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uaestiones

Entomolosicae

A periodical record of entomological investigations,

published at the Department of Entomology,

University of Alberta, Edmonton, Canada.

VOLUME 25

NUMBER 1-2

WINTER-SPRING 1989

Publication of Quaestiones Entomologicae was started in 1965 as part of a

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Issued May 1989

QUAESTIONES ENTOMOLOGICAE

ISSN 0033-5037

A periodical record of entomological investigation published at the Department

of Entomology, University of Alberta, Edmonton, Alberta.

Volume 25 Number 1 1989

CONTENTS

O’Hara-Systematics of the Genus Group Taxa of the Siphonini (Diptera:

Tachinidae) 1

Commentary 231

Book Review 239

SYSTEMATICS OF THE GENUS GROUP TAXA OF THE SIPHONINI

(DIPTERA: TACHINIDAE)

James E. O’Hara

Department of Entomology

University of Alberta

Edmonton, Alberta, Canada Quaestiones Entomologicae

T6G2E3 25:1-229 1989

ABSTRACT

The Siphonini are a monophyletic tribe including 386 named species (including

subspecies and nomina dubia, but excluding nomina nuda and misspellings ), of

which 294 are recognized as taxonomically valid species. The genera and

sub genera of the tribe are revised for the world fauna, based on study of adults of

most described and more than 100 undescribed species, and first instars of 68

species. Seven genera are recognized, each hypothesized as monophyletic:

Goniocera Brauer and Bergenstamm, Proceromyia Mesnil ( elevated from

subgeneric status within Ceromya and including Nipponoceromyia Mesnil and

Shima, n. syn.), Entomophaga Lioy, Ceromya Robineau-Desvoidy (including new

synonym Actinactia Townsend), Actia Robineau-Desvoidy, Peribaea

Robineau-Desvoidy and Siphona Meigen. Ceromya is informally divided into

Ceromya sensu stricto and the Ceromya silacea species group. The concept of

Siphona is broadened to include the following nine subgenera, each previously

recognized as a genus and each (except for one, as noted ) hypothesized as

monophyletic: Actinocrocuta Townsend (new subgeneric status ), Aphantorhapha

Townsend (new sub generic status ), Aphantorhaphopsis Townsend (new sub generic

status; an unresolved and possibly polyphyletic group of Old World species,

including Asiphona Mesnil, n. syn.), Baeomyia O’Hara (new subgeneric status),

Ceranthia Robineau-Desvoidy (new subgeneric status), Pseudosiphona Townsend

(new subgeneric status), Siphona Meigen sensu stricto, Siphonopsis Townsend (new

subgeneric status) and Uruactia Townsend (new subgeneric status). Three species

groups, and a group of unrelated and unplaced species, of undescribed New World

Siphona s.l. species are reviewed but not formally named or described.

New combinations resulting from reclassification of the Siphonini are:

Nipponoceromyia pubioculata Mesnil and Shima moved to Proceromyia; Actia

amblycera Aldrich, Actia comuta Aldrich, Actinactia lutea Townsend and Actia

normula Curran moved to Ceromya; Actia stiglinae Bezzi moved to Peribaea; Actia

panamensis Curran and Aphantorhapha pulla Reinhard moved to Siphona s.l. (and

unplaced to subgenus); and the following 20 species are provisionally placed in S.

(Aphantorhaphopsis) (most formerly placed in Asiphona, a junior synonym of S.

2

O’Hara

(Aphantorhaphopsis)): Crocuta (Siphona) alticola Mesnil, Actia angustifrons

Malloch, Actia brunnescens Villeneuve, Crocuta (Siphona) crassulata Mesnil,

Siphona (Asiphona) fera Mesnil, Siphona (Asiphona) laboriosa Mesnil, Actia

laticomis Malloch, Actia mallochiana Gardner, Siphona (Asiphona) nigronitens

Mesnil, Actia norma Malloch, Asiphona picturata Mesnil, Siphona (Asiphona)

pudica Mesnil, Actia samarensis Villeneuve, Actia selangor Malloch, Thryptocera

selecta Pandelle, Gymnopareia siphonoides Strobl, Siphona (Asiphona) speciosa

Mesnil, Actia starkei Mesnil, Actia verralli Wainwright and Siphona (Asiphona)

xanthosoma Mesnil. Actia heterochaeta Bezzi, a nomen dubium, is left unplaced

within the Siphonini. Removed from the Siphonini is Actia aberrans Malloch

( referred to Eurysthaea Robineau-Desvoidy by Crosskey, pers. comm.).

The status of named species has, in general, been maintained as listed in current

literature, though the following new synonymies were recognized and are herein

proposed: Actia brevis Malloch synonymized with A. darwini Malloch, Peribaea

subaequalis (Malloch) with P. orbata (Wiedemann), Actinocrocuta chaetosa

Townsend with Siphona (Actinocrocuta) singularis (Wiedemann), and Siphona

janssensi (Mesnil) with S. (Siphona) cuthbertsoni Curran. Elevated from subspecific

to species status are: Ceromya similata Mesnil, Siphona (Siphona) infuscata

(Mesnil) and S. (Siphona) nigrohalterata Mesnil. Lectotypes are designated for

Ceromya cibdela (Villeneuve) and S. (Pseudosiphona) brevirostris Coquillett.

A chapter about structural features of the Siphonini is followed by a key to

genera and then by descriptions of genera and subgenera. Keys are also provided to

the species of Proceromyia and Entomophaga and for the subgenera of Siphona s.l.

A list of known siphonine hosts is included. A cladistic analysis based on ingroup

( Siphonini ) and outgroup (other Tachinidae) comparisons is used to partially

reconstruct the phylogenetic history of the major siphonine lineages. Geographic

distribution of siphonine genera and subgenera is discussed in relation to historical

events during earth history and the presumed phylogenetic history of the Siphonini.

RESUME

La Siphonini est une tribu monophyletique qui comprends 386 especes nommees

(y compris les sous-especes et nomina dubia, mais a V exclusion de nomina nuda et

ses mauvaises epelations ). De ces especes, 294 sont reconnues comme etant

taxonomiquement valides. Les genres et sous-genres de la tribu sont revises pour la

faune mondiale, ce base sur une etude des adultes de la plupart des especes

decrites, des adultes de plus de 100 especes non- decrites, et des larves de premier

stade de 68 especes. On reconnait sept genres, et on hypothese que chacun de

ceux-ci est monophyletique: Goniocera Brauer et Bergenstamm, Proceromyia

Mesnil (eleve du statu sous-genre dans Ceromya et y compris Nipponoceromyia

Mesnil et Shima, n. syn.), Entomophaga Lioy, Ceromya Robineau-Desvoidy (y

compris le nouveau synonyme Actinactia Townsend), Actia Robineau-Desvoidy ,

Systematics of the Genus Group Taxa of the Siphonini

3

Peribaea Robineau- Desvoidy et Siphona Meigen. A titre non-officiel, Ceromya est

divise en deux groupes: Ceromya sensu stricto et le groupe d’especes Ceromya

silacea. Le concept de Siphona est elargi pour inclure les neuf sous- genres suivants,

chacun prealablement reconnu comme genre et chacun (a V exception dun tel, que

note) etant par hypothese monophyletique : Actinocrocuta Townsend ( nouveau statu

sous-generique), Aphantorhapha Townsend ( nouveau statu sous-generique),

Aphantorhaphopsis Townsend ( nouveau statu sous-generique; un groupe irresolu et

possiblement polyphyletique d’especes de Vancien monde, y compris Asiphona

Mesnil, n. syn.), Baeomyia O’Hara ( nouveau statu sous-generique ), Ceranthia

Robineau-Desvoidy (nouveau statu sous-generique), Pseudosiphona Townsend

(nouveau statu sous-generique), Siphona Meigen sensu stricto, Siphonopsis

Townsend (nouveau statu sous-generique). Trois groupes d’especes, et un groupe

d’especes sans liens de parente et d’especes non-classees, de Siphona s.l. du

nouveau- monde sont examines sans etre nommes ou decrits formellement.

Les nouvelles combinaisons resultant de la reclassification des Siphonini sont:

Nipponoceromyia pubioculata Mesnil et Shima passe a Proceromyia; Actia

amblycera Aldrich, Actia comuta Aldrich, Actinactia lutea Townsend et Actia

normula Curran passent a Ceromya; Actia stiglinae Bezzi passe a Peribaea; Actia

panamensis Curran et Aphantorhapha pulla Reinhard passe a Siphona s.l. (et

non-classe au niveau du sous-genre); et les 20 especes suivantes sont

provisoirement placees dans S. (Aphantorhaphopsis) (plus anterieurement placees

dans Asiphona, un synonyme junior de S. (Aphantorhaphopsis)): Crocuta (Siphona)

alticola Mesnil, Actia angustifrons Malloch, Actia brunnescens Villeneuve, Crocuta

(Siphona) crassulata Mesnil, Siphona (Asiphona) fera Mesnil, Siphona (Asiphona)

laboriosa Mesnil, Actia laticomis Malloch, Actia mallochiana Gardner, Siphona

(Asiphona) nigronitens Mesnil, Actia norma Malloch, Asiphona picturata Mesnil,

Siphona (Asiphona) pudica Mesnil, Actia samarensis Villeneuve, Actia selangor

Malloch, Thryptocera selecta Pandelle, Gymnopareia siphonoides Strobl, Siphona

(Asiphona) speciosa Mesnil, Actia starkei Mesnil, Actia verralli Wainwright et

Siphona (Asiphona) xanthosoma Mesnil. Actia heterochaeta Bezzi, un nomen

dubium, demeure non-classe parmi les Siphonini. Actia aberrans Malloch est enleve

des Siphonini (et rapporte a Eurysthaea Robineau-Desvoidy par Crosskey, comm,

pers.).

Le statu des especes nommees a ete maintenu en general, tel qu inscrit dans la

litterature, a V exception des nouvelles synonymies reconnues et proposees comme

suit: Actia brevis Malloch synonyme de A. darwini Malloch, Peribaea subaequalis

(Malloch) synonyme de P. orbata (Wiedemann), Actinocrocuta chaetosa Townsend

synonyme de Siphona (Actinocrocuta) singularis (Weidemann), et Siphona janssensi

(Mesnil) synonyme de S. (Siphona) cuthbertsoni Curran. Les suivantes sont elevees

de sous-especes a especes: Ceromya similata Mesnil, Siphona (Siphona) infuscata

(Mesnil) et S. (Siphona) nigrohalterata Mesnil. Des lectotypes sont designes pour

Ceromya cibdela (Villeneuve) et S. (Pseudosiphona) brevirostris Coquillett.

Quaest. Ent., 1989, 25 (1,2)

4

O’Hara

Un chapetre sur les traits structuraux des Siphonini est suivi d’une clef pour les

genres, et ensuite par les descriptions des genres et sous-genres. Des clefs sont

aussi incluses pour les especes de Proceromyia et Entomophaga et pour les

sous-genres de Siphona s.l. Une liste des hotes reconnus pour les siphonines est

incluse. Une analyse cladistique basee sur des comparaisons en groupe ( Siphonini )

et hors groupe (autres Tachinidae) est utilisee pour reconstruire partiellement

I’histoire phylogenetique des lignees majeurs siphonines. La distribution

geographique des genres et sous-genres des siphonines est discutee en relation avec

les evenements historiques de revolution de la terre, et I’histoire phylogenetique

presomptueuse des Siphonini.

TABLE OF CONTENTS

Introduction 5

Materials and Methods 6

Materials 6

Methods 9

Structural features 13

Historical review 13

Structural features of the Siphonini 14

Classification of the Siphonini 34

List of major references to the Siphonini 34

Diagnosis of adults of Siphonini Rondani 34

Review of major keys to genera and subgenera of the Siphonini 36

Key to adults of the genera of the Siphonini 37

Notes about classification chapter 39

Goniocera Brauer and Bergenstamm 41

Proceromyia Mesnil 44

Key to adults of Proceromyia species 44

Entomophaga Lioy 47

Key to adults of Entomophaga species 48

Ceromya Robineau-Desvoidy sensu lato 5 1

Ceromya Robineau-Desvoidy sensu stricto 52

Ceromya silacea (Meigen) species group 63

Actia Robineau-Desvoidy 67

Peribaea Robineau-Desvoidy 77

Siphona Meigen sensu lato 84

Key to adults of the subgenera of Siphona sensu lato 84

Hosts 158

Introduction 158

Explanation of parasite-host list 158

Evolution of the Siphonini 168

Systematics of the Genus Group Taxa of the Siphonini

5

Introduction 1 68

Monophyly of the Siphonini 169

Monophyly of genera and subgenera 1 7 1

Monophyly of the Proceromyia Mesnil and Entomophaga Lioy lineage 178

Monophyly of the Peribaea R.-D. and Siphona Meigen lineage 178

Phylogenetics of non -Siphona s.l. siphonine lineages 179

Phylogenetics of Siphona subgenera 181

Historical zoogeography 189

Concluding remarks 200

Acknowledgements 200

Literature Cited 201

Postscript 218

Index 220

INTRODUCTION

The Siphonini are a monophyletic tribe of about 300 described species and more

than 100 undescribed species, of uncertain phylogenetic position within the

Tachinidae (some authors favoring its placement in the Goniinae, others the

Tachininae). Adultk are smaller than most tachinid flies, generally 3-5mm in length,

and larvae are almost exclusively parasitoids of larval Lepidoptera. Siphonines are

widely distributed throughout the world, with greatest diversity in non-rainforest

areas of the tropics.

