae Familie Familie Phasiidae Familie Dexiidae

Rohdenclorf (1 937) dividecl the Sarcophagiclae into six families.

Rohclendor f' s classification

Familie Familie Sarcophagidae Subfamilie Chrysogrammatinae Subfamilie Subfamilie Sarcophaginae Subfamilie Metopiinae Tribus Tribus Sarcophagini Snbfamilie Macronychiinae Tribus Tribus Raviniini Subfamilie Mimodexiinae Subfamili Subfamili Agriinae 224 224 Katsushige HOR!

Senior-White ， Aubertin and Smart (1940) divided the Calyptrata into four families; families; Muscidae ，Oestridae ，Calliphoridae ，and Tachinidae. The Sarcophagids are retained retained as a subfam i1 y in the Calliphoridae.

Senior-White ，Aubertin and Smart's classification Calyptrata Calyptrata Subfamily Chrysomyiinae Fam i1 y Muscidae Subfamily Rhiniinae Fam i1 y Oestridae Subfamily Sarcophaginae Fam i1 y Calliphoridae Family Tachinidae Subfam i1 y Calliphorinae

Hennig (1 941) recognized twelve fam i1i es in the superfamily Calyptratae. The

Rhiniids ，乱 f巴lanophorids ，Phasiids ，Dexiids ，Prosenids ，Exoristids ，etc. ， are treated as a distinct family respectiv e1 y. In t. he Anthomyiieae eight subfam i1i es; An thomyiinae ， Fanniinae ，Lispinae ，Coenosiinae ，Phaoniinae ， Eginiinae. Muscinae ，and Stomoxydinae ， are are listed. The genus Ophyra is included in the subfam i1 y Phaoniinae.

Hennig's Hennig's classification

Superfamilie Superfamilie Calyptratae Fam i1i e Exoristidae Familie Familie Co rdyluridae Familie Anthomyiidae Familie Familie Rhiniidae Subfam i1i e Anthomyiinae Fam i1i e Ca lliphoridae Subfamilie Fanniinae Familie Familie Sarcophagidae Subfamilie Li spinae Familie Familie Melanophoridae Subfamilie Coenosiinae Fam i1i e Phasiidae Subfamilie Phaoniinae Familie Familie Dexiidae Subfamilie Eginiinae Familie Familie Oestridae Subfamilie Muscinae Fam i1i e Prosenidae Subfamilie Stomoxydinae Fam i1i e Tachinidae

Crampton (1942 ・4) divided the Calypteratae into two principal superfam i1i es ， Muscoidea and Tachinoidea ，on the basis of the study of the male genitalia. The superfam i1 y Muscoidea contain four families; Scatophagidae ，Glo 悩 inidae ，Anthomyiidae and Muscidae. The superfamily Tachinoidea include the Phasiidae ， Dexiidae ，

Tachinidae (or Larvaevoridae). 0 己stridae ，Cuterebridae ， Sarcophagidae (or Metopiidae) ， Calliphoridae ，and Gastroph i1 idae. The genitalic characters of the families Cuterebridae and Gastrophi 1i da 巴 were generally consider 巴d sufficient to place them as a separate Oestroid Oestroid superfamily. He thought that if the Cordyluridae were to be regarded as a true true Calyptratae ， it might be advisable to place them in a separate superfam i1 y Cordy luroidea.

Crampton's classification Calypteratae Calypteratae Family Anthomyiidae Superfamily Superfamily Muscoidea (or Muscomorpha) Fam i1 y Muscidae Family Family Scatophagidae Family Glossinidae Phylogeny Phylogeny of Calyttraiae 225

Sllpcrfamily Sllpcrfamily Tachinoidea Family Cllterebri c1 ae Family Family Phasiida 巴 Family Sarcophagidae Family Family Dexiidae Family Calliphori c1 ae Family Family Tachinid 乱e F、amily Gastrophilidae Family Family Oestridae

Tokunaga (1 943) concurred with Williston 's view (1908) ， ancl clivicle c¥ the

Calyptratae Calyptratae into five families; Anthomyiidae ， Musciclae ，Sarcophagiclae ，0 巳stridae and Tachiniclae. The Muscinae ， Calliphorinae ， Glossininae ， Stomoxyidinae and Philaematomyiinae are retainecl respectively as a separate subfamily in the Muscidae. ln ln the system propose c1 by Mesnil (1 944) ， three families ，Musciclae ，Calliphori c¥ ae and Larvaevoridae ， are recognized in the Thecostomata ， or Calyptratae. The

Calli Calli phorina 色， Hypoderminae ，Sarcophaginae and Rhinophor ~nae are al1 retained as a subfamily subfamily in the Calliphoridae.

Mesnil's Mesnil's classification

L I:- I凡plostomata c. Sarcophaginae 1. 11. Thecostomata c1. Rhinophorinae A. A. Muscidae C. Larvaevoridae B. B. Calli phori c1 ae a. Phasiinae a. a. Calliphorinae b. Dexiinae b. b. Hypoderminae c. Larvaevorinae

The system adopted by Essig (1 947) in his text 四 book followed in many points that that of Enderlein. HaU (1 948) diviclecl the Calypteratae into two groups ， Anthomyiaria and Oestromuscaria ，and the latter into three superfamilies; Oestroldea ，Muscoidea and Sarcophagoidea. Sarcophagoidea. In the Sarcophagoiclea ，two families ， Sarcophagidae ancl Calliphoridae ， are are adopted. Th 巴 Calliphorida 己 are further dividecl into five subfamilies.

Hall's Hall's classification

Calypteratae Calypteratae Family Calliphoridae Anthomyiaria Anthomyiaria Su bfamily Mesem brinellinae Oestromusωna Subfamily Rhiniina 巴 Oestroidea Oestroidea S 1.l bfamily Chrysomyinae Muscoidea Muscoidea Subfamily Calliphorinae Sarcophagoidea Sarcophagoidea Su bfamily Polleniinae Family Family Sarcophagidae

ln ln the system adopt 己d by Colyεr and Hammoncl (1 951) four families ，Corclyluridae ，

恥1uscicla 巴， Calliphorida 己 and L 乱rvaevorida 色， are recogmz 巴d in the Calypterat 巴s. The family family Calliphoridae are clividecl into three subfamilies; Calliphorinae ， Sarcophaginae and Rhinophorinae. The family Muscidae are also subdivicled Into eight subfamilies. 226 226 Katsushige HORI

Colyer and Hammond's classification

Cyclorrhapha Cyclorrhapha Family Muscidae (Anthomyiidae) Acalypterates Acalypterates Subfamily Muscinae Calypterates Calypterates Subfamily Gasterophilinae Family Family Cordyluridae (Scatophagidae) Subfamily Phaoninae Family Family Larvaevoridae (Tachinidae) Subfamily Lispinae

Fam i¥ y Calliphorida 邑 Subfamily Mydaeinae Subfamily Subfamily Rhinophorinae Subfamily Fucellinae Subfamily Subfamily Sarcophaginae Subfamily Anthomyiinae Subfam i1 y Calliphorinae Subfamily Coenosiinae

Zimin (1 951) divided the subfamily Muscinae into two tribes ， Muscini and Stomoxydini ，and the genus Graphomyia Is transferred to th， e tribe Stomoxydin i.

Zimin's classification

Family Family Anthomyidae Genus Dasyphora Family Family Muscidae Genus M orellia

Subfamily Subfamily Muscinae Genus Mese 附 brina

Trib 色 Muscini Tribe Stomoxydini Genus Orthellia Genus GraPho 附 :yia

Genus M 'zI， sca Genus St o. 附 oxys Genus Pyrellia Genus H aematobia Genus Rytellia Genus Lyperosia

The Calyptratae 昌re divided into three superfam i1i es ，Muscoidea ，Sarcophagoidea and Oestroidea Roback (1 951). The f乱nnines ar 巴 advanced to a family of the Muscoidea. The Scopeumatinae are included together with the Anthomyiinae in the family Anthomyiidae. Th 巴 Stomoxyids are recognized 出 a separate subfamily

Stomoxyina 色 in the Muscidae. 111 the Sarcophagoidea two families ，Calliphoridae and

Sarcophagidae ，are listed ，

Roback's classification

Calyptratae Calyptratae Superfamily Sarcophagoid 邑a Superfamily Superfamily Muscoidea Family Calliphoridae Family Family An thomyiidae Subfamily Mesembrinellinae

Subfamily Subfamily Anthomyiina 巴 Subfamily Rhiniinae Subfamily Subfamily Scopeumatinae Subfamily Chrysomyinae

Family Family Fanniida 巴 Subfamily Calliphorinae Family Family Muscidae Subfamily Polleninae Subfamily Subfamily Coenossiinae Family Sarcophagidae Subfamily Subfamily Li spinae Subfamily Miltograminae Subfamily Subfamily Phaoniinae Subfamily Sarcophaginae Subfamily Subfamily Muscinae Superfamily Oestroidea Subfamily Subfamily Stomoxyinae

In the Muscidae (=Anthomyiidae) Ringdahl (1 954-9) rεcognized nine subfamilies.

The genera Gral うhornyia and Muscina are includεd in the Muscinae. Phylog 仰 y of Calyttratae 227

Ringdahl's classification

Holometopa Underfa l11. Fanninae Schizometopa Schizometopa Unclerfa l11. Mydaeinae Fam. Cordyluridae Unclerfam. Li spinae Fa l11. Muscidae (Antho l11 yiidae) Underfa l11. Coenosiinae Underfam. Muscinae Und 巴rfam. Fuce !li nae Underfam. Stomoxydinae Underfa l11. Anthomyiinae Underfam. Phaoniinae

Van Emden (1 954) divided the Calyptrata into four families; Cordyluridae ，

Muscida 巴， Calliphoridae and Tachinidae. The Calliphorinae and Sarcophaginae appear respectively as a separate subfamily in the Calliphor ・idae.

van Emdεn's classification

Calyptrata Calyptrata Subfam i1 y Tachininae Fa l11i1 y Cordyluridae Subfamily Goniinae Fa l11 ily Muscidae Fa l11i1 y Ca 1li phoridae Fa l11 ily Tachinidae Subfamily Rhinophorinae Subfa l11 ily Phasiinae Subfam i1 y Sarcophaginae Subfa l11 ily Dexiinae Subfamily Polleniinae

Subfa l11 ily Macquartiinae Subfamily Calliphof 一 inae

1n his text-book Shiraki (1 954) adopted the system proposed by Enderlein (1 936). Hennig (1955-64) retained the fannines in the subfamily status ，and included the genus Gra ρhomyia together with Myostila in his Mydaeinae and the genera

Othyra ，Muscina ，etc. ， in his Phaoniinae. The genera Stomoxys ，Sithona ，巴tc. remain as a tribe in the Muscinae.