Mesnil established the modem concept of the Siphonini (as Siphonina) in his

illustrious tachinid contribution to “Die Fliegen der palaearktischen Region”

(1962-5). Prior to that time, several genera now relegated to the Neaerini were

included because of their external similarity to members of the Siphonini (e.g.

Mesnil 1939, 1954; van Emden, 1954). Herting (1957), in his interpretive survey of

female genitalia in the Calyptratae, was first to recognize the error in this

classification and his findings were subsequently incorporated into Mesnil’ s

(1962-5) revision of the Palearctic Siphonini. Crosskey maintained Mesnil’ s concept

of the tribe in his treatments of the tachinid faunas of the Australian (1973), Oriental

(1976a) and Afrotropical (1980) regions, as did Andersen (1983) in his recent

generic revision of the Old World Siphonini and Herting (1984) in his catalogue of

Palearctic Tachinidae.

In marked contrast to the recent attention conferred upon Old World siphonines,

the New World fauna is without a modem treatment at any level. Townsend’s

(1940) revision of the “Siphonini” and “Actiini” in his “Manual of Myiology” is an

untenable classification comprising a heterogeneous and polyphyletic assemblage of

genera. A valuable step toward a modem classification of North American Siphonini

was achieved by Sabrosky and Amaud (1965). These authors brought together

almost all the siphonine taxa of America north of Mexico within their subtribe of the

Siphonini, the Siphonina, but included therein a number of unrelated genera.

Quaest. Ent., 1989, 25 (1,2)

6

O’Hara

Guimaraes (1971) catalogued the tachinids of America south of the United States,

and though following the broad concept of the Siphonini advanced by Sabrosky and

Amaud, he did not similarly recognize subtribes; consequently his genera of the

Siphonini sensu Mesnil are interspersed with a variety of unrelated taxa.

This study was undertaken with the aim of revising the supraspecific taxa of the

Siphonini and in particular reclassifying the New World siphonines in light of recent

advances concerning the Old World fauna. This aim has been tempered somewhat

by the diverse and largely undescribed fauna of the Neotropics, which requires

detailed study and description of species before a satisfactory supraspecific

classification can be established. The results of a preliminary study of many

undescribed Neotropical species are included here as an aid to future systematic

research on the Neotropical Siphonini.

In addition to revising the Siphonini and providing a framework about which

future revisions of genera can proceed, this work is also an exploration into the

phylogenetic relationships among the supraspecific taxa and character state diversity

among species. To attain this goal, most available types and specimens of many

undescribed species were examined.

Included in this paper are lists of species names in the Siphonini, arranged

according to a revised classification of the tribe. Descriptions of the genera and

subgenera are provided along with keys to the genera of the Siphonini, subgenera of

Siphona s.l. and species of Proceromyia and Entomophaga. Known hosts of the

Siphonini are listed. Character state diversity is tabulated and discussed, and the

phylogenetic and geographic history of the supraspecific taxa is analyzed.

MATERIALS AND METHODS

Materials

This paper is based on the morphological study of adult specimens of about 270

of the 294 described species of the Siphonini, more than 100 undescribed New

World species, and numerous undescribed Old World species. Included among these

specimens were 264 primary types and eight paratypes; types not examined include

31 known to be lost (22 of these being Robineau-Desvoidy nominal species) and 13

not located and possibly lost. Male genitalia of more than 250 species and female

genitalia of about 90 species were examined. First instars of 68 species were also

examined (see O’Hara in press “a”).

Codes used in the text for designating institutions and private collections appear

below with the names of their respective curators.

AMNH Department of Entomology, American Museum of Natural

History, Central Park West at 79th Street, New York, NY

10024; the late P. Wygodzinsky and K.A. Schmidt.

Biological Laboratory, College of General Education, Kyushu University,

Ropponmatsu, Fukuoka 810, Japan; H. Shima.

BLKU

Systematics of the Genus Group Taxa of the Siphonini

7

BMNH Department of Entomology, British Museum (Natural History), Cromwell

Road, London, England SW7 5BD; R.W. Crosskey and A.C. Pont.

BPBM Department of Entomology, Bernice P. Bishop Museum, P.O. Box 6037,

Honolulu, HI 96818; N.L. Evenhuis.

CAS Department of Entomology, California Academy of Sciences, Golden

Gate Park, San Francisco, CA 94118; P.H. Amaud, Jr.

CEA Coleccion de la Estacion Experimental Agronomica, Universidad de

Chile, Maipu, Chile (curator not named).

CIE Coleccion del Centro de Investigacions Entomologicas, Universidad de

Chile, Santiago, Chile; J. Herrera.

CNC Biosystematics Research Centre [formerly Institute], Central

Experimental Farm, K.W. Neatby Building, Ottawa, Ontario K1A 0C6;

D.M. Wood.

DEI Institut fiir Pflanzenschutzforschung Kleinmachnow, [formerly the

Deutsches Entomologisches Institut], Akademie der Landwirt-

schaftswissenschaften, Bereich Eberswalde, 13 Eberswalde-Finow 1,

Schicklerstrasse 5, German Democratic Republic; H.J. Muller and R.

Gaedike.

DPI Entomology Branch, Department of Primary Industries, Meiers Road,

Indooroopilly, 4068, Queensland, Australia; B. Cantrell.

ETH Entomologisches Institut, Eidgenossische Technische Hochschule,

Zentrum, CH-8092 Zurich, Switzerland; W. Sauter.

FRI Forest Research Institute, Dehra Dun, Uttar Pradesh, India. (I was unable

to establish contact with this institution.)

FSF Forschungsinstitut Senckenberg, Senckenberganlage 25, D-6000

Frankfurt 1 , Federal Republic of Germany; J. Rademacher.

HDE Hope Department of Entomology, University Museum, Oxford, 0X1

3PW, England; M.J. Scoble.

HNHM Zoological Department, Hungarian Natural History Museum, H-1088

Budapest, Baross u. 13, Hungary; F. Mihalyi.

INPA Instituto Nacional de Pesquisas da Amazonia, Caixa Postal 478, 69.000

Manaus, Amazonas, Brazil; J.A. Rafael.

IRSN Institut Royal des Sciences Naturelles de Belgique, Rue Vautier 31,

B-1040 Bruxelles, Belgium (curator not named).

JEOH Private collection of the author.

MBR Division Entomologfa, Museo Argentino de Ciencias Naturales

“Bernardino Rivadavia”, Avenida Angel Gallardo 470, Buenos Aires,

Argentina; A.O. Bachmann.

MCSN Museo Civico di Storia Naturale, 20121 Milano, Corso Venezia 55, Italy;

C. Leonardi.

MCZ Museum of Comparative Zoology, Harvard University, Cambridge, MA

02138; M.K. Thayer.

Quaest. Ent., 1989, 25 (1,2)

8

O’Hara

MNHN Entomologie, Museum National d’Historie Naturelle, 45 bis Rue Buffon,

Paris Ve, France; L. Matile.

MRAC Entomology Section, Musee Royal de l’Afrique Centrale, B-1980,

Tervuren, Belgium; E. De Coninck.

MSU Department of Entomology, Michigan State University, East Lansing, MI

48824; R.L. Fischer.

MZF Museo Zoologico de “La Specola”, Universita Degli Studi, Via Romana,

17-50125 Firenze, Italy; S. Mascherini.

NMBA Naturhistorisches Museum der Benediktiner-Abtei Admont, A-8911

Admont, Austria; B. Hubl.

NMV Entomology Department, Naturhistorisches Museum, Zoologische

Abteilung, A- 1014 Vienna, Burgring 7, Austria; R.

Contreras-Lichtenberg.

NRS Entomology Section, Swedish Museum of Natural History, S- 10405

Stockholm, Sweden; P.I. Persson.

OSU Department of Entomology, Oregon State University, Corvallis, OR

97331; M.D. Schwartz.

PHA Private collection of P.H. Amaud, Jr., c/o CAS.

PPRI Department of Agriculture, Plant Protection Research Institute, National

Collection of Insects, Private Bag XI 34, Pretoria 0001, Republic of South

Africa; M.W. Mansell.

SMNS Staatl. Museum fur Naturkunde, Rosenstein 1, D-7000 Stuttgart 1,

Federal Republic of Germany; B. Herting.

SPHTM Entomology Section, School of Public Health and Tropical Medicine,

Sydney University 2006, Sydney, New South Wales, Australia; M.L.

Debenham.

UASM Department of Entomology, Strickland Museum, University of Alberta,

Edmonton, Alberta T6G 2E3; G.E. Ball.

UAT Department of Entomology, University of Arizona, Tucson, AZ 85721;

F.G. Werner.

UCB Division of Entomology and Parasitology, University of California,

Berkeley 4, CA 94720; J.A. Chemsak.

UCD Department of Entomology, University of California, Davis, CA 95616;

R.O. Schuster.

UCR Department of Entomology, University of California, Riverside, CA

92521; S.I. Frommer.

UCS Facultad de Agronorma, Universidad de Chile, Casilla 1004, Santiago,

Chile; R. Cortes.

UKL Department of Entomology, Snow Entomology Museum, University of

Kansas, Lawrence, KS 66045; C.D. Michener and G.W. Byers.

UNH Department of Entomology, University of New Hampshire, Durham, NH

03824; J.F. Burger.

Systematics of the Genus Group Taxa of the Siphonini

9

UNLP Division de Entomologia, Facultad de Ciencias Naturales y Museo,

Universidad Nacional de La Plata, Paseo del Bosque, 1900 La Plata,

Argentina; L. de Santis.

USNM Systematic Entomology Laboratory, SEA, U.S. Department of

Agriculture, United States National Museum, Washington, DC 20560;

C.W. Sabrosky (retired) and N.E. Woodley.

USP Museu de Zoologia, Universidad de Sao Paulo, Avenida Nazare, 481,

Caixa Postal 7172, 04263, Sao Paulo, sp, Brazil; J.H. Guimaraes.

UZI Department of Zoology, Zoological Museum, Helgonavagen 3, S-223 62,

Lund, Sweden; R. Danielsson.

WLD Private collection of W.L. Downes, c/o MSU.

WSUP Department of Entomology, James Entomological Collection,

Washington State University, Pullman, WA 99164; W.J. Turner.

ZIL Zoological Institute, USSR Academy of Sciences, Leningrad 199164,

USSR; V.A. Richter.

ZMA Instituut voor Taxonomische Zoologie, Zoologisch Museum, Universiteit

van Amsterdam, Afdeling Entomologie, Plantage Middenlaan 64, 1018

DH Amsterdam, Netherlands; T. van Leeuwen.

ZMHU Museum fur Naturkunde an der Humboldt-Universitat zu Berlin, Bereich

Zoologisches Museum, DDR 104 Berlin, Invalidenstrasse 43, German

Democratic Republic; H. Schumann.

ZMU Division of Entomology, Zoological Museum of the University,

SF-00100 Helsinki 10, Finland; B. Lindeberg.

ZMUC Zoologisk Museum, Universitetsparken 15, DK 2100, Copenhagen,

Denmark; S. Andersen.

Methods

Ranking of taxa. — Ideally a classification should be practical to the general

taxonomist, informative to the specialist, and an accurate reflection of the

hypothesized cladistic relationships among included taxa. In reality few groups can

be so perfectly classified, as the three criteria are often to a greater or lesser extent at

odds with one another. The problem is particularly acute when cladistic

relationships are inadequately known, as in the Siphonini. In light of this problem,

and in an effort to adhere as closely as possible to the three criteria given above, a

classification for the Siphonini is proposed using the formal supraspecific categories

of genus and subgenus and informal category of species group, with different

criteria for each.

In addition to the accepted criterion of being phenetically distinct from one

another (an admittedly vague concept), genera recognized herein are hypothesized

as monophyletic (holophyletic) lineages which are distinguishable by non-genitalic

characters (except for a few species possessing key-character parallelisms). Though

in combination these criteria are still subjective, they are stated here simply to give

Quaest. Ent., 1989, 25 (1,2)

10

O’Hara

the reader some indication of the guidelines followed in erecting the Siphonini

classification proposed herein; genera (like all taxa) are by nature subjective entities

undefinable by objective criteria. Most significantly, the criteria used here have led

to a broadening of Siphona to include a number of previously recognized genera

(mostly of Townsend). This broadened concept of Siphona comprises a

monophyletic group recognizable by the non-specialist; lower ranks within the

genus are used (as explained below) to serve the needs of the specialist.

The rank of subgenus has been used sparingly during the taxonomic history of

the Tachinidae, particularly by the most prolific describer of New World tachinids,

C.H.T. Townsend. Townsend had a very restricted generic concept, generally

erecting new genera of questionable value for the inclusion of a very few species.