Hennig's classification

Familie Familie Muscidae Gattung Hydrotaea Unterfamilie Unterfamilie Fanniinae Tribus Phaoniini Gattung FannIa Gattung Muscina Unterfamilie Unterfamilie Mydaeinae Gattung Phaonia Tribus Tribus Azeliini busTri Dichaetomyini

Tribus Tribus Mydaeini Gattung Dichaeto 附 yia Gattung Myostila Unterfa l11i1i e Muscinae Gattung H elina Tribus Muscini Gattung Gattung Gra. ρ 'homyia Gattung M ω embrina Tri Tri bus Lil11 nophorini Gattung Orthellia Gattung Gattung Limnothora Gattung Rytellia Tri Tri bus Li spini Gattung Pyrellia Gattung Gattung Liste Gattung Dasythora

Tribus Tribus Co 巴nosiini Gattung M orellia Gattung Listocethala Gattung Musca Unterfa l11 ilie Phaoniinae Tribus Sto l11 oxyini Tribus Tribus Achantipterini Gattung Stomoxys Tribus Tribus Hydrotaeini Gattung Stygeromyia Gattung Pegonomyia Gattung Siph 側耳 Gattung Ophyra 228 228 Katsushige HORI

In the Calyptratae Herting (1 95 '7) recognizes six familiεs; Muscidae ，Cordy luridae ，

Glossinidae ，Calliphoridae ，Tachinidae and Oestrida 日， The Muscidae are divided into into three subfamilies; Anthomyiinae ，Fanniinae and Muscinae ， The genera Fucellia ，

Pegomyia togeth 巴r with Hylemyia ， Anthomyia are includ 巴d in the subfamily Anthomyiinaε ，and the genera OJ うわ lra ，Liste ， 1tfusc 仰 a and Graphomyia are all placed in the subfamily MuscinaεThe Calliphorids and remain

respectively respectively in a subfamily rank 固

Hεrting's classification

Fam ， Muscidae Subfam. Sarcophaginae Subfam. Subfam. Anthomyiinae Subfam. Rhinophorinae Subfam. Subfam. Fanninae Fam. Tachinidae Subfam. Subfam. Muscin 日記 Subfam. Echinomylinae Fam ， Cordyluridae Subfam. Dexiinae Fam. Glossinidae Subfam. Phasiinae

Fam ， Calliphorida 己 Fam ， Oestridae Subfam. Subfam. Calliphorinae

(1960) divided th 色 into two subfam i1i es ，and continu 己d to treat treat the Ca l1i phorids and Sarcophagids resp 巴 as a i11 the Calliphoridae 。

A system adoI 比吋 in A 0/ the Dil うtera 0/ America North Mexico American Dipterists (1965) is as follows: 一

Section Section Calyptratae Fam i! y Calliphorida 日

Sup 日rfamily Muscoidea Subfamily Rhiniinae Family Family Anthomyiidae Subfamily Chrysomyinae Subfamily Subfamily Scatophaginae Subfamily Calliphorinae Subfamily Subfamily Fucelliinae Subfamily Polleniina 己

Subfamily Subfamily Anthomyiinae F 旦mily Sarcophagidae Family Family J¥在 uscida 巴 Subfamily Miltogramminae Subfamily Subfamily Coenosiinae Subfamily Sarcophaginae Subfamily Subfamily Li spinae Family Tachinidae

Subfamily Subfamily Li mnophorinae Subfamily Rhinophorina 官 Subfamily Subfamily Mydaeinae (Melanophorinae) Subfamily Subfamily Fanniina 巴 Subfamily Phasiinae Subfamily Subfamily Phaoniinae Snbfamily Proseninae Subfamily Subfamily Muscinae Subfamily Tachininae Subfamily Subfamily Stomoxyinae (Larva 巴vorinae) Superfamily Superfamily Oesteroidea Subfamily Dexiinae

Hennig (1 965) recognized nine subfamilies in his Muscid 配 (s. and maintained that the Fanniinae have a sister-group relation to his family Muscidae.

Hennig's classification

Familie Familie Muscidae Unterfamille Phaoniinae Unterfamilie Unterfamilie Fanniinae Unterfamilie Mydaeina 日 Unterfamilie Unterfamilie Achanthipterinae Unterfamilie Li mnophorinae Unterfamilie Unterfamilie Muscinae Unterfam il1 e Coe 日osiin

Unt 巴rfamilie Cryptoneurinae Familie Hippoboscidae Unterfamilie Unterfamilie Stomoxydinae Fam i1i e Nycteribiidae Familie Familie Glossinidae Familie Streblidae

In In his monograph Fan (1 965) recognizcd four superfamilies in the Calyptratae. The superfamily Muscoidea are divided into three families. The genera Fannia and Lisl りe are each treated as a subfamily in the family Muscidae. The genera 01 りhyra ，

Muscina ，and Gra ρhomyia ar 巴 contained in the subfamily Phaoniinae of the Muscidae. The Calliphorids and Sarcophagids are trcated respectivdy as a family in the

sup 巴rfam i1 y Calliphoroidea.

Fan's Fan's c1 assification

lyptratae Ca lyptratae Superfam. Calliphoroidea Superfam. Superfam. Muscoidea Fam. Calliphoridae Fam. Fam. Anthomyiidae Subfam. Polleniinae Subfam. Subfam. Fucelliinae Subfam. Calliphorinae Subfam. Subfam. An thomyiinae Subfam. Chrysomyiinae Fam. Fam. Muscidae Subfam. Phormiinae Subfam. Subfam. Fanniinae Subfam. Rhiniinae Subfam. Subfam. Li spinae Fam. Sarcophagidae Subfam. Subfam. Co enosiinae Subfam. Sarcophaginae Subfam. Subfam. Phaoniinae Subfam. Agriinae Subfam. Subfam. Muscinae Subfam. Metopiinae Subfam. Subfam. Stomoxydinae

Relevance of Characters to the Relationships of the Calyptrate Taxa Studied

(a) (a) The male accessory glands

The male accessory glands are wanting in the families Scatophagidae and Muscidae here examined ， while they are fully developed in the Calliphoridae ， SarcQphagidae ，Phasiidae ， Dexiidae and Tachinidae (s. st r.) (Hori ， 1960). In the Anthomyiidae they are wanting in the genera Othyra and Pegomyia ， but they are developed developed in the genera Fannia and Anthomyia. Thus ，on the basis of presence or absence absence of the male accessory glands ， the Calyptratae are separable into two main groups. groups. The absenc 巴 of th 巴 male acc 己ssory glands and the sim i1 arity of other internal male organs in addition to the kn~wn facts ab~ut the terminalia and other adu 1t characters characters link the genus 0) 幼yra with the Muscidae as assumed by some recent Di pterists as Crampton (1 944) ，Hennig (1 954-64) ，Huckett (1 965). The shape of the male accessory glands of the calyptrate muscoid Diptera also makes them roughly divisible into three groups; (a) spherical or ellipsoidal which occur occur in Fannia ，Anthomyia ，Isomyia ，Echinomyia ，Chrysosoma ，Succingulum and Ectothasia ， (b) banana-shaped which are observed in Hydrothoria ，Metotia ，Prosena and nine genera of the Calliphoridae ， and (c) co i1 cd which are peculiar to the Sarcophagidae Sarcophagidae (Hori ， 1960). 230 230 Katsushige HORI

(b) (b) Shape of the testes

The testes of the adult Calyptrate Dip 生era grea t1 y vary in shape with the advance of the age after the emergence. This intraspecific age variation is caused mainly by the discharge of the spermatozoa. Consequen t1 y the shape of the testes in a newly emerged fly has taxonomic significance. The shapes of the testes of the Calyptrate Calyptrate Diptera are ，from this point of view ， roughly divisible into three groups: (a) (a) oval ， (b) spongecucumber-shaped ， (c) lamp-shaped (Hori ， 1960). The morphological evidence evidence of the testes indicates that the genus Lispe resembles those of Scatophagidae rather rather than those of the other Anthomyiidae or Muscidae ，and Eciophasia also appears a to to be related to the Sarcophagidae rather than to the rest of the Tachinidae (s. lat.).

(c) Colour of the testes

The colours of the testes of the adult flies just emerged are divided into three groups: groups: (a) reddish orange ， (b) brown or fuscous ， (c) pale greenish yellow (Hori ， 1960). 1960). In the Tachinidae (s. str.) examined ， the testes are pale greenish yellow in colour ， while only in the genera Ect o. ρ hasia and Succingulum they are reddish orange ， resembling resembling those of the Sarcophagidae ，Calliphoridae ，and many others.

(d) (d) The fat-body

The fat-body which densely invests the adult t四 tes of the Calyptrate Dipt 疋ra is observed observed in all the members of the Calliphoridae and also in some Tachinids (Hori ， 1960). 1960).

(e) (e) The ejaculatory apodeme

The ejaculatory apodeme is a chitinous sclerite of the ejaculatory sac. In the three species species of the genus Fannia ，no trace of this is to be found ，which indicates their probable probable primitive condition. This morphological evidence indicates that the genus Fannia has a very primitive nature considerably different from most of the Antho- myiidae examined. The shape of the apodeme was roughly classified into the following two types: (a) rod-shaped ，and (b) fan-shaped ， the latter being advanc 巴d (Hori ， 1960).

(f) (f) Length of the ejaculatory duct There can be little doubt that a straight tube connecting the vas deferens with its its aedeagus is the ejaculatory duct of the primitive insects ，a longer one meaning a later later developmen t. To compare the length of the ejaculatory ducts of the Calyptratae ，

the the ejaculatory duct indices of 83 sp 巴cies are measured (Hori ， 1960). The Sc ato 司 phagidae ，Anthomyiidae ， Calliphoridae and Tachinidae examined all belong to the first first group in which the indices are below 50. In the Muscidae ， the genera Musca ， Graphomyia ， Pyrellia ， DasYPhora ， Orthellia ， Myiospila and Muscina of the subfamily subfamily Muscinae all belong to the second group in which the indices are .generally Phylogeny Phylogeny of Calyttratae 231

ovεr 50 ， While the values of the genera Stomoxys and Siphona which belong to the subfamily subfamily Stomoxydinae remain below 50.

In In the S司rcophagidae genera Leu CO' myia ，Bl α esoxitha and several members

of of the genus SarcoPhaga (s. 在t.) which have three dorso 四 centrals all belong to the

first first group ，and the othε 1' s of this genus belong to the second group 喧

(g) (g) TO l" sion ()f th8 eja むulato :r y duet

The modes of the torsion of th 己 ejaculatory duct round the hind-gut are of two typ 色s; counter c1 ockwise torsion ，and clockwis 色torsion. Of 8.3 species examine c1， only only Lisj うe belongs to the fl 1' st type (Hori ， 1960). This evidence as well 乳s the lmown facts facts of other internal organs indicates that Lispe is considerably different from those of of the typical Anthomyiid or' Mu.scid Diptera. According to Milani and Rivosechi (1 957) the direction of the torsion in lV fusca determined by a single g色ne. Therefore the taxonomic value of this charactεr must avvait furthεr study.

(h) (h) The l' ectal

The arrangement of 1: h日 rectal papillae of the ma1 巴s was divided into three types: (a) (a) bilateral type ， (b) cruciatεtype A ，and (c) cru cI ate type B. (Hori ， 1962a).

( i) Positum of the rectal valve

The rectal valve indices く 100; 1 -length of the distal intestine ，L 四 length

between the commencemεnt of the hind 回 gut and the rectal pouch) of 82 adult flies are are measur 吋 (Hori ， 1962a). The values inclicate the relative position of the rectal

valve valve from th 巴 commencemεnt of the hind 叩 gu t. speaking the valuεs among

a single genus or allied groups are nem 勾 ly constan t. 1n most of the Anthomyiidaεthe values values are 50 to 40 ，whilεin うe the value is ovεr 90 ， differing c1 early from the

rest rest of the same family. It deal 縄 that is remotely related to other Anthomyiids examine c1.

( j) Co iI ing degree of the lnoximal l. ntestine

The coilingεs of th 色 intestine of the male adult flies are obsεrved (Hori ，196:2a). The are smaHer in th 日Scatophagicla 色， Anthomyiidae ， Sarco- phagi c1 ae an c1 Tachinida 日， and larger in the Muscidae and Ca l1i phoridae. 1n the resp 巴ctive subfamiliεS of the Anthomyiidae ，Muscidaε ，Ca l1 iphoridae and Sarcophagidae ， the the degrees incr 巴乳 se proportionately from more primitive to more aclvanced (see Table

1). 1). Th 巴 8ma11 valu 巴s of th 日 d巴grees of the Tachinidae are probably the result of the

parasitic parasitic habit in th 巴ir larval stages. It is notice 日lble that the values of the Sarco 回 phaginae examined are to but little variatio l1， in contrast to those of the others.

This may indicate that the food habits of the Sarcophaginae 乳re nearly the samε 。 232 232 Katsushige HORI

(k) (k) Lengt lI of the m :i. d 同 gut

The double coils of the proximal intestine are in most cases mor 巴 or less irregula r. Consequently Consequently the measurement of the indices ， besides the coiling degree of

that that part ， is necessary ⑩ The mid 司 gut as has been expectεd ， are inclinεd to increase increase i l1 many cases parallel with the values of the degr 田 (Hori ， 1962a).

0) A r:r angement of the Malpighian tubules

The arrangement taken by the anterior pair of th 邑 tubules was grouped into the following following three main types: (a) Y-shaped ， (b) M-shaped ，and (c) fused (Hori ， 1962 b). b). The arrangements of the tul コules of the Phasiinae and Dεxiinae appear transition 乱l from the Tachinidae stL) to the ム Sarcophagidae and other families. ln ln the Tachinidae examined ， the Malpighian tubules are generally short ， but their diameters diameters are usually very large. This is regarded as a compensatory of the tubulεs for the 10s8 of the of the tubulεs. These three kinds of arrangement ， including including the transitional one ，may well have developed in the following direction:

→ 一一歩 → fU8 色d

The tubules of う fused into a loop must definitely be regarded as one rare rare noticeable case of Judging from the morphological characters of the the tubules as well as other εxt 吃rnal and internal Graphomyia appears to be considerably considerably different from the rest of the NIuscidae.

(m) Number and arrangement of the sper ・matheeae

1n 1n the majority of the Calyptrate Diptera examined ， the spermatheca 巴 are usually three three in number ， but in cases such as three species of the Stomoxydinae

and the lesser house 圃 fly ， are tvvo 1961). At this time ，however ， it is difficult difficult to arrive at any firm conclusion regarding their p1' imitive number in the

v 巴1' y the number two appears to be the most probab1e primitive primitive of these organs in the Calyptrate Di ptera.

This fact as well as other external characters such as thεmouth 四 parts sugg 巴sts that that the as 干vell as Glossinidae occupy an interesting position in the

Calyptratae Calyptratae genealogy. The spermathecae 011 the common oviduct a町 bilaterally arranged arranged in three ways: 2:0 or 0:2; 2:1 or 1:1:1 (Hori ，1961).

(n) (n) The female accessory

From the morphological evidences the female accessory glands are divided into three three groups: (a) oval or elongate oval ， (b) banana-shaped ，and (c) c1 avate. Among th 日se three types the first mεntioned type 18 the most primitive ，and the second is is the most while the last type occupies a position interm 己diate between these two 1961). Relations among these th 1' εe types are considered PhylogM 里y of Caly ρ'tratae 233

as as follows: ー

Oval orεlongate oval → banana -sha p巴d → clavate of of the female accessory glands together with the the shape 01' t11e testes ， t11e and Sarcophagidae appear to have S0111e affinity. affinity.

(0) (0) The uterova g;:i m: !l tube

From the morphologicalεγidences the uterovaginal tube is divisible into three groups: groups: (a) a simple-shaped ， a straight tube with a large incubatory pouch ，and (c) coiled when matured. these three groups the first group is ， without without dou i.比， the most primitiγε ，and tlle sξcond and third groups are deriv 出l independen tI y from the first 1961).

The arista

In In the higher Di ptera ， the sβgments clistal to the tllird antenna1 segment ar 巴 greatly greatly reduced and appear to he a slencler appεndage of the antenna. They

are are called arista. The diffel でntiation of thεarista from the fourth and its succeeding

terminal terminal antennal segments is first r日 in the Nematocεran Diptera by

Robineau 四 Desvoidy In the Calyptrate Diptera ，therefore ，a bare or sparsely haired haired arista is ，事 rithout doubt ， in the primitive condition. The significance of the arista arista in the taxonomy of thεDiptera was recognized by Linnaeus (1 758). 1n I: he Calyptratae it Is said that the arista is we l1 developed 3nd affords taxonomically important important specific characteristics. In In the Scatophagidae which are supposed to be the most primitive ， the arista is bare ， representing their least mod i:f ied condition. In the Anthomyiidae it is bare in 0ρ hyra and Pannia ， short p1umωe in and Lispe ，plumose in I-l ydroPhoria. In In the Muscidae it Is plumose in jVf uscina ，Morellia ，Qrthellia ，Dasyphora ，Pyrellia ， β 1usca and Grathomyia a11 of vvhich aγe here includεd in the subfamily Muscinae;

while while it is s合miplumose in and which belong to the subfamily

Stomoxydinae ，inciicating some resemblan 町 to that of Glossina. In the Stomoxydinae ，

howevεr ， this condition of the a 1' ista is not a gmup character ，for ，even within th 田 e

two gεl1 era ， several exotic forms in c1 uding Stomox)'s chinensis ，S 必hona stimulans and some others have a more or less plumose arista.

111 111 the Ca lI ipho 1' ida 己 it is short plumose in Stomorhina ， while a11 other members

have a de 己ply o-n~.".v~合 arista. 1n the Sarcophagidae it is ba 1' e in Meto] う pubescent in in Leucomyia ， while it is well plumose in aH the members of the genera Blaesoxitha

and Sarcothaga 1a 1:.) examined. ln the Phasiidaεand T 旦chinidae examined it is

bare ，wh i1 e the Dexiids examined bεa 1' short pubesc 色nce. From the above mentioned ， it is that the arista of the CaJyptratae 234 Kat 弓ushige HORI presents presents of hairiness to their systεmatic and the filiform filiform type of arista indicates the basic or primitive type ， haired ones bεing a later The hairiness of the arista must ， generally speaking ，have

fro ロ1 the to the the tvvo 日equenCeS:

b メ，， ¥;...-U 1...H.ド山 liV':'C → 10 昭 plumose are are -;. nl1h ，po'N"" く

冶 short plumose ザ plumose → (many Muscids)

(q) (q) Distal bend of the fou :r th Iongitudinal vein

It It is aw 巴11 known fact that theγenation of the veins of inseds is so stable within within a but between different 巴s or groups it is considerably so variable that that it is very useful for purposes of classification ， In the Calyptratae the fourth

vein 官1→2) is of considerable since it Is very long and stout

and anchors the postεrior half of the to its 司nterior part a cross 四 veinη 〕 and a1so connects it with the end of costa ， cつnsider thεfourth vein to be a fusion of 111- 1 and

')']1 1 21 'v¥-rh i1 er 巴ce 立 Lower (1 951) beliεves inεIVluscoidea calyptratae such as

and this is not 11'1 1 as understood ， but an union of r 5 and m r， bending upwards and the latter of thεformer as a stub vein.