As Townsend’s work is re-evaluated, most of his genera are being combined with

others or his original generic limits expanded. With respect to the Siphonini, most of

his genera proposed for non -Siphona s.l. species have been combined with other

genera, leaving only his Siphona group genera, representing mostly New World

taxa, unrevised. Whereas Townsend’s genera previously combined with

non -Siphona genera required no change to the concepts of those taxa, the same is

not true of his Siphona group names. Each of these names (i.e. Actinocrocuta,

Aphantorhapha, Aphantorhaphopsis , Pseudosiphona, Siphonopsis and Uruactia)

designate lineages of unknown relationship to one another. Either they must all be

retained as genera or the generic limits of another genus broadened for their

inclusion, if one is to follow the criterion of monophyly stated above for genera. In

addition, most of these lineages are phenetically similar, and only distinguishable by

male genitalic characters. A compromise position has been adopted here by reducing

Townsend’s Siphona group genera to subgenera within a broadened concept of

Siphona. Siphona s.l. is strictly monophyletic under this classificatory scheme, and

lower units are given the equal rank of subgenus because of the unknown cladistic

relationships among them. Thus such distinctive groups as Baeomyia and Ceranthia

are also assigned subgeneric rank along with Townsend’s names because their

exclusion would make Siphona s.l. paraphyletic. By retaining Townsend’s genera in

a formal classification, albeit at lower rank, the taxonomic and phylogenetic unity of

each of these taxa is retained, which would otherwise be lost if Siphona was

broadened and no lower ranks recognized. Ideally, I would like to see some New

World subgenera of Siphona s.l. combined with one another in the future, but at

present no synapotypies are known upon which to base such action. More

taxonomically and phylogenetically useful characters need to be discovered so that

the numerous undescribed species of Siphona s.l. are not classified into new Siphona

subgenera of unnecessarily narrow limits.

All Siphona subgenera are considered strictly monophyletic except one. An

exception has been made for S. ( Aphantorhaphopsis ) on practical grounds. Old

World species of the Siphona group not belonging to Siphona s.s. or S. ( Ceranthia )

have been assembled together in S. ( Aphantorhaphopsis ) as a preliminary measure

Systematics of the Genus Group Taxa of the Siphonini

11

until the group can be adequately revised. Many of its described species are known

only from holotypes - some male, others female - severely restricting their detailed

study. Given also the rich undescribed fauna that belongs within this group, a

revision of this assemblage is not attempted within this work. Preliminary study of

these species suggests they have few, if any, close relationships with New World

Siphona s.l. species, and almost certainly no close relationships with the New World

Siphona subgenera recognized here.

The lowest supraspecific rank used is that of the informal species group. This

rank is used for strictly monophyletic assemblages of species, and is used in two

parts of the classification, once in Ceromya s.l. and again in Siphona s.l.

The species of Ceromya s.l. are arranged in Ceromya s.s. and the C. silacea

species group, each tentatively hypothesized as monophyletic. This arrangement is

presently incomplete because each can only be recognized by examination of male

genitalia and not all species of Ceromya s.l. were examined for male genitalic states.

The two groups of Ceromya s.l. are referred to as above to reflect the tentative

nature of these findings and to avoid premature changes to the classification. If

future studies support the monophyly of each group and their sister group

relationship, then both could be formally recognized as subgenera of Ceromya s.l.

Nearly 100 undescribed siphonine species are known from the New World. More

than half belong to such distinctive taxa as Actia, Ceromya and Siphona s.s., or are

included in the revised concepts of such Siphona subgenera as Pseudosiphona and

Siphonopsis. The remainder do not exhibit clear relationships with any of the named

taxa, although relationships are evident among some of these species. To reflect

these relationships, document diversity, and to guide future studies on the

assemblage, three numbered species groups are separately discussed, and the

remaining 20-odd species treated as a whole. Thus the assemblage of known but

undescribed species of New World Siphona s.l. is discussed in as detailed a manner

as possible at this time. The naming of new species and possibly supraspecific taxa

awaits a more detailed systematic examination of this diverse and undescribed

fauna.

Treatment of undescribed species. — No new species is described in this work,

though the characteristics of many are incorporated into the descriptions of the

supraspecific taxa. I consecutively numbered (according to country or region of

origin) undescribed species within each supraspecific taxon so that information

obtained from these undescribed species could be usefully expressed herein. The

examined specimens were appropriately labelled to permit their recognition in

collections during future systematic studies of the Siphonini.

The numbering of undescribed species includes two or more higher taxa in two

groups. The first involves Ceromya s.l., in which undescribed species are numbered

consecutively according to country or region of origin regardless of their placement

in Ceromya s.s. or the Ceromya silacea species group. The second group involves

species included in New World Siphona species groups 1 and 3 and unplaced

Quaest. Ent., 1989, 25 (1,2)

12

O’Hara

species of New World Siphona s.l. : these were numbered together consecutively

from 1 to 34. [Groups 1 and 3 were recognized after the original numbering system

was adopted, so their numbers were not changed when the species were shifted to

their present classification. Siphona species group 2 was recognized before the other

groups, and includes ten species numbered from one to ten.] One species from the

above series, sp. 31, was recognized as near S. ( Uruactia ) uruhuasi and transferred

to S. ( Uruactia ) without a change in number. Due to other taxonomic changes, no

species in the Siphona s.l. series is numbered 9, 11 or 18 (i.e. these numbers are not

used to designate undescribed species).

Terms. — Terms used here are those proposed by Me Alpine (1981), with several

changes as discussed in O’Hara (1983a: 269-70). Two additional changes are made

with respect to male genitalia because of conflicting theories about genitalic

homology. Griffiths’ (1972, 1981) interpretation of homologies in male genitalia

differs from that of Me Alpine (1981), so the neutral terms pregonite and postgonite

here replace McAlpine’s controversial homology of these structures with,

respectively, the gonopod and paramere. I also use the common terms epandrium

and surstylus, though for descriptive purposes only and not in rejection of Griffiths’

periandrial hypothesis about development of these structures.

The male abdominal sternum 5 is a sclerite of varied shape among siphonines.

There do not appear to be standard terms available for detailed description of this

structure, so I have chosen arbitrary terms for this purpose, and explain their

meaning in the Structural Features chapter in characters 38 to 40. Features of this

structure are labelled in Fig. 47.

Many terms used for head features, thoracic and abdominal setation, and

genitalic structures are labelled in figs. 1-12 in O’Hara (1983a: 345-8); wing veins

are labelled here in Fig. 21, features of first instars in Figs. 157-158, and varied

external and genitalic features are labelled on scanning electron photomicrographs.

Figs. 23-46.

Examination and illustration of specimens. — Adult specimens were examined

with a Wild M5 stereoscopic microscope, with standard light source, at

magnifications to 50X. Genitalia and first instars were studied with a Leitz

SM-LUX compound microscope at magnifications to 400X. Though most genitalic

features are visible at 50X with a dissecting microscope, there are certain

taxonomically and phylogenetically important characters of the male genitalia,

particularly those of the pregonite (characters 44-46), not readily discernible unless

examined at 100X with a compound microscope or high resolution dissecting

microscope (as discussed under appropriate characters in the Structural Features

chapter).

Structures were drawn with the aid of a camera lucida or drawing tube, as

explained in O’Hara (1983a: 268-9). Certain structures were gold-coated and

examined and illustrated using a Cambridge M-250 scanning electron microscope.

Illustrations referred to in other works are cited as “fig.”, original illustrations shown

Systematics of the Genus Group Taxa of the Siphonini

13

in this work are cited as “Fig.”.

Genitalic dissections. — Male and female genitalia of siphonines provide some

of the best character systems for taxonomic study and phylogenetic analysis. The

genitalia of more than 600 siphonines were dissected during this study, following

the method of O’Hara (1983a: 271-2). Briefly stated, this procedure involved

removal of the abdomen from an adult specimen, partial clearing of the abdomen in

10% NaOH, dissection of genitalia, reattachment of abdomen to specimen, and

storage of genitalia in glycerine. The abdomen generally changed little from its

pre-dissection condition except for slight clearing and some shrivelling. Perhaps

because of these minor changes, plus the time involved in preparation, most past

workers have avoided study of siphonine genitalia despite the wealth of taxonomic

and phylogenetic information they contain.

Genitalic preparations were stored during this study in short vials of glycerine in

specially designed trays (O’Hara and McIntyre 1984), permitting direct examination

and comparison among genitalia. Each dissection was numbered such that adult

specimens and genitalia could be stored and examined separately and later easily

re-associated. Genitalia were placed in microvials and pinned below their respective

adult specimens for return of institutional loans. Dissection tags were placed in each

microvial and attached to each pin to help prevent future accidental dissociation of

genitalia and adult specimens, and to allow particular dissections referred to, or

illustrated, herein to be identified in collections and restudied.

Each dissection was color-coded and numbered. The following colors were used,

and a one letter abbreviation of each color identifies the appropriate dissection series

in the text: blue (B), green (G), orange (O), red (R), white (W) and yellow (Y).

STRUCTURAL FEATURES

Historical review

The very long geniculate proboscis of Siphona s.s. species was recognized as a

fundamental feature of that genus from the earliest writings about the Siphonini, but

it was not until publication of Mesnil’s (1954) work entitled “Genres Actia

Robineau-Desvoidy et Voisins” that the concepts of other large siphonine genera

reached their near-modem forms.

Mesnil (op. cit .) perceptively recognized the diagnostic value of two structural

features of siphonines that had been noted by previous authors but not used at the

generic level. One was Villeneuve’s (1924: 22) discovery that the anal vein

extended to the wing margin in certain species, which Mesnil used in the first

couplet of his 1954 key (pp. 8-10) to distinguish Strobliomyia ( =Peribaea ),

Ceranthia, Siphona s.s. and Siphona (Asiphona) (=S. (Aphantorhaphopsis)) from

other siphonines. This state is now considered synapotypic of that group of taxa.

Mesnil (op. cit.) also recognized the generic value of Malloch’s (1930b: 120)

discovery that a group of species possesses a long, downwardly directed, lower

proepimeral seta. Mesnil used this feature to differentiate Strobliomyia from other

Quaest. Ent., 1989, 25 (1,2)

14

O’Hara

genera in which the anal vein is extended to the wing margin, and to separate what

are now considered neaerine genera from other siphonines.

Mesnil (op. cit.) further distinguished between two groups of genera based on

basal setulation of wing vein R4+5. In one group R4+5 was setulose from base to

crossvein r-m or beyond, in the other only a single setula was present basally.

Within the former group Mesnil keyed out Goniocera and established the modem

concepts of Actia (as Entomophaga, as explained in O’Hara 1985) (based on the

presence of a row of hairs on the katepistemum), and Ceromya (based on absence of

this row of hairs). Mesnil’s latter group was later transferred to the Neaerini by

Herting (1957), based on study of female genitalia in the Calyptratae.

Mesnil synthesized the contribution of Herting (1957) into his revision of

Palearctic tachinids in “Die Fliegen der palaearktischen Region” (1962-5). In that

work Mesnil firmly established the modem concept of the Siphonini (as Siphonina),

recognizing presence of a row of setulae on R4+5 as one of the fundamental features

of the tribe. His generic classification, little changed from his 1954 publication, has

become the touchstone of modem siphonine systematics.

Characterization of the Siphonini sensu Mesnil suffered from one shortcoming,

namely the lack of a single feature which could be considered synapotypic of the

tribe. Such was the situation when this work began in 1981. By then Crosskey

(1973, 1976a, 1980) had revised the siphonine fauna of all Old World non-Palearctic

regions, but the siphonine fauna of the New World was essentially unrevised and the

applicability of Mesnil’ s tribal concept to the Neotropical fauna, in particular, was

untested. At the suggestion of D.M. Wood, Biosystematics Research Centre,

Ottawa, I investigated the possibility of two spermathecae in the female

reproductive system being synapotypic of the Siphonini. All dissections of females

during this study suggests that this state is synapotypic, and the same conclusion

was reached independently by Andersen (1983). This fundamental feature of adult

female siphonines is used here to define the world Siphonini and supplement former

external characterization of the tribe based on Old World members.

Structural features of the Siphonini

The two major revisions of the Siphonini are those of Mesnil (1962-5) and

Andersen (1983). Both are primarily based on the Palearctic fauna, and hence do not

provide detailed information about the distribution of character states throughout the

tribe. Such information is desirable not only to provide a sound phylogenetic basis

for supraspecific concepts and descriptions (and an accurate key), but also as an aid

to future revisers of siphonine taxa. Here follows an explanation of the characters

used in the descriptions of the supraspecific taxa of the Siphonini, with designation

of the states as they appear in Table 1. Table 1 is primarily intended as a quick

reference to the distribution of character states within the Siphonini, both to

summarize similarities and differences among supraspecific taxa and to augment the

phylogenetic analyses of these taxa in the Evolution chapter.

Systematics of the Genus Group Taxa of the Siphonini

15

Character 1. Length. — Adult siphonines are relatively small tachinids, ranging

in total length from 2.0-6.5mm (as measured in profile from apex of flagellomere 1

in resting position to apex of abdominal tergum 5), with most species averaging

between 3. 0-5. Omm. Like most tachinids, adult siphonines vary intraspecifically in

size, probably because of nutritional constraints placed upon them during their

parasitic larval stage. It is thus not unusual to observe a considerable size range

among conspecifics. This notwithstanding, the size of most individuals is closely

grouped about the mean of that species, such that each species has a characteristic

size. Ranges in total length given in Table 1 represent the size range observed

among examined specimens of each taxon, and are rounded to the nearest 0.5mm.