In In the with the vein 1' 4 十軒

The almost similar course pattern of the fourth v巴in is a1so seen in a11 the members of of the here examined ， The Muscinae wεre former 匂 S巴parated

fr 均 om the others of the Muscida 色 on the basis of the bend of this vein. In the

however ，th 巴 fouth vein is bent ant 巴riorly tov ，.氾 rds the third vein before before in the genera Muscina and curved in thεgenera

bent bent in the genus Musc 仏 and bent bent up in a11 the members of this family. ln ln the Sarcophagidae and Tachinidaεstr.) examined a1so thεfOUI 廿1 vein is to to the third vein and corresponding

Th 巴se facts may bεtakε11 to indicate that the fourth vein of the

PI ・imitive Di ptera is a little diverted from the primitive type which runs almost straight to the margin as shown in nearly a11 the extant extant Acalyptrate and tha t. the curved on 巴s of the fourth vein are found in the the evolved flies. This distal bend of the fourth the directing forwards to the ante fI or

costal costal margin ，九ア hich is (1 803) to separate the Anthomyiids Calyttratae 0/ Calyttratae 235

fW l11 the Ot! lfol ・ Muscid flies ， frequently occured in the higher Calyptrate Diptera as

abo"'/e 明 mentioned. 1 propos 巴 to tご1" 111 th .i s phenomenon he 日“cos f: αliz αuon" ，which is pf)oab1y pf)oab1y eff ョctive in increasing the strength of the wing indispensable for the quick and strong vibration of the wing in the “higher ， or larger ，flies 勺 with due result of ::o th ::o degeneration of the hind part of lI1 e wing and a1so with due change of the general form of it. From above it is clcar th.a t. the dis l. al bend of the fourth vein of the Calyptratae Calyptratae is an evolutionary trend and increases with the advance of their phylogenetic phylogenetic position as shown above. This view has ，however ，met opposition in van

Emden (1 958). According to him this bend of the fourth vein ，h O¥羽 八、 J be 巴n known in so mar 日ly Ul 日lr 己1at 巴d ior 口.1 8 tha1 its u仁Ll 8巴 as a group cha n:乳 lct 巴下 should be abanc 吋仁don 色cl. The inv 己stigation of this possibility must be a subj 巴ct of :f uture study.

(r) Size of the squamae

The squamae or calypters キ are f1 ap. ，like membranou 詰 region at the base of the wing ， the upper bεing the a1ar or upper sq u乱111a which is attach 巴d to the wing ，and the the 10wer is the thoracic or 10we 1' squama which is attachecl to the thoraxo The uppεr squama is an apparently moclified of the basal m 巴mbrane of the wing ，¥vhile the thoracic thoracic squama is op 巴11 to considerable doubt as to its primitive status or origin ， and is inconspicuous in most Diptera including the Acalyptrata 丸 but in the higher

Diptera Diptera it is usually well d巴velopecl and often conce 乱ls the ha1t 巴re. On the basis of this this they a1'巴 call 己d the Calypt 1' ata 色 or Calypterae.

Th 己 Cal かypt 色白rata 巴久 accoωr I七寸司dピE一1i加 ng 白o Os 叫te 白凹n' 冶.

D~おsvoωidy as t11ε firs 凶t，fr 伺刀om はth日己 la 釦r令ge 呂別 12 犯ε Oぱf thelr sq l1 ama 己勾 ぱofh 11 一 おS 1凶廿1in 巴 g釦r‘oup 呂 of

Myοdair 晶 es in 1830. This divlsion of 己1' a， howeve 1'， was not equivalent to to the Calyptrata ::o of late 1' authors ， because it did not include the Anthomyiidae. 1n othe 1' words th 巴 Calyptrata 己 here in qu 己stion are a group of “hi. gher tlies" characterized by the gOJd d~v eIopm~nt of the thoracic sq l1 ama which used to be considerεd to be taxonomically taxonomically of no importance by some Dipterirts.

In In order to indicate the degree of tb 己 development of the thoracic squama into the the alar squama ， the mean value squamal index ，which is the Iength of the thoracic sqama divided by that of the alar squama ，was measured for each speciεs as shown in in Table 1. The squamal indices are 002 in the Scatophagidaeo 1n th 巴 Anthomyiidae the the indices are 0.8 in Pegomyia ， Othyra ， Anthomyia ， Hydro) 幼 oria ， O. 9 in F annia ， whereas in Lis) うe they are 1.5. 1n the Muscidae they are 1.2 in Siphona ， 1.5 in Stomoxys (Stomoxydinae) ， whereas in the Muscinae they are 1.2 in IVl uscina ，

Myospila ， 108 in !v Iusca ， 200 in liA orellia ， Orthellia ， DasYPhora ， Pyrellia and

Gra] うhomyia.

111 111 the Calliphoridae th 巴y are 1.2 in Isomyia and Stomorhina ， Protoj うhorrnia ，

斗 The terms tegulae ，alulae ，etco ， are a!s o used by earlier Dipteristso 236 236 Katsushig 官 HORI

1， 5 in 1 号 B in Lucilia. Blaesoxitha Blaesoxitha and う la t.) here examined. In In the Tachinidae (s. la t.) are 2.0 in Echinomya ， 2.2 in and Sturmia.

Although the function of the squ 証maεof the is not studiεd and remains subjected to considerable it is that the thoracic squama

1S 1S little 温 or in the primitive mεmbers even within the Calyptratae. From the above mentioned it is that the index increases gradually from the primitive to the advanced accoring to the Ir司 systematic

(s) Number of the b: dst]es

The bristles are t巴rmed the brlst1es or simpIy simpIy two ， as seen in thεin the families Scatophagidae ， and Tachinidae here examined.

In In the Sarcophagida 巴， there are two in lvleto iJ 向。 but in Leucomvia there are are one or two sma ll. bristles i11 addition to the usual two strong ones ，and there there are constantly two small bristle 呂 besides the usual two strong ones in Sarco- thagao thagao Thus the number of the bristles of the ranges ， within within narrow limits ，from the llsual strong two (0 in accordance vvith their their systematic position sεen from internal anatomy. This evolutional trend will be supported supported by the fact that some such as the Dn コ have two notopleural notopleural bristles.

(t) 。f the bristKes The bristles simp1y simp1y

h巴re examined.

sorne sorne short hairs arεpresent in the gener 忍 the the Calliphoridae the groups such as and Stomorhina have

severaI severaI short bristles on their hypopleura ， vvhile the gεn 巴ra and Lucil ia ha ve a row of well

there there are sev 巴ra1 short hairs on the hypopleura

of of the genera Metotia ， and Blaesoxi iJ h仏むld the hvpopleura of all the members of the genus Sarco ρ(s. laL) hereεxamined have a row of bris t1巴 s. All the the Tachinid flies here studied have a row of wellεd hypopleural bristles.

On the basis of presence or absence of th 巴 bristles ， it was customary to to divide the into two familiεs ， or and

Tachinida 日 bv older Diote .r ists sincεOsten - SackefL to Shann( コn( the Phylogelly Phylogelly 0/ Caly ρtratae 23 '1 prcs':nce prcs':nce or absence of the hypopleur 叫 bristles is mainly correlated with their larval

Iood Iood habits ，旦ncl the groups lacking the hypopleural bristles are the most primitive. But the developmental status of the hypopleural bristles of the Calyptratae is not

always clear cut among the different groups of the Calyptr 命 atae ，and many lntermediate conditions ， as above mentioned ， are frequen t. These morphological evid 己nces suggest that that the hypopleuron of the ancestral calyptrate prototype must have been Lare or poorly poorly haired. From the above menticned it is clear that theεvolutionary trends of the the hypopleurals are the change from a bare or weakly haired conditicn through a haired haired to a bristling with the advance of their phylogenetic position. Hence ， their taxonomic values are of great importance not only on the family level but a1so on the genus level ， or som 巴times on the species leve l.

(u) (u) Head widths of the adult flies

For the purpose of estimating the body sia of the adult flies the head widths of the the l11 ales of 83 species were ，on an average ，measured (Table 1). The head wiclths are 2.2-2.3mm in the Scatophagidae. 1n the Anthomyiiclae they are are 1. 6-2.0mm in Fannia ， 1. 7mm in Anthornyia ， 1. 81mll in Pegomyia ， 2.0mm in Liste ，

2.1111111 2.1111111 in Hydrothoria ， 2.1 ・2.4mm in Ophyra.

111 111 the Musciclae th 巴y are 1. I!1l1 11 in Sithona ， 2. 2m 日1 in Stmnoxys (Stomoxyclinae) ，

1. 1. 61l1 m in Pyrellia. 1. 8-3. 3mm i l1 Musca ， 2. 4nnll in 11 ゆめゆila ，2.7-3.4 凹 in Muscina ， 2.8mm 2.8mm in Morellia ，2.9111m in Orthellia ，3.0111111 In Dasythora ，3.9nllll in Graphomyia.

In In the Calliphoridae they are 2.4mm in StOJnorhina ， 2.51 旧日 1 in IS O'I nyia ，2.2nnll in

Melinda ，3 同 3111111 in Tricerato ρ'yga ，3.9 “ 4.2mm in Call i. ρhora ， 2. 9llnll in Hem i. ρ'Y1'ellia ，

2.4 閑 3.6nnll in Lucilia.

1n 1n the Sarcophagiclae tbey are 2.5 日1111 in Metotia ，2.6 11l m in Leucomyia ， 2.3-2.6mm

in in Blaesoxitha ，2.0 回 4.311nll in Sar ・cothaga (s. la t.). In In the Tachiniclae (s. la t.)， they are 2.7-3 .1unll in Succingulum ancl Prosena

(D 巴xiidae) ，3.2nllll in Ecto) 叫 αsia (Phasiidae) ，3 ，5 咽 4.8mm in the Tachiniclae (s. st r.). From the above it is noticeable that the head widths of Scoteuma here examined are are much larger than those of most of the lower Calyptratae. But l11 0st of the Scato- phagidae ，especially those of the primitive of this group ， according to Sack (1 937) ， are are sma11 slender flies about 3-6 1l1Ul in bocly length. The head width of these primitive

Scatophagids ，therefore ，must be much smaller than those of Scoteuma here l11 easured.

Fro l11 the ab:we it may be said that the head widths increase in size in an ascending series ，with some exceptions ，fro111 the primitive forms to the evolv 巴d， exhibiting a considerable considerable variation of more 工han twice the width of th 巴 smallest. This evolutionary trend in the Calyptratae is also backed up by eviclence from the

Acalyptratae ，where a small or slender body is th 巴 most com111on condition. 238 238 Kat 巴ushige HORI

Table L Characteristics of th 己 adult fiies of the Calyptratae

Male Fan1Uy &:

S開 cl 崎 Subfamily

Mu~ddae Gratlw 岬， y岬閉acuiata Musci 回 e Mus ωdω 地 S附 G 出円削

M 問問 sorbens

Mus 叩 hervei

M 閑""印刷 exilrl 由絡 PyreWa .cadaverin2

Dasyphora cya 捌 colo 'i'

Orlhe lz. 叫印¢叩!四

MoreU 国幻附'PI 鉛""糊昭

Myio 塾、 la 醐 alitabimda M uscina stabulans J，.Wl~SC担 :l aw;wJtifro ?i s

M 問cinz 'Jう :z lnuon 掛包

StO mG XYS 回 1A itra 畑

Sip !w n:;; exi r. n日 。占 hyra OPhy r> !m ，，，， stoma

Ophyra cha! 叩， iJ otste 'f p 昭市nyra VWf Jm

Sωβ ew: 四'l ste 1'l ωrarh 棚

Scota 抑 1a 脚立物s L以i5桝 μo

Fa 酎加"悶nF悶n削σ sc 悶庁悶材芳吋l陶αar 円13

Fσ" 柑 a 却5p. Fannfa ca師部~laris

f1 "tlw 恥J問 iU ocata ;ljl;1711lj H ydro)horia ruralis ;ljl;1711lj lsomyia se 田 'mera 了 dev e1 0ped St ()J'P 叩>rhin !t obsoleta

Pro!oph 却 r刷 ia ierraa n

P Jw nnia re;;in :'A

Chrysm 珂ya mo 仰向，phala

Chrys 四nya PI 珂 ms

M elinda pusill l'l

Tr 国 eYa 如'pyga 印 llip 恥 roides 仇 lliplwra !ata

Calli 主hora fJ rah:u 柏町

H 即 'ti pyrellia ligltrri~ lI s Lu dlia 的ifoniVG 〆6 lA<ニョ lia cu) 町四

Ltd 抑制YI 岬 ta

L1.dlia L1.dlia ill 附 tris

Lu c:i l凶 caeSG γ

Lu 刃lia am 戸t. llac 回

心t;i lia PO ゆ，hyrin~ 1../ 口 lia μ'jnansis Metopia 1即""田 :p hala 一一 抑制戸αnZ11 即

Blaesozipha Blaesozipha laticor- 捌 S Blaesox ψ 'ha japonensis

Sarcophaga Sarcophaga 11 昭 la 附 ra Sarcoph3g3 Sarcoph3g3 Sp.

SarcoPhaga SarcoPhaga uga 刑 sk 勾z Thyiogeny Thyiogeny 0/ eai y. ρtratae 239

In 陪'rnal characters

Ferr 国Je Ex 怯rnal ch 田 acte 四

MalpIghian MalpIghian tubules Re productive 世田国 Anterior Anterior Arrangement Arrangement 白 Ilour Spermathecae tubules tubules Utero .. H 田 d Agclcaensdsos Agclcaensdsos ry vtaug 悼inal A 口S祖 3E E width 叫国 3 Atunbteurlieosri|ptousbteurlieos Atunbteurlieosri|ptousbteurlieos r Atnubteurlieosr |1Ptousbte ri or Min 回 dexllFdeugsriene g iii Ei

ba 田 na .. gecntulry ved f国 ed 3 fuscous fu 配 ous 2.7 7.0 3 1:2 shaped simple plumo 揖 2.0 2 short hairs 3.9 .. M .. 1 yellow yellow 2.0 11.2 s 1:2 c1 avate shacruprlvy ed 1. 8 2 国 re 2.4 .haped .haped " " " 1 " " 2.3 8.5 s 1:2 " " " " 1. 8 2 " 1. 8 " 1 " " 3.1 13.8 3 1:2 " " " " 1. 8 2 " 2.6 " 1 " " 3.0 7.0 3 1:2 " " " " 1.8 2 " 3.3 " 2 " " 2.8 6.8 3 1:2 " " " genctulry vd 2.0 2 " 1.6 " 2 " " 3.0 5二2 3 1:2 " " " " 2.0 2 short hairs 3.0 " 2 " " 3.5 6.9 3 1:2 " " " " 2.0 2 同 re 2.9 " 1-2 " " 3.0 10.2 3 1:2 " " " " 2.0 2 " 2.8 " 1 " " 1. 8 5 .4 " bowed 1. 2 2 " 2.4 " 1-2 ." " 2.5 16.7 3 1:2 cU¥ vate simple " " 1. 2 2 " 3.0 " 1-2 " " 2.8 15.9 3 1:2 " " " " 1. 2 2 " 2.7 " 1-2 " " 3.9 12.2 " " 1.2 2 " 3.4 " " 2.0 11.1 2 0:2 c1 ava 胞 simple 日 mi.plumosc " 1. 5 2 " 2.2 " " 2.1 11. 3 2 0:2 " " " " 1. 2 2 " 1. 1 " s 1. 5 4.7 bare straight 0.8 2 " 2.3 " 3 1. 5 5.3 3 1:2 davate simple " " 0.8 2 " 2.4 s 1. 4 5.9 3 1:2 0.8 2 11 2.1 " " " short " " " 2 1. 6 4.3 plumose " 0.8 2 " 1. 8 v l Ml M l M l151 321 3 1 1 ぺ。叫 simple 同時 | " 10212| " 2 I 1. 6 I 3.6 I 3 I 1:2 I " 0.2 I 2 " 2.3 1 2 .4 4.8 3 1:2 short plumo&e 1. 5 2 2.0 " " " 岡田抽・" " " " 11 11 1 white 1. 5 4.1 s 1:2 bare 11 0.9 2 11 1.9 " shaped " 11 1 1. 7 3.9 0.9 2 11 1.6 " " 岡田 na .. " " 1 1. 5 4.4 2 0:2 宇田ple 0.9 2 11 2.0 " " " shaped short " " 11 11 2 yellow 1. 7 5.6 3 1:2 11 0.8 2 1. 7 " pplulummoos e田 " " 2 1. 8 6.0 " 0.8 2 2.1 " " " banana. " " 11 11 11 1. 6 5.8 3 1:2 simple 11 1. 2 2 畠 s shaped gently cu t' vcd maU brlntles 2.5 " 3 11 11 2.4 5.5 3 1:2 11 11 short plumoHe 11 1. 2 2 11 2.4 ヲー IT ご一一 11 11 3 11 11 2.5 7.7 3 1:2 clavate plumose C urIvTed - 1. 2 2 3.3 I " 三高 石 developed 11 11 11 11 11 s 11 2.6 8.5 3 1:2 11 1. 2 2 11 3.2 11 3 11 11 3.2 9.8 3 1:2 11 11 " " 1. 5 2 " 4.5 11 11 3 11 I 3.4 8.5 3 1:2 11 " 11 11 1. 5 2 11 4.5 11 1 11 11 1. 0 7.7 " 11 1. 5 2 11 2.2 11 11 s 11 11 3.1 8.3 3 1:2 clavate simpIe 11 11 1. 5 2 11 3.3

11 11 3 11 11 3.0 7.1 s 1:2 11 11 11 11 1. 5 2 11 4.2 11 11 s 11 11 3.0 9.3 3 1:2 11 11 " " 1. 5 2 11 3.9 11 1 11 " 3.0 8.1 3 1:2 11 " 11 11 1. 5 2 11 2.9 11 11 11 1. 8 2 11 2.4 .. M .. s yellow yellow 2.2 3 1:2 clavate simple 11 1. 8 2 11 2.6 shaped shaped 6.9 " " 3 11 " 2.8 6.8 3 1:2 11 〆f " 11 1. 8 2 11 2.9 11 3 11 11 2.8 9.6 3 1:2 " 11 " 11 1. 8 2 11 3.2 11 11 s 11 " 3.2 10.1 3 1:2 11 11 11 " 1. 8 2 11 3.6 11 3 " 11 3.6 9.0 3 1:2 11 11 " 11 1. 8 2 11 3.3 11 3 11 11 2.8 8.0 3 1:2 " 11 Zシ 11 " 3.3 11 11 3 " 11 3.5 9.1 3 1:2 I 11 Aγ 11 11 1.81 2 11 3.4 一 bare I 2.0 2-4 町田 11 brisUes 2.5 f3r1 1. 81 7.91 3 11:1:1 oval in~~~~~ory ¥ pU 国 cent 1 1 2.0 4 " .2.6 4 グ 1 0.91 7.61 3 11:1:1 I plumo 時 I 2.0 " 2.6 1 0.9 1 10.4 1 3 11:1:1 I 2.0 4 2.3 well well ".. 1 1. 51 5.31 3 11:1:1 1 2.0 4 developed 3.4 3 1 2.0 1 6.6 一一 一 I 2.0 4 " 2.3 3 1 1. 21 8.6 一一 ー 1 2.0 4 " 2.5 240 240 Katsushige HORI

Table 1 (Continued)

Ma le Family Family & Reproductive Reproductive orga 田 Digestive organs Sp らci 田 Sublamily Test 田 Ejacula 柏， ry duct Mid -g ut R 田 tal Agclcaen 田dos ry (;1→ iffe 白ren 山ti atioOn 田 f) Ftaits-sbuoe dy valve Shape Shape ICo lour index 3Z2E|lgZz 盟