Character 2. Proclinate orbital setae. — Two proclinate orbital setae are present

on each side of the vertex of both male and female siphonines. Two states are

recognized: one in which the anterior seta is longer than the posterior one (A), the

other in which both setae are subequal in length (S). This character is not always

useful taxonomically since some species possess both states and in others males and

females differ (with the anterior seta larger in the female than in male in these

species). There is also no evident trend toward one state or the other among the

major siphonine lineages. In addition to the two states recognized here there is also

marked variation, though difficult to quantify, in length and thickness of the

proclinate orbital setae (particularly the anterior one) among siphonine lineages.

Character 3. Frontal setae. — Siphonines have a row of reclinate frontal setae

which extend from about pedicel height to the anterior reclinate orbital seta. The

frontal setae alternate short and long in all species except Proceromyia pubioculata,

in which they are all long and unusually thin (Fig. 3). Also unique to P. pubioculata

are 8-12 frontal setae, as other siphonines have no more than five (rarely six in

atypical specimens). In many siphonines the fifth (uppermost) frontal is very short,

and in a few lost altogether (often intraspecifically varied); a few species among

different lineages characteristically possess only three frontal setae, though a series

of specimens is needed to determine the stability of this state. The number of frontal

setae in each taxon is listed in Table 1, and an asterisk beside a five is used to

indicate that three or four frontals occur in some species (either characteristically or

as an aberration).

Character 4. Eye vestiture. — Siphonines, as a rule, have bare (B) eyes (i.e. only

a very few short and widely spaced hairs visible). The exception is Proceromyia

pubioculata , which has a densely haired eye (D).

Character 5. Male eye height. — Eye size is markedly varied among siphonines,

and as such is one of the most useful external characters for species recognition.

Variation among species renders this character of little value at the generic level. In

some lineages males and females of a species are apparently very similar in eye size,

in other lineages the male or female may have the larger eye; this distinction was

difficult to document with accuracy because of interspecific variation and because

both male and female specimens of a species were not always available for

Quaest. Ent., 1989, 25 (1,2)

16

O’Hara

comparison. Therefore, I have made generalizations in the generic descriptions

based on specimens available. Male eye size was selected as the standard for

comparison among genera because males generally vary more in eye size

interspecifically and possess the better characters for species identification ( i.e .

features of the male genitalia).

Overall eye size is difficult to measure, so eye height is used here for

comparisons. Eye height is taken as the maximum vertical distance from top of eye

to bottom, relative to vertical head height (excluding ocellar triangle; as in O’Hara

1983a). Some authors ( e.g . Andersen 1982) measure eye height on a diagonal axis,

resulting in slightly greater values than given here. Ranges in eye height are

recorded in Table 1 and in descriptions; qualitative differences as well are given in

the latter as an aid for comparisons, according to this arbitrary guide: small, eye up

to 0.65 head height; medium-small, 0.66-0.69; medium, 0.70-0.76; medium-large,

0.77-0.84; and large, 0.85 and greater.

Character 6. Male flagellomere 1 length. — Characteristics of flagellomere 1

(termed third antennal segment by some authors), like eye size, are among the best

external features by which to recognize species. One aspect that is easily quantified

is that of flagellomere 1 length, as measured in profile from the suture on the pedicel

to the most distant point apically, and expressed in relation to vertical head height

(as in O’Hara 1983a). Male flagellomere 1 length has been used here for

comparisons among genera because males of most species have the larger

flagellomere 1 (i.e. longer and broader) and thus exhibit more interspecific variation

in this character than females. As with eye height, a qualitative guide to

flagellomere 1 length accompanies the numerical range given in descriptions, as an

aid for comparisons among taxa. The following arbitrary length classes are used:

short, less than 0.40 head height; medium-short, 0.41-0.48; medium, 0.49-0.60;

medium-large, 0.61-0.69; and large, 0.70 and greater.

Character 7. Special modifications of male flagellomere 1. — Though the

majority of siphonines possess an average shaped (i.e. not lobed) flagellomere 1,

males (and only males) of a few species have unusual modifications, the functional

significance of which is unknown. Included in this category are: bilobed

flagellomere 1 (B; e.g. figs. 3a, 4a and 5a in Shima 1970a); trilobed flagellomere 1

(T; e.g. fig. 2a in Shima 1970a); and pectinate flagellomere 1 (P; e.g. Fig. 11, and

fig. la in Shima 1970a). Absence of such modifications is indicated by an A in

Table 1. The bilobed condition of male flagellomere 1 is found in a very few other

tachinids besides the Siphonini (e.g. some species of the neaerine genus

Phytomyptera Rondani), and a similar type of pectinate flagellomere 1 is found in

males of Borgmeiermyia Townsend (figs. 1-5 in Amaud 1963).

Character 8. Shape of male flagellomere 1 . — Considered here is the general

shape of male flagellomere 1 , excluding the extreme modifications discussed under

character 7. It is difficult to characterize flagellomere 1 shape into classes because

its shape varies from one form to another in a continuous, non-graded manner. The

Systematics of the Genus Group Taxa of the Siphonini

17

following states are broadly defined to cover the range of observed variation in

shape of male flagellomere 1, and the cited figures illustrate examples of each. The

states are: L, linear (Fig. 14); A, average (Fig. 17); B, broad (Fig. 6); S,

subquadrangular (Fig. 15); and T, large and triangular.

Character 9. Aristomere 1 length. — The majority of siphonines have a short

aristomere 1 (S), with a length half or less that of its width. Aristomere 1 is slightly

elongate (E) - subequal in length and width - in a few species of several taxa. Most

useful taxonomically and phylogenetically is state L (Figs. 4-5, 15), in which length

of aristomere 1 is much greater than width (usually 2-4 X longer than wide).

Character 10. Aristomere 2 length. — Aristomere 2 is elongate in siphonines and

at least 1.5X longer than wide, with most species having an aristomere 2 in the

range of 2-4X longer than wide. Length of aristomere 2 varies markedly within

genera, and to a lesser degree even within some species, so this character is of

limited taxonomic value above the species level. As a generalization, species with a

very large flagellomere 1 also have a long aristomere 2, so there is some correlation

in size between these structures in some lineages. Similarly, a long aristomere 2

generally accompanies a bilobed, trilobed or pectinate flagellomere 1. In Table 1 the

range in size (length divided by width) of aristomere 2 is given for each taxon.

Character 11. Aristomere 3 length. — Aristomere 3 is varied in overall length

and also in the degree to which it is tapered along its length; with length, and

thickness along length, generally correlated. For this reason I have taken a

subjective approach to this character rather than a strictly numerical one, using

states that are visually interpretable. However, the difference between one state and

another in the following series is subtle and the cited figures should be referred to as

a guide for their discrimination. The states are: V, very short and either evenly

tapered to tip or thickened to near tip (Fig. 15); T, short and thickened to near tip

(Fig. 8); S, rather short and usually evenly tapered to tip (Figs. 10, 16; a very

subjective interpretation of an aristomere 3 intermediate in form between T and the

next state); and L, long and evenly tapered to fine tip (most figures, e.g. Figs. 6, 12,

18). Clear differences in shape and length of aristomere 3 among some lineages are

useful both taxonomically and phylogenetically, even though intermediate states are

difficult to categorize.

Character 12. Arista vestiture. — Amount of pubescence on the arista is constant

within a few lineages and greatly varied in most. Lack of apparent pubescence is

referred to as almost bare (B; most figures, e.g. Figs. 14-16). Other states form a

graded series from micropubescent (M; Figs. 7, 20), pubescent (P; Figs. 10, 17),

short plumose (S; Fig. 13), to medium plumose (L, long).

Character 13. Clypeus. — The clypeus is visible as a small sclerotized structure

above the maxillary palpi when the proboscis is at least slightly extended. It is

varied in form from narrow and partially enclosed laterally in membrane (N), to

slightly (S) or distinctly broadened (B), to U-shaped (U).

Quaest. Ent., 1989, 25 (1,2)

18

O’Hara

Character 14. Maxillary palpus. — Most siphonines have a rather short, apically

clavate palpus (S; most figures, e.g. Figs. 13-15). Particularly in species with an

elongate proboscis, the palpus is correspondingly longer (L; Figs. 9, 20) and also

apically clavate (e.g. Siphona 5.5.). In S. (Ceranthia) species and two species of S.

(Aphantorhaphopsis) the palpus is cylindrical (C; Fig. 16) along its length and

varied in length from very short (reduced) to average.

Character 15. Prementum length. — Almost all species with padlike labella have

a short (S; e.g. Figs. 1-2, 5-6) to medium length (M; e.g. Figs. 14, 18) prementum

( prem ). Species with elongate labella almost always have an elongate prementum

(E; Figs. 9, 20) as well. I have not classed these states into discrete sizes, but as an

approximate guide short is less than half head height, elongate approximates or

exceeds head height, and medium is intermediate.

Character 16. Labella. — Of considerable taxonomic and phylogenetic

importance is the length of the labella (Ibl). The groundplan condition is padlike

labella (P; e.g. Figs. 1-8), and this state is widely distributed throughout the

Siphonini. A state common to many lineages but independently derived within most

of them is slightly lengthened labella (S), which are slightly longer than wide. Less

common are the following states: M (Figs. 13, 17), labella moderately lengthened

and subequal to or slightly longer than half prementum length (characteristic of most

S. (Pseudosiphona) species); E (Fig. 18), labella elongate and slightly shorter to

slightly longer than three-quarters prementum length (typical of most S.

(Siphonopsis) species); and L (Figs. 9, 20), labella as long as or longer than

prementum (all Siphona s.s. species and a few species in other lineages).

Character 17. Prosternum. — The prostemum of most siphonine species is

setulose (S), generally with a single pair of setulae but in a few species with several

pair. A bare prostemum (B) is characteristic of a few species, though more

commonly a bare specimen belongs to a species in which the prostemum is typically

setulose.

Character 18. Lower proepimeral seta. — All siphonines have a strong, upturned

seta (u prepm s) on the lower portion of the proepimeron. In addition, a strong lower

seta (/ prepm s) is present and directed downward in Peribaea (state P; Fig. 27). In

other taxa the lower proepimeral seta (prostigmatic seta of some authors) is hair-like

or absent (A; Fig. 28).

Character 19. Anepisternum. — The upper region of the anepistemum has a

patch of hairs and one to three larger setulae (anepst s), as recorded in Table 1 (one

large setula shown in- Fig. 32, two in Fig. 31). In most species the number of setulae

is apparently constant, in others varied; as well, in certain genera there is a

characteristic number of setulae while in others the number is varied.

Character 20. Katepisternum anterior to mid coxa. — The katepistemum is

considered bare (B) anterior to the mid coxa if several hairs are present along (and

restricted to) the lower portion adjacent to the ventral midline (Fig. 34). In most

Actia species and Entomophaga exoleta a row of hairs (kepst h) extend upward

Systematics of the Genus Group Taxa of the Siphonini

19

anterior to the mid coxa from the ventral midline almost to the lower katepistemal

seta (S; Fig. 33).

Character 21. Lower katepistemal seta. — Three setae are in a triangular

arrangement on the upper portion of the katepistemum. The lower seta (/ kepst 5) is

varied in length, and compared with the upper anterior seta is shorter (S; Fig. 31),

approximately equal in length (E), or distinctly longer (L; Fig. 32 [seta appearing

equal in length to upper anterior seta due to perspective]).

Character 22. Postsutural dorsocentral setae. — Three or four postsutural

dorsocentral setae are present, as indicated in Table 1. As a generalization, the setae

are longer and thicker (i.e. stronger) in species with three postsutural dorsocentrals

than in those with four. Though the states of this character are constant within some

species and lineages, they vary within a few species ( e.g . several species of Siphona

s.s. ).

Character 23. Preapical anterodorsal seta on fore tibia. — This seta is varied

from apparently absent to long. As compared with the preapical dorsal ( d) seta on

the fore tibia, it is either shorter (S) or approximately equal in length or longer (E).

Character 24. Anterodorsal seta on mid tibia. — Most siphonines possess one

strong anterodorsal seta (ad s) on the lower half of the mid tibia (Fig. 37). This seta

is lacking from S. (Baeomyia) species (Fig. 38) and several species of Actia , and a

row of ad setae is present in Goniocera. Number of ad setae present on the mid tibia

is shown in Table 1.

Character 25. Tarsomere 5 modification. — An average (A) tarsus is common to

the groundplan of the Siphonini. One to several species in separate lineages possess

a modification in tarsomere 5 of females (not known in males). Tarsomere 5 is

enlarged in these species - broader and longer than in other siphonines (e.g. fig. 22

in Richter 1980 [female mislabelled as male]) and in most species with a dense

patch of sensory haiis ventrally. This specialization is restricted to the fore leg in

most species (F), but is present on all legs in a very few (L).