Sarcophoga Sarcophoga caudagalli " " " " " " " 55.2 2.0-2. 1. 2 Sar 四ophaga 町民to 60.0 2.0 1. 1 " " " " lH drf r " " " Sorcop 加 ga josephi " " " " " " " 53.2 2.5 1. 4 Sa γ回 Iphaga kagaensis " " " " " " " " 55.0 2.5 1. 1 Sor ω，'P 加 ga .i ezensis 54.7 2.5 1.2 " 。" " " " " " " So γω 'Phago boso !i s " " " " lOQg " " 50.0 2.0 1. 4 Sar四日， ~haga β例gri:師 " " " " " short " " 55.3 2.ι2.5 1. 6 Sa γ回 ，phaga septentrionalis " " " " " " " " 52.0 2.5 1. 5 Sorcop 加 ga similis " " " " " long " " 51.0 2.5 1. 5 Sor 印，'p 加'ga ka 附則ω回 is " " " " " short " " 62.0 2.0 1. 2 Sar 四β加 ga antilo μ， 11 11 11 " " 52.0 2.5 1.1 Sar ω ，phaga albiceps " " " " " " 11 11 11 long " 11 56.3 2.5 1.3 Sa 叩'op 加 g(l tsus 胸蜘 11 " " " 11 " " " 55.2 2.5 1. 2 Sorcophaga Sorcophaga b向 vicornis 。 11 " " 11 11 11 11 60.6 2.5 1. 5 Sarc ψ，加 ga misera 11 11 11 11 " " 11 11 11 55.3 2.5 1. 4 Sarl 印 'Ph.ga harp 口 11 11 11 " " 11 11 " 57.5 2.5 1. 8 Sarcop 加 ga kobayashii 11 11 " " " " " " 53.6 2.5 1. 3 Sar ω ，'phoga ho 加吋b 剛 sis " " " " " " 55.7 2.5 1. 3 Sar ω9， 加 ga crassI. μ lpis " " " M " 52.2 2.5 1. 4 schutzei schutzei " " " " " Sar.ωIp."'~ga " " " " " " " 11 52.0 2.5 1. 3 Sarcophaga 加'zawa 11 ， " 11 51. 0 2.5 1. 5 Sarcophaga Sarcophaga gravelyi " " " " " " " " " " 11 " 11 52.0 2.5 1. 5 Sar 印，'phaga 加 kusana 4 " " 11 " " " " " 50.0 2.5 1. 5 Ect， ゆ加s臼抑te>> sis Phasiidae ! " I short rod ..g haped Icruciale BI 97.0 11.0 ー1.51 1.7

Suai 切ぱ酬， tran sVit ta 加 n D 虹 iidae 一bhs 一Mpn-ha an2eaa nH-eEaEl 生lレ「号 h一A一m{a 一prFe 一-d --yel ，lr ow I 11 I /1 三五五-l1 -lbila 回 196.0 寸:0 11:2 Pros 即幅 siberita rfoadE1- ・eshhappe 四:1 " 1 %.0 1 1. 0 1 1.1 sha pt: d " developed I " 1 " Servill ，回 ja 胸 1/' 劇画 Tachtnidae ~n- _ _1 11 183.02.0 1 1. 0 I " I 1'1 developed 城間>m 鍬古 E ya udo 1 1.1 I 11 i .(1 I " 11 1 83.0 11.5- 2.01 1. 0 Chrys ，^ 凶剤師 aura 抱 " " " developed " 11 11 " 77.0 2.0 1. 7 St 師 W 自 s.n 回'riae pap i1 lary un ・ " " developed 11 11 %.0 2.0 0.7 一 " "

Synthetic Synthetic Consideration of the Phylogeny of the Calyptratae

Some internal and external characters of the Calyptratae examined are arranged in in descending or ascending successive orders of their morphological resemblance on the basis basis of the fact that the degree of morphological resemblance in the true sense which exists exists between different organisms indicates usually the closeness of their phylogenetic relation relation (Table 1). As it is clear from this Table ， the Japanese Calyptrate Diptera here treated were ， by the pr 白巴nce or absence of the accessory glands of the male reproductive system ， divided divided into two groups; (a) that which has the organs ，and (b) that which lacks them. them. The Scatophagidae and Muscidae are included in the group (a) ，and the Calliphoridae ， Sarcophagidae and Tachinidae are all included in the group(b). The An thomyiidae here studied are divided into two groups: (1) to which genera Fannia ， Plzylogeny Plzylogeny of Ca!Yl うtTatae 2，11

I l1 ternal characters

F 円 nalc Exterm¥l d 1il racters

Arbta

2 ，0 ii11 什2 ，1 3 ，3

2 ，0 i 4 I 2 ，8

3 2.0 4 2 ，3 り何のんヴ 1nd dod 77 2.0 4 : fノ 3 ，2 Qυ 2 ，0 4 3 ，3

2 ，3I 1004 2.0 4 3.5 2.8 2.8 4 3 ，0

2 ，0 2 ，0 4 2 ，5

2 ，0 4 2 回日

2.2 2.2 I 5.6 2 ，0 4 3 ，5

2 ，0 4 I N 2 ，9 1， 9 2 ，0 4 3 ，1 2.2 2.2 2.0 4 3 ，1

2 ，0 4 3 ，1

2 ，0I 6 ，7 I - ， 2 ，0 4 4 ，3

2 ，0 4 2 ，7 l!lI1111Jii::tUjl l!lI1111Jii::tUjl jlijijljljj 2 ，0 4 4 ，2

Anthomyia and Hydrol うん oria belong ， ancl (2) to vvhich genera Ophy Ta ancl Pe[!omyia belong. Thus the lack of the male accessory glancls is i11 the Calyptratae of of great taxonomic vaIue since it inclicates that the Scatophagiclae ancl Musciclae shoulcl shoulcl be consiclerecl the more primitive of the Calyptratae. The Scatophagiclae ， or Corcly Corcly luridaαon the basis of their retention of so many of the features of the archetype ， e.g. the well separated eyes in both sexes ， the weak thoracic squamae ， the

r巴tention of five ，sometimes six evident abdom .i nal segments ，one sternopleural bristle ，

etc. ，ar 巴 usually consiclered the most primitive of the Calyptratae ，and in the past ， therefore ， they were sometimes in c1 udecl in the Acalyptratae as a distinct family or a

subfamily subfamily rank by some olcler workers such as Schiner (1 862 旬 4) ， Coquillett (1901) ， Williston Williston (1 908) ， Comstock (1 924) ， Imms (1925) ， etc. Enderlein (1 936) raised the Scatophagidae ， including the Corclylurids ， to a new series Protomuscaria as a distinct

rank co-ordinate with the Acalyptratae or the C 乱lyptratae 1n the Schi; ;;o phora of 242 242 Katsushige HORl

Di ptera. Essig (1947) ， Shiraki (1 954) ，etc. ， adopted this system of Enderlein's. On the other hand ， it is a known fact that the Sc atophagidae have Anthomyiid- like like male genitalia together with some other primitive characters peculiar to the Calyptratae Calyptratae (Crampton ， 1944). Therefore ， recent specialists such as Hendel (1928) ， Crampton (1 942 ，1944) ，Hennig (1 94 1)， Colyer and Hammond (1 951) ，Roback (1 951) ， Oldroid Oldroid (1 954) ，van Emden (1 954) ， Ringdahl (1 954) ，Vockeroth (1 965) ，etc. ， regarded the the Scatophagidae as belonging to the Calyptratae and forming a link between the lower lower and higher Schizophora. Recently Roback (1 951) takes the position that the Scopeumatinae and the Anthomyiinae have almost without change the generalized pattern pattern of the ancestral Calyptrate muscoid postulated by himself. Van Emden (1 941) thought thought the Scatophagidae ， or Cordyluridae ， should be regarded as a mere subfamily

of of the Muscidae. Judging only from the absen 田 of the male accessory glands the Scatophagidae Scatophagidae (s. str.) here examined are internally similar to the typical Muscidae ， but but the shape of the testes of this Scatophagid Scopeuma here examined do resemble not not those of the Muscidae ， but those of the Sarcophagidae (Hori ， 1960). As many pf the the Scatophagids ，especially primitive smaller on 出 are not internally examined ， their

affinities affinities and systematic position can but be ， at pr 田 ent ，provisiona l. It It is a we l1 known fact that the Anthomyiidae have been considered by many

Dipterists Dipterists to be the m Cl st primitive of the Calyptratae ，and also to be c1 0sely related to to the Scatophagidae on one hand ，and to the お1uscidae on the other hand. Coquillett (1 90 1)， Williston (1 908) ，Comstock (1924) ， Enderlein (1 936) ，Crampton (1942 ，1944) ， Tokunaga (1 943) ， Essig (1 947) ，Zimin (1 951) ，Roback (1951) ， Shiraki (1954) ，Fan

(1 965) ， Huckett (1 965) ， etc. ， all r巴garded the Anthomyiids as a distinct family separable separable from the Muscidae. Some authors in c1 uding Girschner (1893) ， Stein (1 918) ， Seguy (1923) ，Imms (1 925) ，Hennig (1 941) ，etc. ，in c1 uded the Muscids in their family Anthomyiidae. Anthomyiidae. Senior-White (1924) ，Karl (1 928) ，Hendel (1928) ，Malloch (1934) ， Curran (1 934) ， Colyer and Hammond (1 951) ， Herting (1 957) ，etc. ，on the contrary ，

amalgamat 巴d the Anthomyiids into their family Muscidae. Roback (1951) believes that the the Anthomyiidae have div 巴rged very lit t1 e from the generalized adult and larval patterns patterns of the prototy Phylo [l eny 01 Calyttralae 243

fannines fannines to family rank 丘nd placecl them betvveen the Anthcmyiidae ancl Musciclae.