Character 26. Tarsal claw length. — Claws and pulvilli are short in most

siphonines (S; e.g. figs. 4 and 6 in O’Hara 1983a). In a few species the claws are

large (L; e.g. fig. 5, op. cit.), and generally larger in males than females. M

designates a medium or intermediate-sized tarsal claw.

Character 27. Wing vein Rj dor sally. — - Setulation of vein Rx dorsally can be

discretely divided into three states: B, bare (Fig. 22); D, setulose distally; and E,

setulose along its entire length (Fig. 21). Species do not vary between states D and

E, though many species vary between B and possessing one to several setulae

distally near the costal margin.

Character 28. Wing vein R} ventrally. — Vein R j ventrally is either bare (B) or

distally setulose (D). As with character 27, some species vary between state B and

possessing one to several setulae distally near the costal margin.

Character 29. Wing vein R4+5 dorsally. — Setulation of vein R4+5 is the most

useful wing character taxonomically other than character 33. It is setulose (with four

Quaest. Ent., 1989, 25 (1,2)

20

O’Hara

to many setulae) in all siphonines and is one of the diagnostic features of the tribe.

Two states are recognized, and only a very few species vary between one state and

the other: P (proximal), R4+5 setulose between base (bifurcation of R2+ 3 and R4+5)

and crossvein r-m (Fig. 22); and E, R4+5 setulose from base to beyond crossvein r-m

(Fig. 21).

Character 30. Wing vein CuA j dorsally. — Most siphonines have either a bare

(B; Fig. 21) or partially setulose vein CuAx. This vein is either setulose from

crossvein bm-cu outward (S; Fig. 22), or in a very few species is additionally

setulose proximal to bm-cu (P). State P is constant in those species with that state,

but a few species vary between state B and possessing several setulae on CuAx (a

row of setulae on CuAx is a good species-constant state).

Character 31. Setulation of other wing veins. — Setulation described in

characters 27-30 are average for the Siphonini (A). Setulation on normally bare

veins is noted here for several species: B, Ceromya Brazil sp. 5, with veins Sc and

/?2+3 setulose ventrally; C, Actia ciligera , with veins R2+3, ^4+5 and M setulose

dorsally and ventrally; F, Actia fallax, with vein M setulose ventrally; M, Peribaea

modesta, with vein R2+ 3 setulose dorsally and ventrally; and S, some specimens of

the S. ( Actinocrocuta ) singularis complex, with vein Sc setulose ventrally.

Character 32. Loss of vein M distally. — Vein M is complete (C; i.e. extended to

wing margin) in the majority of siphonines; in some species of Actia (involving

several species groups) it fades out before its characteristic bend towards the wing

tip in the apical one-fourth of wing (N). In a few Actia species the bend of M is faint

but still visible.

Character 33. Anal vein. — This is one of the most significant characters in the

Siphonini. The anal vein either fades out before the wing margin (N; Fig. 21) or is

extended to the wing margin at least as a fold (E; Fig. 22). The latter state is rare in

other tachinids, and of great diagnostic and phylogenetic importance in the

Siphonini.

Character 34. Position of crossvein dm-cu. — The position of crossvein dm-cu is

varied from near wing margin (Fig. 21) to relatively far removed from it (Fig. 22). A

measure of dm-cu position was obtained by dividing the distal length of CuAx

(distance from dm-cu to wing margin) by proximal length of CuAx (distance from

crossvein bm-cu to dm-cu). The range in values for each taxon is given in Table 1

along with the number of species examined (generally one specimen measured per

species), the mean, and standard deviation (SD). This character is discussed in detail

in O’Hara (in press “b”).

Character 35. Median marginal setae on abdominal terga 1+2. — Median

marginal setae are considered absent (A) from abdominal terga 1+2 (Tx+2) if they

cannot be distinguished from adjacent setulae. If they are slightly longer and thicker

than adjacent setulae then they are considered weak (W), and if subequal in size to

the median and lateral marginals on f3 they are strong (S) (states A and S are

illustrated in figs. 7-8, O’Hara 1983a). Some species with state S are constant for

Systematics of the Genus Group Taxa of the Siphonini

21

this state, while in other species all states are present among different individuals or

some specimens have a strong median marginal on one side and lack one on the

other.

Character 36. Lateral marginal setae on abdominal terga 1+2. — States A

(absent), W (weak) and S (strong) are the same as described for character 35 (see

figs. 7-8 in O’Hara 1983a for states W and S). This character varies intraspecifically

as in character 35. though generally such variation is between states A and W; most

species with strong lateral marginal setae (S) rarely vary from this state.

Character 37. Setation on abdominal terga 3 to 5.- — The average condition (A)

is one in which tergum 3 (Tf) has a pair of strong median and lateral marginal setae

and tergum 4 (T4) and tergum 5 ( T5 ) each have an evenly spaced row of six marginal

setae (figs. 7-8 in O’Hara 1983a). Additionally, in some species T3-T5 have weak to

strong lateral discal setae (D) and/or an extra pair of lateral marginal setae (L).

Character 38. Posterior processes of male abdominal sternum 5. — Male

sternum 5 ( S5 ) consists of an anterior (basal) plate ( bs p) and two posterior

processes. The processes are separated from one another by a median cleft ( med c)

and bordered anteromedially by a broad desclerotized area ( descl a) of varied size

(Fig. 47). Shape of the processes is greatly varied, but two general areas are

recognized on each process: the apical lobe (ap /) and median lobe ( med 1 ) (Fig. 47).

In most Actia species the median lobes are undifferentiated and the apical lobes are

long and broadly continuous with the processes, resulting in a more or less V-shaped

S5 (A, Actia- type; Figs. 58-59). In a few other siphonines S5 resembles the A state of

Actia except the median lobes are distinctly differentiated and partially enclose the

median cleft (V; Figs. 61, 64). In most siphonines the apical lobe is more narrowed

preapically, and in combination with a distinctly differentiated median lobe

produces a sharply or broadly defined U-shaped posterior margin (U; e.g. Figs. 47,

50, 65-70) or one in which the angle between apical and median lobes is obtusely

angled (O; i.e. an intermediate state between V-shaped and U-shaped, e.g. Figs.

48-49, 60). A few siphonines have an undifferentiated apical lobe and rounded

median lobe, producing a more or less transverse sternal margin posteriorly (T; Figs.

51-52, 57).

Character 39. Apical lobe of male abdominal sternum 5. — Within lineages

possessing a preapically narrowed apical lobe and U-shaped S5 posteriorly (see

character 38), there is a trend toward curving inward (i.e. medially) of the apex of

the apical lobe. State A designates the average condition in which the apical lobe is

directed posteriorly (e.g. Figs. 47, 50, 66-68); C, an apical lobe distinctly curved

inward (Figs. 56, 62-63, 65, 69); and S, an intermediate state in which apical lobe is

slightly curved inward (Figs. 53-54).

Character 40. Median lobe of male abdominal sternum 5. — This feature of

considerable variation is here defined as the region bordering the median cleft

posteriorly, and continuous with the posterior process laterally. It is undifferentiated

(U) in those species of Actia in which the sternal margin posteriorly is V-shaped and

Quaest. Ent., 1989, 25 (1,2)

22

O’Hara

median cleft not bordered by lobes (Fig. 58). In the simplest developed form the

median lobe is rounded along its median margin (R) and partially encloses the

median cleft (e.g. Figs. 55, 64, 67). In more developed forms the median lobe is

truncate (T; e.g. Figs. 63, 66) along its medial or posteromedial margin or very

broad and longitudinally elongate (E; e.g. Figs. 53, 69). In Goniocera species and a

few other species the median lobe forms a broad, truncated plate flattened in a more

or less posteromedial plane (F; Figs. 47, 54). State F is a highly derived form of

median lobe, and between it and state T are a range of intermediates here classed as

state I, characterized by a truncate margin and rather flattened surface

posteromedially (Fig. 68). Some species, particularly within Siphona s.l., have an

accessory lobe on the median lobe posteriorly (A; Fig. 70).

Character 41. Setulation of male abdominal sternum 5. — Sternum 5 is bare on

the basal plate, anterior to the desclerotized area. The posterior processes are

sparsely (S), moderately (M), or densely (D) setulose. The processes may also have

one pair of setae distinctly larger than the surrounding setulae, but presence or

absence of these setae has been interpreted as too intraspecifically varied to be

usefully characterized here.

Character 42. Size of male abdominal ter gum 6. — Male tergum 6 (T6) is

separated from the epandrium by a narrow strip of membrane. In a few siphonines

T6 is unsclerotized (A, apparently absent), while in most it is varied from tiny to

small lateral sclerites (S), to larger lateral sclerites (L), to a narrow to broad dorsally

continuous sclerite (D).

Character 43. Width of ejaculatory apodeme. — The ejaculatory apodeme in the

male genitalia has a bulbous base and a fan-shaped apical portion of varied size.

Width of the fan-shaped portion is expressed herein as a function of the preapical

width of the hypandrial apodeme (as measured dorsally). The relative width of the

fan-shaped portion is given either as a measured range or expressed in relative terms

in Table 1 and descriptions. The following states of the latter are recognized: N,

ejaculatory apodeme slightly narrower than hypandrial apodeme; S, apodemes

subequal in width; and W, ejaculatory apodeme slightly wider than hypandrial

apodeme.

Character 44. Shape of pregonite. — Pregonites are structures to either side of

the aedeagus, articulated at their bases with the hypandrium. A thin membrane

extends anteriorly from the apex of each pregonite to the ventral surface of

hypandrium. Pregonite shape is varied in many and subtle ways and cannot be

classified into states without grouping similar forms in a subjective manner. I have

tried to select and describe states I perceive as most meaningful in a taxonomic and

phylogenetic sense, though it must be noted that even a continuous transformation

series cannot be hypothesized since most states could easily have been derived from

one of several others. The following states are recognized, and are arranged

alphabetically because of the complexity of this character: A, average, smoothly

curved along posterior margin or bent rather sharply at midlength, and pointed or

Systematics of the Genus Group Taxa of the Siphonini

23

rounded apically (Figs. 73-76, 89-94, 96-106); C, C-shaped (open side facing

anteriorly) and broad at midlength (Figs. 87-88); F, fused basally with hypandrial

apodeme (only observed in S. ( Aphantorhaphopsis ) nigronitensf J, broad

subapically and more or less J-shaped (Figs. 85-86); L, linear and only slightly

curved apically (Fig. 72); M, anterior membranous portion enlarged, apex of

pregonite generally not curved anteriorly (Figs. 77, 79-81); O, ring-shaped (Fig. 78),

though part of ring incomplete in a few species; P, elongate and curved posteriorly

(unique to Ceromya lutea. Fig. 83); S, thin and sickle-like, with basal arm slightly

extended anteriorly (Fig. 84); T, basally broad and apically truncate (Fig. 71); and

W, broadened or wide along most of length, with short pointed apex (Fig. 95).

Character 45. Pregonite modifications. — The pregonite has a smooth outer

surface in the groundplan condition. Spines (sp) or spinules (spin) adorn the

pregonite in a number of genera, and the size, pattern and location of these are

classed into discrete, non-continuous states. However, not all of the following states

can be seen with a dissecting microscope (at 50X), so it is necessary to examine the

pregonite at a power of 100X or more with a compound microscope to discern with

certainty whether small spinules are present or absent (A; e.g. Figs. 72, 76, 84,

87-88, 90-94). The spinose states are: L, longitudinal ridge on anterolateral surface

with spines in single row along edge (Figs. 73-75, 103-105); M, anterior

membranous portion ( memb p) of pregonite expanded, usually with field of spinules

on lower portion (Figs. 39-40, 71, 83, 79-81; these spinules are too small to be seen

with most dissecting microscopes in specimens of most species); E, similar to state

M, but in addition has laterally directed spines on anterolateral portion of sclerotized

area (Fig. 82); R, ring-shaped pregonite (state O in character 44) with spinules

anteriorly (Fig. 78); S, sclerotized portion ( scl p) of pregonite with field of spines

distally, with largest spines along apical margin (Figs. 41-42, 85-86); and T,

sclerotized portion of pregonite with field of tiny spinules distally (Figs. 89, 95, 100;

as with state M, these spinules are very small in specimens of some species, and not

readily discernible with most dissecting microscopes).

Character 46. Posterior seta on pregonite. — Some members of Siphona s.l.

possess a tiny to long seta at about midlength on the posterior surface of the

pregonite. Different setal size classes are recognized because certain lineages

possess a seta with a characteristic length. However, setal length is varied among

some species, and varied between absent and small to medium in a few. Given the

varied nature of this character, the following imprecisely specified states are

recognized: A, absent (e.g. Figs. 71-82); T, very tiny seta (Figs. 89, 93, 97-98,

103-104, 106; not readily discernible without aid of a compound microscope); M,

short to medium-sized seta (Figs. 90-92); L, long, thick seta (Fig. 94; as

characteristically present in species of S. ( Ceranthia )); and F, two to a few tiny to

short setae (Fig. 95).

Character 47. Epiphallus. — An epiphallus is a subbasal, posterior outgrowth of

the aedeagus. A small epiphallus is best seen in dorsal or ventral view, where

Quaest.Ent., 1989, 25 (1,2)

24

O’Hara

unobstructed by epandrium or postgonite. The following states are recognized: A,

absent; S, small (short); N, relatively large but narrow; and P (present), relatively

large and average width.

Character 48. Posterior margin of distiphallus. — The distiphallus is a

cylindrical structure of varied size, shape and sclerotization. Several characteristics

of the distiphallus appear to be more or less independent of one another, and three

are considered here as characters 48 to 50. Character 48 describes the relative

amount of sclerotization along the posterior margin of the distiphallus. The posterior

margin is either entirely sclerotized to near apical margin of distiphallus (E; pos

marg in Fig. 1 16), or is incised or desclerotized to varied degrees (I). The latter state

can be recognized in lateral view in most species of Siphona s.l. by the presence of a

pointed, sclerotized projection on the distiphallus posteriorly, below the narrowed

attachment of basiphallus and distiphallus ( e.g . scl proj in Figs. 125, 127).

Character 49. Posterolateral margin of distiphallus. — The lateral surfaces of the

distiphallus, as viewed in profile, provide among the best features for species

separation and recognition within the Siphonini. Shape of the distiphallus is too

varied to be fully categorized at the generic level, but two aspects have been

selected for description as characters 49 and 50. Character 49 refers to the

posterolateral margin (apically) of the distiphallus. In some siphonines it is reduced

(r lat marg in Fig. 108) or not differentiated ( i.e . not incised; undif p slat marg in

Fig. 124) from the rest of the distiphallus (A; e.g. Figs. 108-111, 1 15-124). In other

siphonines the posterolateral margin of the distiphallus is incised from the rest of the

lateral margin and slightly to markedly elongate, mostly independently among

different lineages. This is a difficult character to classify into states, though the

following series gives a rough indication of the diversity of this character among the

recognized siphonine lineages: S, posterolateral margin separated from lateral

margin by narrow incision and not extended beyond apex of lateral margin (present

in Goniocera io, but not visible in Fig. 107 because posterolateral margins are

curved inward); M, moderate apical extension of posterolateral margin beyond rest

of lateral margin (m pslat marg in Fig. 125) and usually pointed or spined (either

continuous with lateral margin or incised from it; Figs. 125, 131, 134); and E, as in

state M but apically extended well beyond apex of rest of distiphallus (/ pslat marg

in Figs. 1 12 and 127) (Figs. 1 12, 1 14, 127, 133, 136).

Character 50. Spines on lateral margin of distiphallus. — The lateral and

anterolateral surfaces of the distiphallus are adorned with small spinules, especially

conspicuous anteriorly and apically. In some siphonines no larger spines are present

(A; Figs. 107, 115-118, 132-135), in others the lateral margins are armed with

conspicuously larger, usually recurved, spines along anterior and/or apical margin

(P; Figs. 108-111, 114, 121, 124, 128). The degree to which these spines are

differentiated is very useful taxonomically, particularly at the species level, but is

too varied to be categorized here.

Systematics of the Genus Group Taxa of the Siphonini

25

Character 51. Postgonite size. — Paired postgonites extend posteriorly from

either side of the aedeagus basally. They are large (W, well developed) and

conspicuous in most siphonines. Postgonites are short (S) in some siphonines (about

as long as width of basiphallus subbasally), or reduced (R) to scarcely discernible

lobes at base of basiphallus.

Character 52. Sur stylus basally.- — Each surstylus articulates basally with the

epandrium and cerci. In most siphonines it is attached membranously (M; Figs.

139-144), in a few there is a narrow sclerotized connection between the epandrium

posteriorly and surstylus basally (F, epandrium and surstylus fused; Figs. 137-138).

Character 53. Shape of surstylus. — The average condition (A) of the surstylus,

as viewed in profile, is one in which it is narrowed at about midlength and either

straight or slightly curved posteriorly near apex (Fig. 141). The following states are

qualitative departures from the average condition (see cited figures for examples): S,

short (Fig. 142); B, distinctly broader than average (Fig. 138); C, markedly curved

posteriorly (Fig. 139); and L, elongate (Fig. 137).

Character 54. Shape of male cerci. — The cerci of siphonines are fused medially

and tapered to a single slender apex. Their shape is described in lateral view, with

average (A) being slender on lower half, smoothly curved at midlength and curved

anteriorly near apex (Fig. 141). The following shapes are recognized, relative to the

average shape: S, short (Fig. 138); B, broadened; E, elongate (Fig. 140); L, linear or

straight along posterior margin in profile; R, same as state L but short (Fig. 139);

and I, sharply inflexed at midlength, short to long in length (Figs. 137-138, 140;

state I may be accompanied by one of the former states).

Character 55. Female ovipositor length. — The ovipositor, or female genitalia, is

primitively short and only slightly extensible. More extensible ovipositors have

evolved independently within different lineages, perhaps as an adaptation for

depositing larvae directly upon hosts (as suggested by Andersen 1983). Extensible

ovipositors are telescopic and characterized by broad regions of intersegmental

membrane, and often elongated sclerites as well. Degree of extensibility is classed

as follows: S, short and only slightly extensible (Figs. 43-46, 146-148, 151-153); M,

moderately extensible (Figs. 145, 149-150, 154); E, elongate (Fig. 155); and V, very

elongate (Fig. 156).

Character 56. Shape of female sternum 6. — The most apparent modifications to

the female genitalia, other than extensibility, involve shape of sterna 6 (S6) and 7

(S-j). Modification of S6 is not common among siphonines, and is apparently always

accompanied by similar or more pronounced modification of S7. In the average (A)

condition S6 is slightly rounded on its ventral surface (Figs. 43-46, 145-153).

Several types of modification are recognized: W, weakly or slightly keeled ( kl )

posteromedially; S, sharply keeled posteromedially (Fig. 154); M, moderately

elongate (Figs. 154, 156); and E, elongate (Fig. 155).

Character 57. Size of female tergum 6. — Female tergum 6 ( T6 ) is varied in size

as follows: A, absent; S, very small lateral sclerites (Fig. 152); L, moderate-sized

Quaest. Ent., 1989, 25 (1,2)

26

O’Hara

lateral sclerites (Figs. 145, 148, 151, 155-156); M, dorsomedian sclerite, not

extended laterally; D, single sclerite extended dorsally and laterally (Figs. 153-154);

and N, as in state D but narrowly discontinuous dorsally (Figs. 146-147, 149-150).

Character 58. Anterior apodeme on female sternum 7. — Female sternum 7 (S7)

is without an anterior apodeme (A, absent) if its anterior margin is broadly rounded

(Figs. 145, 149, 151-156). An apodeme is present if a sclerotized arm extends

anteriorly from the anteromedian margin of S7, and is classed as either short (S; Fig.

150) or long (W, well developed; Figs. 146-148, ant apod in Fig. 147).

Character 59. Shape of female sternum 7. — As mentioned under character 56,

modifications to sterna 6 (S6) and 7 (S7) are the most conspicuous in female genitalia

other than extensibility. The following states are similar to those given for character

56: A, average condition, S7 slightly rounded ventrally (Figs. 43, 45, 145-146,

148-153); W, weakly or slightly keeled ( kl ) posteromedially (Fig. 147); S, sharply

keeled posteromedially (Figs. 44, 46); E, elongate (Figs. 155-156); and L, elongate

and keeled posteromedially (Fig. 154).

Character 60. Size of female ter gum 7. — Female tergum 7 (T7) is varied in size

as follows: A, absent; S, very small lateral sclerites (Figs. 151-152); L,

moderate-sized lateral sclerites (Figs. 145-150, 153); and F, lateral sclerites fused

with S7 (Figs. 154-156).

Character 61. Location of spiracle of segment 7. — The position of the spiracle

of segment 7 is varied. In some siphonines it is located in segment 7, close to or in

T7 or dorsal to margin of S7 (state P, posterior; Figs. 148, 153-156). In other

siphonines the spiracle is located dorsolaterally in membrane between segments 6

and 7 (state I, intermediate; Figs. 145, 150-152) or anteriorly in segment 6 in or near

T6 (state A, anterior; Figs. 146-147, 149).

Character 62. Characteristics of female sternum 8. — Female sternum 8 (S8) is

varied in several ways. In the average (A) condition it is a small sclerotized plate

partially covered by S7, and sparsely haired posteriorly (Figs. 43-46, 146, 148-153;

average condition but bare in Peribaea species). Other states are as follows: L,

lacking or absent (Figs. 147, 154-156); R, reduced in size (Fig. 145); W, much

wider than average; B, bare or almost so; and S, with thick setae posteriorly

(setation not shown in figures).

Character 63. Size of female tergum 10. — Tergum 10 (Tl0) is located dorsal to

the cerci ( cer ). It is small and sparsely haired in most siphonines, and present as

either a median sclerite (M) or as two slightly separated sclerites (P, paired; a few

species possess both states). In a few siphonines T10 is unsclerotized (A, absent) or

reduced in size (R).

Character 64. Labrum of first instar. — The anterior end of the

cephalopharyngeal skeleton of first instars of the Tachinidae is extended forward

between the lateral sclerites, and is tentatively homologized with the labrum by

Wood (1987: 1196) (often termed “mouth hook” by previous authors). As in other

tachinids, the labrum of first instar siphonines varies from hook-like (N, narrow in

Systematics of the Genus Group Taxa of the Siphonini

27

dorsal-ventral plane; Figs. 159, 161-162) to hatchet-like (B, broadened in

dorsal-ventral plane; Figs. 158, 160, 163). In a few siphonines the labrum is

intermediate between these states, and here termed slightly broadened (I). First

instars of the Siphonini are described by O’Hara (in press “a”).

Character 65. Dorsal cornu of cephalopharyngeal skeleton of first instars. — In

most larval siphonines and other larval tachinids there are two cornua posteriorly on

the cephalopharyngeal skeleton, one dorsal ( d corn) and one ventral (v corn ) (P;

Figs. 158-159, 161-163). The dorsal cornu is absent (A) from all nine Actia species

examined for this character (Figs. 160). The possible presence of a dorsal cornu in

Actia dubitata, illustrated in Farinets (1980), is indicated by a “7P1” in Table 1.

Character 66. Posteroventral margin of abdominal segment 6 of first instars. —

First instars of the Siphonini are equipped with spinules or hooks externally. The

greatest modification and variation in these structures are found on the last several

abdominal segments, particularly the posteroventral margin of segments 6 and 7.

Segment 6 is bare or has small spinules (B) posteroventrally (Figs. 157-161), or

large spinules to hooks. If large spinules or hooks are present (Figs. 162-163), then

their number is entered in Table 1 .

Character 67. Posteroventral margin of abdominal segment 7 of first instars. —

Segment 7 possesses ventrally a row or rows of spinules along its posterior margin,

anterior to the anus. Two states are recognized: R, two or more even or uneven rows

of spinules (Figs. 157-162); and S, a single dominant row of spinules (Fig. 163).

Quae st. Ent., 1989, 25 (1,2)

Table 1. Distribution of character states in the Siphonini. Refer to text for explanation of characters and states. (Superscripts indicate

the number of species possessing a state, an asterisk denotes a state shared by most species in a taxon, an arrow signifies the presence of

states intermediate to those on either side of arrow, and a diagonal slash (/) indicates that two states are present in the single examined

species of that taxon.)

28

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Taxon shape shape length shape setation seta setation mid coxa setae dorsocentrals fore tibia mid tibia

Systematics of the Genus Group Taxa of the Siphonini

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Table 1 (cont.). Distribution of character states in the Siphonini

Systematics of the Genus Group Taxa of the Siphonini

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Quaest. Ent., 1989, 25 (1,2)

Epiphallus Male distiphallus Postgonite Male surstylus Cerci Female Female

Taxon size posterior posterolateral lateral shape basally shape shape ovipositor sternum 6

32

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Female Female sternum 7 Female Segment 7 Female Female 1st instar Dorsal Posteroventral margin

Taxon tergum 6 apodeme shape tergum 7 spiracle sternum 8 tergum 10 labrum cornu abd. seg. 6 abd. seg.

Systematics of the Genus Group Taxa of the Siphonini

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34

O’Hara

CLASSIFICATION OF THE SIPHONINI

List of major references to the Siphonini

Thryptoceratae Robineau-Desvoidy 1830: 82-102 (unavailable name).

Siphonae Rondani 1845: 31 (original description).

Siphonina Rondani 1856: 59 ( Siphona ; other siphonine genera in Tachinina).

Siphoninae Rondani 1859: 9-11 (Italian species of Siphona; other siphonines in Tachininae).

— [Muscidae], Schiner 1862: 517-522 (early classification; species of Austria).

Ceromydes and Thryptocerides, Robineau-Desvoidy 1863: 665-728 (early classification).

Thryptoceratidae, Brauer and Bergenstamm 1889: 101-106 (early classification).

Section Thryptocera , Brauer and Bergenstamm 1893: 148-152 (early classification).

— , Bezzi and Stein 1907: 381-392 (early Palearctic catalogue).

Section Thryptocera, Villeneuve 1924: 34 (Palearctic reclassification).

Group Thryptocera, Stein 1924: 123-148 (reclassification and revision of mid-European species).