On the basis of their p己culiar a己deagi which have no c1 0se resemblance to the other Muscoid flies ， Chillcott (1 960) suggests that the ianniines ha c1 a sel ，arate evolution insteacl insteacl of having any relationship with the res t. Hennig (1 965) follovvecl suit and treated treated them as a subfamily of the Muscidae ancl thought that the Fanniinae are

monophyletic monophyletic ancl have doubtlessly a sister 情 group relaticn to the Musciclae la t.)。

It It may be remarked here that the m 旦le internal sexual organs of two species of the

Fannia here examined w 巴re distinctly Anthomyoicl in t11 巴ir characteristics (Hori ， 1960). 1960). If the evidence of the internal crgans is reliable ， the fannincs bear c1 0se relationship relationship to Anthomyia illocata which is consiclered the typical Anthomyiid ， not to the the Musciclae. The shape of the male accessory glan c1 s， the lack of the sperm pump sc1 erite ，etc. ， are also characteristic to this genus Fannia. Another point of interest is is that the larvaεof the fanniines show much more clear cut diffεrences from those of of the typical Muscids ， ancl may possibly be taken 1: 0 inclicate som 日 resemblance to those those of the Phoriclae. To clear the systematic status of the Anthomyiidae (s. st 1".，)， including the fanniines ， more accurate comparative studies on both the external and int 巴rnal morphologies morphologies of many mor 色 matεrials are to be waite c1. Crampton (1 944) considers the Anthomyiiclae Anthomyiiclae (s. str.) are the key group to solve the relationship among the diffεrent groups groups of the Calyptratae upon which many Calyptrate f呂milies converge. The present present author too believes that this view of Crampton's is very rεasonable. The genus Ophyra together with Pegomyl:a is classecl in the Anthomyiidae by many older workers. Karl (1 928) ，Ring c1 ahl (1 954) ，Hennig (1 941 ， 1962) ，Fan (1 965) ，

Huckett (1965) ，etc. ，placecl the genus 0ρ わJra in th 巴 subfamily Phaoniinae of the

Muscidae. Muscidae. Herting (1957) placed the genus Ophyra in the Muscinae by its Hydrotaea 崎 type type of ovipositor. Van Emden (1 943) thought that the genus Othyra together with the the g巴nus Hydrotaea is closely related 1: 0 his Limnol うhora-group of thεsubf 乳mily

Phaoniinae Phaoniinae ancl these two g巴nera seem to connεct the laUer group with the Fanniinae. But their female reprocluctive organs such as accessory glands 244 244 Kats l1 shige HORI

1960)0 1960)0 The other intεrnal of Liste such as the rectal valve a1so is anomalous and clearly too different from thε Muscids to warrant it in the

Muscidae ，and the shape of proboscis ， predatory et c" are more suggestive of those of of the Scatophagidae ， Thεauthor believ 田 that the differences ，mainly with respεct to the the male internal sexual organs and thεexternal characters are syst 邑matically so important important that th 己 g巴nus Liste should be excluded from the typical Anthomyiidae or the the 1¥在 uscidae (so st [，) 0 The internal organs of and its τelatives are ， at present ， poorly poorly known and it is believed that more evid 色nces in them would do much to clear their their relationship among the Calyptratae. Thus ， the position of Lis 戸g shown in the Figo 1 can be at pr 巴sent tentative. From the above it is clear that most ， if not a11 ，Anthomyiids 1at.) examined here ，even if they were so limited in show many morphological divergences in in their internal organs. This fact suggests that the Anthomyiidae as we define them in in thi3 paper contain 30m 巴 h色terogenou3 or distantly related groups. Many Di pterists such as Sεnio ト羽Thite ( Malloch (1 934) ，van Emden (1 939) ， Hennig (1 941 ，1965) ，Tokunaga (1 943) ，Roback (1 95 1)， (1 954) ，Fan (1 965) ，

Huckett Huckett (1 965) ，etc. ， a11 thεas in the IV l: uscidae ， Some authors in c1 uding Karl (1 928) ，Townsend (1 935) ，Zimin (195 1)， Hennig (1 964) ，

etc. ， retained th 己m as a tribe in thεMuscidae. Enderlein ( raised the 8tomoxyids to a separate family StomoxidaιShiraki ( follovved this Enderlein's system and adopted his family name Sto111oxidae. Essig ( named this family Sto111oxyidae ， while recently ( 1957 戸 adopted the family name Stomoxydidaeo The onlv one but decisive character of adopted these

workers workers is thεpiercing mouth~parts" Th 色se piercing mouthι包~p

mιore or 1邑ss dεveloped even in the different familiεs ， e.g. (8 にatophagidae) ，

Liste Liste and 渉oce ρhala yt is dear ，therefor 久 that these mouth 中 arts are undoubtedly undoubtedly adaptive and thIs alone does not constitute a great distinction

on the family or subfamily leve l. The g色nera Musca ， Pyrellia ，

Dasy ρ Orthellia ， and うho 幻a of the Muscidae are all distinct in lacking the male accessory glands (Hori ， 1960). But on

the the basis bf the arrangem 色nt of the rectal the number and arrangement of the the spermathecae ， the of the ejaculatory duct ， the Muscidae are divided into two groups ， the Muscinae in which l'vf uscina ，etc. ， are included ， and (b) the Stomoxydinae in which ancl are included (Hori ，1960 ， 1961 ， 1962a). The length of the the position of the rectal valve ， the number of the the of the proximal intestine ， the hairiness of of the arista ， the distal bencl of the fourth the head width ，etc. ， a l1 indicate that

the the Stomo 叉γids are more primitive than the typical Muscids. These primitive

characters characters of the internal together vyith the piercing mouth 四 parts offer a

キ Hennig (1964) ， p. 1033. Plzylogeny Plzylogeny of Calyttratae 245

very very valuable basis for separating th 巴 StJ 日(

Many Di 均pt 伐erist 匂s 凶nclue dピ心一li 日il1 g Seguy (1923 幻)， Karl (1 928) ， H 巴nnig (1 941) Colyer and

Hammond (1 95 1)， Ringdahl (1954) ，etc. ， placecl GraphOJnyia in the Muscinae 011 the basis basis of the sharp distal bencl of the fourth vein ， plumose arista ， holoptic eyes of males ， etc. Enderlein (1 936) shiftecl the genus GrapJ lO myia to the Morellinae on the basis basis of the bencl of the fourth vein ， the absence of the posterior bristle on seconcl tibia. tibia. Zimin (1 951) transfen 吋 the Gr ゆ homyia f1' om the Muscini to the Stomoxydini by the presence of the concaved posterio 1' margin of the compound eyes ， slender prob :J scis ，a small labellum ，narrow thoracic squamae ， the absence of a1 コcsterior bristle of of thircl tibia ， etc. But this Zimin's transfer is not generally favoured ye t. Van EInden

(1 956) believes that the Muscina 色'v vere formerly separated from the rest of the Muscidae by the bend in the vein m ， and thus Graphomyia was included in this subfamily. subfamily. This bend ， according to him ，was now known in s.o many unrelated forms that that its use as a subfamily character had to be abancloned. The external pattern ，head shape ， the dense pollinosity of the adult ancl aquatic larval type ，etc. ， of Gral うhomyia ， according according to van Emclen (1 956) ， are suggestive of those of the LimnoPhor ル group. Hennig (1 958) placecl Gra ρhomyia in the Mydaeinae on the basis of the absence of

the the postero 幽 clorsal b 1' istle on third tibia. So far as the evidences of these foregoing works are concerned ， it is clear that the genus Gra ρhomyia has some affinity to the LimnoPhora and also to the Stomoxys and its al1 ies ， ancl may be originatecl from the common ancestor of these flies. We can ，on the basis of th 巴 above eviclences and those those of internal morphology ，say that the Graphomyia has no such close kinship to the the typical Muscinae as many oIcler Dipterists claimecl. The exact placement of the genus Gra ρhomyia ，however ，may b巴 a difficult problem. Seguy (1923) ， Karl (1 928) ， Colyer ancl Hammond (1 951) ， Ringdahl (1 954) and many other olcler authors placed the Muscina in the subfamily Muscinae. Enderlein

(1 936) transferred the genus Muscina to his subf 呂m 日y Morellinae together with the genera genera Gr ゆ homyia ，A'V1 yios) うila ，Mesembrina ，Morellia ， where the second tibia is without without a poterior bristle. Recently Hennig (1 962) in c1 uded ， 246 246 Katsushige HORI are are of the typical Muscids such as the genera Morellia ， etc. etc. (see Roback's plate IV)ω Girschner Girschner (1 893) recognized the Calliphorids as a subfamily rank ， or a subfamily group ， in the family Tachinidae ，and in c1 uded the genera

Acrothaga ， 幼 ora and Lucilia in this subfamily 匂 Some older

Di pterists ， e.g. Williston consider 巴d that the Calliphorids are closely related to the

Muscids and included most of the typical Calliphorid g巴nera such as Callithora ， Lucilia ， Phormia ，Mesemorina ， etc. ， in his Muscidae. Crampton (1 942) considers that that the evidences of the male terminalia indicate that the Ca 1li phorids appear to be

somewhat closer to the Anthomyiidae than are to the Muscidae ， This view of

Crampton's may a1so be support 邑d the male internal sexual organs secured the p肥 sent writer (1 960). Senior-White Senior-White ( Imms (1 925) ，Lundbeck ( Hendel (1 928) ， (1. 928) ，etc. ， all recognized the Calliphorids as a of the Tachinidae. Shannon

(1 923) ，Comstock (1 924) ， Enderlein ( Crampton (1 942) ョ Tokunaga (1 943) ， Essig ( Roback (1 951) ，Ha l1 ( Fan etc" a11 treated the Ca 1li phorids as a

distinct distinct family ， S匂 uy (1 928) ， Senior-W 削除， Auber ・tin and Smart (1 940) ，Mesn i1

(1 944) ， Colyer and Hammond ( ，van Emd 巴n( etc" united the Calliphprids

and the Sarcophagids to form the Calliphoridae. Curran ( includ 己s the Calliphorids

and Sarcophagids together with some Muscids in his new Metopiidae ， It however ，c1 ear that the Calliphorids ，on the basis of the presence of the male access o. ry

glands ，and of the fat 阿 body envelopεs 01 the testes ， are c1 early on onεhand ，

fro )TI the Muscids and differ ，on the other from the Sarcophagids by the shape

of of the testes and male and female acc 巴ssory glands ， the arrangement of the spεrm か thecae ， the of the utεro-vaginalεtc The genεra of the Calliphoridae Calliphoridae such as P hormia ，

Triceratotyga Triceratotyga and Lucilia h乳ve many internal characters in common. The banana ・・

shaped male accessory glands ，oval 幽 shaped testes ， organge colour of the testes ， the

pr 邑sence of the fat 司 body of the testεs ， the arrangement of the rectal papillaε ，

etc 吋 are all quite common in these genera ， The male and female accessory glands of the genera and Sto n1 orhina are ，

however ，somewhat different from those of the rest of this family. On the ba 呂is of the the distal bend of the fourth the squamal index ， poorly dεveloped hypopleural

bristl 巴s， smaller width of the head ， the genera and Stomor hi 匁a which belong to to the subfamily Rhiniinae can be eunsidered more primitive than the Chrysomyinae or or Calliphorinae. The valve indices ， the coiling of 士he proximal intestine ， the fusing fusing clegree of the anterior Malpighian tubules ， etc" also incrεase fro111 the Rhininae through through the Chrysomyinae to the Calliphorinae. From the internal morphology (Hori ，1960 ，162a ， 1962h) it is dear that ， in the Calliphorinae ， the primitive characters such as the small size of the ejaculatory Phyloge 幻y of Calyttnitae 247

apo c1 eme ， the coiling degγ 巴三 of the pro 叉im81 intestine ， the arrangement of the posterior Ma11 コighian tubules ， the lDdex of the anterior and the small size

of of the ac1 ult etc. ，were retain 己d in the g巴ηllS .Melind a:. On the other hand in

the the gεnus Lucilia hεre 忠v 設minξd ，two of the "richardsi-group ヲヲ of

K urahashi L. s♂ ricata and L. Cli ヨウ 011 the r:asis of the internal characters

an c1 the s111all size of the 事7ell separated dichoptic eyes i11 the male ， arε

undoubtedly undoubtedly the mo" .t of the tribe Luci Ii ni. Th 日 differences between these

primitive primitive forms and the advanced Luci Zi a ，hov!ever ，do not se 巴m to me so remarkable as to warrant their into different genera. In the Ca !li phoridae ， of of a11 the internal organs ， the differentiation of the digestive organs and the h在旦 lpighian tubules w 巴re most noticeable (日 lori ，1962a ， 1962b). Sorne Sorne older worker .s such as Senior ベNhite ，Imms ， Lundbeck (1 927) ，

Hendel (1928) ，羽 1ainwright 歩合tc. ， a11 the Sarcophagids tqgether with

Calliphorids Calliphorids in the Tachinidae. Senior 回 ViT hite ，Aubertin and Smart (1 940) ， Seguy (J Mesnil ， Colyer and Hammond (1951) ， van Emden (1 954) ， Herting ， Zumpt ， etc. ， a11 amalgamated the Sareophagids with

the the Ca l1 iphorid .s to form the f丘町 Curr 泊 an (1 934) included the Sarco 凶

phagidae togethεr with thεCalliphori c1 ae and some others in his new family Meto 四 piidae. piidae. Coquillett (1号， Williston ， Comstock (1 924) ， Ende r1 ein (1 936) ，

Rohden c1 orf ， Tokunag 乱 (1942 ， ， Essig ， Robaek

(1 951) ， Shiraki ， Downes 邑tc. ， regarded the Sarcophagicls as a distinct family family rank. Crampton believes that thεSarcophagids have ，on the basis of the the terminalia ，some ， not much ， affinity to the Ca Ui phoridae 011 011e hancl ，and to the

Tachinidae on the oth 巴r hand. But the evidences of the internaJ l11 0rphology such as the the testεs ，male and female accessory glands ， the arrangement of the spermathecae ，

the the presence of the incubatory ppuch ， the presence of fOUf 同 notopleurals in the adults ，

etc. ， a11 Indicate that the Sarcophagidae are eonsiderably different from the Calli 明 phoridae phoridae than the latter are from t11εTachinidaιThe shapεs of the testes are l11 0re similar similar to that of thεthan to that of thεCa lJi phoridae. The aedeagi of the Sarcophagids 3llCh as Camtto ρs unicolor are ， accorcling accorcling to Roback (1 954) ， very dose in foτm to thosεof some extant Scatophagi c1 s such as stercorarium ancl cincta. This indicates that the Sarcophagiclae Sarcophagiclae probably arose from the ancestor of the 111me primitive of the Scato- phagids. phagids.

In In the Sarcophagiclae it was noted the al1 thor (1960) that the ejaculatory duct incrεases in 色ngth gεnerally i l1 accordance with the rise of their systematic position.

Generally Generally speaking ， in the ge l1 lls (s. laL) both the ext 巴rnal appearance and the internal organs were seen to have remained remarkably unaltered in contrast to to their male and female terminalia in which considerablevariation had occured. Of the three subfamijies of the Sarcophagidae here treated 'Nhich have bεen 248 248 Katsushige HORI

on the b乳sis of thε degree of the proximal intestine ， the characters tubules tubules as well as the adult characters such as the bare arista ， sma11 sma11 size of adult head ，etc. ， the subfamily Metopiinae can be consider 己d the most and the are ，on th 巴 basis of the we lI dεmale gεnitalia ， the most sp 巴cialized and have become the largest largest group in the

Some D 叩t巴rists in c1 uding Allen ，etc. ， thought that the

8・eneral charaderistics of thεMetopiinae are essentially sarcophagoid and have

arisen arisen from the same ancestor as the Sarcophaginae. The banana “shaped male accessory accessory glands of seem ， expressive that this is a1so a connectant leading leading to the

The g巴nus is closε1y r巴lated to the Sarcophagids. The short ejaculatory three three to five hypopleurals ，etc. ， of

In In the is ， without doubt ， primitive on the basis basis of the coiling 巴e of intestine ， short ejaculatory duct ， the arrangernent arrangernent of the three to five etc.

Th 色 extεrnal chaetota ちを y of the Sarcophaginae with their internal shows a few characters that are of value in determining determining the course of evolution of these but the aedεagi with their wealth of of diverse structures as noticed many have been found to offer valuable systematic systematic evidences upon which relationships and phylogenetic conclusions within the can be based. ， the arrangemεnt of each of the Sarcothaga-group in in Fig" 1 can but at presεnt ， and their definitive arrangement

i1l w i1l have to wait for further 吋 increase in our In In the past tI1 e close betweεn thεSarcophagidae and Calliphoridae has been overrated bぅT S01γle (Curran ， 1934; Sεnior ベiV hite et a1.， 1940; Seguy ， 1941; 1941; and Hammond ， 1951; van Emden ， Zumpt ， 1956). The evidences evidences of the internal organs of the Sarcophagidae together with their peculiar external external moγcharactεristics will separate them from the Call ト phoridae phoridae or Tachinidae ，

Some authors such as Stein ，Senior 国 White (1 924) ，Imms (1 925) ，Seguy (1 928) ， εtc. ， adopted a c1 assification of the Calyptratae which divided divided them into two families: Anthomyiidae ， or Muscidae ，and Tachinidae ， In this Tachinidae Tachinidae both the and together with typical Tachinids are included. included. Comstock ，Crampton (1 942) ， Colyer and Hammond (1 951) ，van Emden ，Oldr 叫 dεtco ， excluded them from the Tachinidae.

80mεauthors such as I1ennig (J ，Crampton (1 942) ， Essig (1 947) ， et c.， recog 四 nized nized the Phasiids and Dexiids as a distinct family respectively. But Crampton

250 250 Kat 活ushige I- IOIU:

"，，"， JU"HC that the on thεbasis of the evidence of the male termi 暢 nalia ，apparεdescended more or less from the Tachinid ancεstOL The

Phasiids Phasiids and Dexiids 呂町 sometimes treated as subfamilies of the Tachinidae ， but are are here as a separate The Phasiids are ， in either case ， considered to be the more of the Tachinidae many Dipterists. As

a matter of fact the testes of a Phasiid うhasia are sim i1 ar to those of of the and their malεaccessory recall those of the Calliphorids

1960). 1960). The lamp 回 testeswhich were observed in the typical Tachinids examined seem to be a characteristic to the Tachinidae

The number of here treated tco small to discuss the relation 回 ship ship of the Tachinidae in any finεr details. From Table I it is seen that the internal organs organs of the Tachinidae such as the canal and the tubul 出 indicate indicate at least their or reduced condition. their their distinctness above mentioned with their l乱rval parasitisrn ， it appears that the ancestral To ，chinids represent a line line which diverted frcrn the rnain stem of the Phasiids Phasiids show several features which connect them to both the Calliuhoridae and Sarcophagidae. Sarcophagidae.

Fro l11 the for 唱。ing it is clεar that the major charact 巴rs which were sufficient sufficient to regroup the families of the into some were here not not found ， thus it may be advisable to them in a separate The of a11 the forms discusad here are tentatively shown in FiιL

1n 1n the past various systems have be 己n for the c1 assification of the Caly 岨 ptratae ptratae mainly on the basis of the external characters of the adult the male the the larval etc. No tV{O have agreed on the the genεric or divisions to be used.

1n 1n the present account a co~ordinate consideration of both the adult internal organs organs and some external characters of 83 speci 田 of the from Japan was rnade. The presence or and the of the male accessory glands ， the and colou ず of the the of the εthe position position of the rectal valve ， the of the th 己 arrange- ment of the コ lUU'U1C"' ，etι ， as well as some external adult characters such as as the hairiness of the the distal bend of the fourth longitudinal vein ， the size of of the squamae ， the bristles ， the head etc. ，have to be of major in the 1n the Calyptrataeω It was made clear from the present study 、 of these that the male ac c: essory glands have great Phyloa e日:y of Calyptmfae 251 taxonomic value above the gεneric leve 1. Thus the Calyptratae are ，on the basis of presence or absence of the male accessory glan c1 s，separable into two main groups.

Several genera such as Anihomyia ，Fannia ，01 うhyra ，Pegomyia ，Lis1 りe，etc. ，which are sometimes included in the Anthomyiidae (s. la t.) by some older Dipterists ，are likewise divided into two groups. This indicat 巴s that these small Anthomyiid Hi es are the key-group to solve the relationships among the Calyptratae as suggested by Crampton (1944). This study emphasizes a1so a need for examination of far more materials of this group than is now available in order to establish the reJationships among the Calyptratae.

The differentiation of the intεrna1 organs has not al 'v vays the same trend in the higher c1 ivisions such as the Muscidae ，Calliphoridae ，Sarcophagidae ， etc. This could indicate indicate that they wereεstablished very early from the common anc 巴stor as a quite different different respective systematic group. Thes 巴 probable relations of a11 the groups are tentatively figured. It is a1so noticeable that the differentiation of the internal organs are c1 0sely correlated to the ext 巴rnal characters such as the body size of the adult flies.

References

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