— [Eutachininae, Group 3], Lundbeck 1927: 448-476 (species of Denmark).

— [Tachininae], Wainwright 1928: 203-209 (species of Britain).

Actiini, Malloch 1930a: 303-310 (species of Australia, all as Actia).

Actiini, Malloch 1930b: 120-148 (species of Malaysia, all as Actia).

Actiini and Siphonini, Townsend 1936: 129-152 (tribal descriptions and generic keys; very broad tribal

limits).

Crocutini, Mesnil 1939: 35-36 (reclassification).

Actiini and Siphonini, Townsend 1940: 187-299 (generic descriptions following 1936 classification).

Siphonini, Emden [van Emden] 1954: 62-68 (species of Britain).

Siphoninae, Mesnil 1954, 41 pp. (reclassification and species of “Congo Beige”).

Siphona group, Herting 1957: 454, 458 (female genitalia).

Siphonini, Herting 1960: 58-63 (biology of Palearctic species).

Siphonina, Mesnil 1962-1965: 795-879 (reclassification and revision of Palearctic species).

Siphonina, Sabrosky and Amaud 1965: 1061-1064 (list of New World species north of Mexico).

Siphonini, Mesnil and Pschom-Walcher 1968: 163-164 (list of species of Japan).

Siphonini, Guimaraes 1971: 164-171 (list of New World species south of United States; broad tribal

concept).

Siphonini, Crosskey 1973: 80-81, 136-138, 176 (species of Australia).

Siphonini, Mesnil 1975: 1399-1400 (changes to 1962-1965 Palearctic revision).

Siphonini, Crosskey 1976a: 1 12-113, 21 1-214, 291 (species of Oriental region).

Siphonini, Crosskey 1976b: 99-100 (list of British species).

Siphonini, Crosskey 1980: 852-855 (list of Afrotropical species).

Siphonini, Dear and Crosskey 1982: 134-139 (species of the Philippines).

Siphonini, Andersen 1983: 1-15 (revision of Old World genera).

Siphonini, Crosskey 1984: 261-262 (key to Afrotropical genera).

Siphonini, Herting 1984: 120-126 (list of Palearctic species).

Siphonini, Tschorsnig 1985: 88-89 (male genitalia).

Siphonini, Rognes 1986: 72-73 (list of Norwegian species).

Diagnosis of adults of Siphonini Rondani

The most convincing synapotypy, and hence best diagnostic feature, of the

Siphonini is the presence of only two (rather than three) spermathecae in the female

reproductive system.' This state seems to be universal among siphonines, and is

known to be shared with only one other (unrelated) tachinid genus, Phaenopsis

Townsend (Andersen 1983).

The following combination of character states separates adults of this tribe from

other tachinids: Generally 3. 0-5. 0mm in length, but varied from 2.0-6.5mm. Head

(Figs. 1-20) with little sexual dimorphism. Two proclinate orbital setae in both

sexes. Eye bare or almost so and three to five frontal setae except in Proceromyia

Systematics of the Genus Group Taxa of the Siphonini

35

pubioculata (eye densely haired and 8-12 frontals; Fig. 3). Aristomere 2 longer than

wide, elongate in most species. Proboscis varied from short to long, labella padlike

to more than 2X head height.

Thoracic dorsum illustrated in O’Hara (1983a, fig. 2). Prostemum with one to

several pair of setulae in most species, bare in a few (intraspecifically varied in a

few species). Katepistemum with three setae arranged in triangle (Figs. 31-32),

lower seta varied from hairlike to slightly longer than upper anterior seta. Row of

katepistemal hairs anterior to mid coxa present (Fig. 33) or absent (Fig. 34). Three

or four postsutural dorsocentral setae. Scutellum with strong basal, lateral and

subapical setae, subapicals longest and convergent to crossed, apicals short in most

species. One ad seta on mid tibia (Fig. 37) in most species, seta absent from a few

species (Fig. 38) and row of setae present in Goniocera species. Wing relatively

short and broad (Figs. 21-22), vein Rx partially incised at midpoint, vein R4+5

setulose dorsally from bifurcation of R2+ 3 and R4+5 to crossvein r-m or beyond, vein

m smoothly curved in apical sector and ending in wing margin near wing tip (ending

close to end of /?4+5), anal vein extended to or not extended to wing margin.

Abdomen (O’Hara 1983a, figs. 7-8) ovoid to elongate, with T1+2 excavate on

basal half. Median discal setae absent from Tl+2 to ^4- ^1+2 with 0-1 pair of lateral

marginal setae in most species, additional pair present in a few species, pair of

median marginal setae present in a few species of Goniocera and Siphona

(Siphona). T3 to T4 with strong pair each of lateral and median marginal setae,

additional marginals present in a few species. T5 with row of marginal setae and in a

few species with row of weak discal setae.

Male genitalia (O’Hara 1983a, fig. 9) with tergum 6 unsclerotized to moderately

broad and sclerotized. Terga 7+8 well developed. Epandrium saddle-shaped.

Pregonite (Figs. 39-42, 71-106) well developed, with membrane anteriorly between

apex and basal articulation with hypandrium (possibly synapotypic of the Siphonini,

as suggested by Tschorsnig 1985 and discussed in Evolution chapter). Ejaculatory

apodeme well developed. Aedeagus with or without an epiphallus. Distiphallus

(Figs. 107-136) narrowly connected to basiphallus, in most species approximately

cylindrical in shape and well sclerotized laterally. Postgonite well developed to

markedly reduced. Surstylus (Figs. 137-141) linear, relatively unmodified, and

basally free or connected with epandrium by a narrow strap-like connection. Cerci

(Figs. 142-144) medially incised basally, fused apically.

Female reproductive system with two spermathecae (synapotypic of the

Siphonini). Female genitalia (Figs. 43-46, 145-156) unmodified to long and

extensible (telescopic), without piercing ability. Tergum 6 absent to dorsally

continuous. Tergum 7 absent to present as two moderate-sized lateral sclerites.

Spiracle of segment 6 associated with ventrolateral margin of tergum 6, spiracle of

segment 7 varied in position from segment 7 to segment 6. Tergum 8 represented by

two lateral sclerites, sternum 8 relatively narrow in most species, absent from a few.

Segment 9 without external sclerites (O’Hara 1983c: 380). Tergum 10 unsclerotized

Quaest. Ent., 1989, 25 (1,2)

36

O’Hara

or represented by small median sclerite or two small sclerites. Sternum 10 and

paired cerci unmodified.

Review of major keys to genera and subgenera of the Siphonini

Keys to genera and subgenera of the Siphonini are available for each of the

biogeographic regions of the world except the Neotropics. Each key reflects a

classification different from the one adopted here, so each also differs from the keys

presented in this work: i.e. key to the genera of the Siphonini, keys to the species of

Proceromyia and Entomophaga , and key to the subgenera of Siphona s.l. To avoid

confusion between those keys and mine, the major differences are summarized here.

Specific information about differences in the placement of species can be obtained

by comparing the species lists accompanying most of the works cited below with the

descriptions and species lists contained herein. Differences in phylogenetic

interpretation are not discussed in this section, but rather in the Phylogenetics

section of each supraspecific taxon described. Recent keys to species are cited in the

Geographic Distribution section of each supraspecific taxon.

The keys considered here are those published in major works since Mesnil’s

revision of the Palearctic Siphonini in “Die Fliegen der palaearktischen Region”

(1962-1965). The most geographically comprehensive is Andersen’s (1983) generic

revision of the Old World Siphonini. This work is broad in scope and contains

important new findings about the Siphonini. Andersen’s key (pp. 13-14) is easy to

use and his classification is similar to the one presented herein. His key differs from

mine in the following respects: the two species of Entomophaga are split between

Actia and Ceromya, Proceromyia macronychia is included in Ceromya,

Proceromyia pubioculata (type species of Nipponoceromyia) was not examined and

hence excluded, Actia species with atypical features key to Ceromya or Asiphona,

taxa here considered subgenera of Siphona s.l. are considered genera, Siphona

(Aphantorhaphopsis) is called Asiphona, a few species of S. (Aphantorhaphopsis)

key to Ceranthia , and S. (Aphantorhaphopsis) and S. (Siphona) [his Asiphona and

Siphona, respectively] not adequately differentiated. Most of these differences stem

from the relatively few species examined by Andersen during his study. Most of the

species which key differently have limited known ranges (mostly non-Palearctic)

and/or are rarely collected, so despite the differences in our keys probably 90% of

Old World siphonine species will key to the same taxon. Because my keys attempt

to provide for accurate generic placement of all species examined (see Materials and

Methods), they are necessarily longer and more complex than Andersen’s.

Other keys to Old World siphonine genera and subgenera are regional in

coverage. The most comprehensive is Mesnil’s Palearctic revision cited above

(1962-1965), in which all the then known species of Palearctic Siphonini are keyed

and described. Mesnil’s revision restructured the concept of the Siphonini into its

modem form and still provides the only means for identification of most species of

Palearctic Siphonini. With respect to the Palearctic fauna, Mesnil’s keys to the

Systematics of the Genus Group Taxa of the Siphonini

37

Siphonini (1962: 797) and subgenera of Ceromya (1963a: 829) differ from mine as

follows: Peribaea called Strobliomyia, Siphona (Aphantorhaphopsis) called Siphona

(Asiphona), Ceranthia accorded generic rank, Proceromyia as Ceromya

(Proceromyia), Entomophaga exoleta in Actia, Ceromya monstrosicornis as type

species of Ceromya ( Stenoparia ) and Entomophaga nigrohalterata as C.

(Stenoparia) nigrohalterata. Nipponoceromyia (here synonymized with

Proceromyia ) was described after Mesnil’s revision. Atypical species of Actia and

S. ( Aphantorhaphopsis ) are non-Palearctic in distribution.

The remaining Old World regions have received recent treatment by Crosskey.

These include a conspectus of Australian Tachinidae (1973), conspectus of Oriental

Tachinidae (1976), review of the Tachinidae of the Philippines (Dear and Crosskey

1982), catalogue of Afrotropical Tachinidae (1980, including Madagascar) and key

to genera of Afrotropical Tachinidae (1984, excluding Madagascar). Excluded from

these works is a treatment of the Tachinidae of the Australian region outside

Australia. These works follow the same classification of the Siphonini, so their keys

are here compared as one with the keys herein: atypical Actia species key to

Ceromya , Siphona s.s. not then recorded from Australia, S. ( Aphantorhaphopsis )

generally not recognized so most of its species are placed in Ceromya and a few

(those with elongate labella) in Siphona s.s., and ranking of subgenera of Siphona

s.l. as genera. These differences in classification are mitigated by Crosskey’s

comprehensive species lists for each region, without which the initial stages of

almost any revision of a non-Palearctic tachinid group would be an arduous task.

Wood’s (1987) recent key to tachinid genera of the Nearctic region provides a

valuable and much needed guide to the identification of Nearctic Tachinidae. Wood

separates tachinid genera in the first couplet of his key according to presence or

absence of setulae on the prostemum. Goniocera io (the only New World species of

the genus) is varied for this character, though it keys as having a bare prostemum.

Other siphonines key as having a setulose prostemum, though a few of these species

are also intraspecifically varied for this character (especially species of Siphona

( Baeomyia )). Wood’s Siphona and Baeomyia are equivalent to my Siphona

( Siphona ) and S. (Baeomyia), and his Ceranthia includes all Nearctic Siphona s.l.

species exclusive of the last two subgenera.

Key to adults of the genera of the Siphonini

1 Lower proepimeral seta strong and directed downward (Fig. 27)

(Old World) Peribaea R.-D., p. 77

T Lower proepimeral seta absent or hair-like (Fig. 28) 2

2 (T) Anal vein not extended to wing margin (Fig. 21); lower

katepistemal seta shorter than upper anterior one (Fig. 31) 3

2' Anal vein extended to wing margin at least as fold (Fig. 22);

lower katepistemal seta subequal to or longer than upper

Quaest. Ent., 1989, 25 (1,2)

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O’Hara

anterior one in most species (Fig. 32) 10

3 (2) Mid tibia with row of three or more ad setae and setulae (four

species; Europe and northeastern North America)

Goniocera B. & B., p. 41

3' Mid tibia with one ad seta in most species (Fig. 37), seta

reduced or absent in a very few (Fig. 38) 4

4 (3') Aristomere 1 distinctly longer than wide (Figs. 4-5) and fore

tibia with preapical ad seta as long as or longer than d seta (two

European species) Entomophaga Lioy, p. 47

4' Most species with aristomere 1 distinctly shorter than wide and

fore tibia with preapical ad seta shorter than d seta, a few

species with one (but not both) of above states 5

5 (4') Katepistemum with row of hairs directly anterior to mid coxa,

extended upward almost to lower katepistemal seta (Fig. 33)

(cosmopolitan) most species of Actia R.-D., p. 67

5' Katepistemum almost bare directly anterior to mid coxa, except

for several hairs in posteroventral comer (Fig. 34) 6

6 (5') Male sternum 5 with posterior margin approximately V-shaped

(Figs. 58-59); pregonite with outer, sclerotized portion short

spinose on apical half or less (Figs. 41-42, 85-86); upper part of

anepistemum with two setulae (Fig. 31) (presently known only

from Malaysia and eastern Australia)

several species of Actia R.-D., p. 67

6' Male sternum 5 with posterior margin U-shaped, obtusely

angled or almost transverse (Figs. 48-49, 52-57); pregonite bare

or with tiny to distinct, uniformly-sized spinules (Figs. 39-40,

73-74, 77-84) (spinules, if present, restricted to membranous

portion of pregonite except in several species with extensively

sclerotized, ring-shaped pregonite, Fig. 78); upper part of

anepistemum with one setula in most species (Fig. 32) 7

7 (6') Dorsal surface of wing vein R4+5 setulose from base to beyond

crossvein r-m (as in Fig. 21) (cosmopolitan)

most species of Ceromya R.-D. sensu lato 8

T Dorsal surface of wing vein /?4+5 setulose between base and

crossvein r-m (as in Fig. 22) (Old World) 9

8 (7) Distiphallus bearing unique, infolded and sclerotized structure

formed from posterior surface (Fig. 112); pregonite bare (Fig.

84) (Old World) Ceromya silacea (Mg.) species group, p. 63

8' Distiphallus without infolded and sclerotized structure (Figs.

113-116); pregonite bare in a few species (Fig. 77), with tiny to

distinct spinules in most species (Figs. 78-83) (spinules, if

present, restricted to membranous portion of pregonite except

Systematics of the Genus Group Taxa of the Siphonini

39

in several species with extensively sclerotized, ring-shaped

pregonite - Fig. 78); (cosmopolitan)

Ceromya R.-D. sensu stricto, p. 52

9 (7') Fore tibia with preapical ad seta as long as or longer than d

seta; male genitalia with distinctive sternum 5 (Figs. 48-49),

pregonite (Figs. 73-74) and distiphallus (Figs.

108-109) Proceromyia Mesnil, p. 44

9' Fore tibia with preapical ad seta shorter than d seta; male

genitalia not as above in C. cephalotes and C. natalensis, not

examined in C. monstrosicornis (these species discussed in

Phylogenetics section of Ceromya s.s .)

three known species of Ceromya R.-D. sensu stricto , p. 52

10 (2') Katepistemum with row of hairs directly anterior to mid coxa,

extended upward halfway to lower katepistemal seta (type

specimen of A. completa from Malaysia) or almost to lower

katepistemal seta (as in Fig. 33; A. fulvicauda from Malaysia

and A. chrysocera from the Seychelles Islands)

three known species of Actia R.-D., p. 67

10' Katepistemum almost bare anterior to mid coxa, except for

several hairs in posteroventral comer (Fig. 34)

Siphona Meigen sensu lato, p. 84

[Subgenera keyed in section on Siphona s.l. ]

Notes about classification chapter

Notes about lists of included species. — The present classification of the

Siphonini includes 386 named species (excluding nomina nuda and misspellings), of

which 294 are recognized as taxonomically valid species. Each description of a

supraspecific taxon of the Siphonini is followed by a list of included species. Names

of species treated as valid are preceded by a letter indicating the region of

occurrence of the species, and all named species are followed by full bibliographic

and type information (sex, type locality and depository, and whether type

examined). Synonyms are listed in chronological order according to date of

description. The following letters are used to denote region(s) of occurrence of each

species:

A Afrotropical, sensu Crosskey and White (1977).

N Nearctic, sensu Griffiths (1980).

O Oriental, sensu Crosskey (1976a).

P Palearctic, as delimited by the Afrotropical and Oriental regions.

S Neotropical, sensu Griffiths (1980).

U Australian.

Quaest. Ent., 1989, 25 (1,2)

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O’Hara

Lists of included species were originally compiled from regional catalogues by

Sabrosky and Amaud (1965), Guimaraes (1971), Crosskey (1973, 1976a, 1980) and

Herting (1984). This information was then checked and where necessary augmented,

many of the types personally examined, and the species and genera reclassified into

the present scheme. Changes to previous classifications are listed in the next section

and are indicated in the lists in bold face. In general, I avoided decisions about

species synonymies and followed current placements. This is particularly true of the

older Palearctic names that have been authoritatively re-evaluated in the works of

Herting (1969-1984). For each listed synonym I have included a recent reference to

its synonymic status rather than attempt to determine the author who first proposed

the synonymy. In a few instances I have recognized new synonymies or have

changed the status of subspecies. Each change of this sort is discussed in the

taxonomic portion of this revision under the appropriate genus. Not discussed are

new combinations, unless the species involved possesses a combination of character

states unusual or atypical of the genus into which it is placed.

Type designations of Coquillett and Townsend. — The type concept became

increasingly popular among taxonomists in the late 1800’s and early 1900’s, and it

was not unusual for workers of that period to adopt the concept at some point in

their careers. Coquillett and Townsend were two such workers, and it is the status of

the type series of their earlier species that is of concern here.

It is evident that Coquillett, by the time of his 1897 “Revision of the

Tachinidae”, was not only choosing type specimens for new species but for his

previously described species as well. He assigned type numbers to each of his

USNM types, though only published numbers for species he described as new in the

“Revision” (Sabrosky, pers. comm.). For previously described species Coquillett

generally appended his redescription with the statement “From the type specimen”.

For the purposes of nomenclatural stability, and because Coquillett’s intent is clear,

I accept his reference to a type specimen in the redescription of a species described

from syntypes as a valid lectotype designation. (This was also the interpretation of

Sabrosky and Amaud 1963.) Similarly, I accept Coquillett’s citation of a type

number in the description of a new species as sufficient for a holotype designation,

provided specimens were labelled appropriately. Under the first situation two

siphonines are involved, S. (Siphonopsis) plusiae and Ceromya palloris, and under

the second S. (Ceranthia) flavipes and S. (Pseudosiphona) brevirostris. In the text

that follows I accept Coquillett’s lectotype designations of S. plusiae and C. palloris

and his holotype designation of S. flavipes. I have had to designate a lectotype for S.

brevirostris because Coquillett’s original holotype designation refers to a pin

bearing two specimens, neither of which was specifically chosen as the type.

There is only one siphonine described by Townsend, Actinocrocuta chaetosa, for

which a holotype was not designated in the original description. However, in his

1940 redescription of the genus (which is herein considered a subgenus of Siphona ),

Townsend cites the “Ht male” of A. chaetosa , clearly indicating his choice of the

Systematics of the Genus Group Taxa of the Siphonini

41

only male of the type series as the type. I accept this citation as a lectotype

designation.

Genus Goniocera Brauer and Bergenstamm

Figs. 1, 47, 71-72, 107, 137, 145, 158

Goniocera Brauer and Bergenstamm, 1891: 354. Type-species, G. schistacea Brauer and Bergenstamm,

1891 (monotypy).

Euthryptocera Townsend, 1916: 624. Type-species, Tachina latifrons Meigen, 1824 (original designation) =

Tachina versicolor Fallen, 1820.

Euchaetactia Villeneuve, 1921: 47 (as subgenus of Actia Robineau-Desvoidy). Type-species, Actia

{Euchaetactia) montium Villeneuve, 1921 (monotypy).

Cartocometes Aldrich, 1929: 9. Type-species, C. io Aldrich, 1929 (original designation). Recent synonymy

by Wood (1987: 1258) in key to Nearctic tachinid genera.

Recognition

This genus of four described species belongs to the group of siphonine genera in

which the anal vein does not extend to the wing margin (i.e. all genera except

Peribaea and Siphona s.l.), and is distinguished from other members of this group

by the presence of several ad setae on the mid tibia. Other siphonines have one ad

seta (Fig. 37) or none (Fig. 38), with the known exception of two undescribed

African species of S. ( Aphantorhaphopsis ) which have two setae - these two species

have the anal vein extended to the wing margin and other Siphona s.l.

characteristics. The distinctive shape of the distiphallus is probably autapotypic of

Goniocera (Fig. 107; refer to Description and Phylogenetics sections).

Three of the four Goniocera species have a densely setulose parafacial (Fig. 1).

The other species, G. versicolor , has the lower parafacial bare (i.e. the region of the

parafacial adjacent to the lower margin of the eye) as in most other siphonines

(several siphonines have hairs on lower parafacial, but fewer hairs than in the three

Goniocera species). Other features shared by species of Goniocera , though only

collectively unique to the genus, include ( cf . Table 1): wing vein R4+5 setulose from

base to beyond crossvein r-m and other veins bare, prominent and flattened median

lobe on male sternum 5 (Fig. 47; similar appearance in a few Ceromya species),

surstylus long and basally fused with epandrium (Fig. 137), and reduced female

sternum 8 (Fig. 145). The male and female genitalia of G. montium and the female

genitalia of G. versicolor were not examined, so it is unknown if these species share

the genitalic features of the other species.

Description

Length: 4.0-6.0mm.

Head (Fig. 1). — Five frontal setae, normal arrangment. Proclinate orbital setae subequal in size in most

species. Lower parafacial bare in G. versicolor, densely setulose in other species. Eye of male small to

medium, 0.58-0.71 head height; eye of female slightly smaller than in male. Flagellomere 1 of male

medium-short to medium-long, 0.42-0.67 head height; shape from linear to slightly broadened; not bifid.

Flagellomere 1 of female slightly smaller than in male. Aristomere 1 short. Aristomere two 2-4X longer than

wide. Aristomere 3 almost bare to micropubescent, rather short (G. montium) to long and tapered. Clypeus

Quaest. Ent., 1989, 25 (1,2)

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U-shaped. Palpus short to medium, clavate. Proboscis with prementum short, labella padlike.

Thorax. — Prostemum bare or setulose. Lower proepimeral seta weak, not directed downward.

Katepistemum bare anterior to mid coxa. Lower katepistemal seta shorter than upper anterior seta. Three

postsutural dorsocentral setae, except four in G. montium. Upper part of anepistemum lacking, or with

single, setula. Fore tibia with preapical ad seta varied: short in G. montium, half to subequal length of d seta

in G. schistacea, and at least length of d seta in G. io and G. versicolor. Mid tibia with row of ad setae, two

to seven long and one to three shorter in most taxa. Tarsomeres normal in size, claws small to medium.

Wing: CuA] with distal portion 0.26-0.47 length of proximal portion (i.e. dm-cu near wing margin; mean

0.34); anal vein not extended to wing margin. Wing setulae: /?, bare dorsally and ventrally; R4+5 setulose

from base to beyond r-m\ CuAi bare.

Abdominal terga 1-5. — Abdomen ovoid in shape. Setae varied in length intraspecifically: some

specimens with median and lateral marginal setae on 7j+2 and lateral discal setae on Tl+2-T5, others without

long setae on 7j+2 and with normal setation on TyTs.

Male genitalia (Figs. 47, 71-72, 107, 137). — S5 (Fig. 47) little varied, posterior margin approximately

U-shaped; processes with apical lobes clearly differentiated, at least as long as wide; median lobe unusually

prominent, in form of broad, truncate plate flattened posteromedially; processes moderately setulose. T6 in

form of single, narrow to broad, dorsally continuous sclerite. Ejaculatory apodeme with fan-shaped portion

subequal to or slightly wider than hypandrial apodeme. Pregonite in profile either narrow and linear (Fig.

72), or broad and truncate with membranous portion spinulose (G. io. Fig. 71). Epiphallus present, narrow

(not absent as stated in Andersen 1983 and shown in his fig. 22). Distiphallus (Fig. 107) large, in profile

more or less truncate apically and evenly tapered basally, posterolateral margin with short pointed sclerite

(not extended beyond apex of distiphallus) separated from broad lateral margin by narrow incision

(posterolateral margin curved toward midline and thus pointed sclerite not visible in Fig. 107), anterior

margin distinctly developed; apex at least as long as broad in ventral view. Postgonite apically narrow or

broad, turned outward. Surstylus long, thin and straight; fused basally with epandrium (Fig. 137). Cerci

elongate, sharply inflexed at midlength; thickly covered with long setae on basal half (Fig. 137).

Examined male genitalia of: G. io, G. schistacea and G. versicolor (last species shown in Andersen 1983,

fig. 22).

Female genitalia (Fig. 145). — Moderately extensible. S6 with very long hairs. T6 distinctly developed,

enclosing spiracles of segment 6; discontinuous dorsally. S7 without anterior apodeme; pointed posteriorly,

with (G. schistacea) or without (G. io) posteromedial keel. T7 present as two lateral sclerites; spiracles of

segment 7 in membrane between segments 6 and 7. S8 small, haired. 7j0 present as two small sclerites.

Examined female genitalia of: G. io. Female reproductive system of G. schistacea shown in Andersen (1983,

fig. 3).

Hosts

Hosts are known for three of the four species, and these are all parasites of

Malacosoma species (tent caterpillars) (Table 2).

Phylogenetics

The only externally autapotypic feature of Goniocera is the row of ad setae on

the mid tibia, though similarities in the male genitalia, specifically sternum 5 and

distiphallus, in three of the four species (male genitalia of G. montium not

examined) provide additional evidence supporting monophyly of the genus. With

respect to male sternum 5, the median lobe is unusually prominent, and flattened on

its posteromedial surface (Fig. 47). A trend in several siphonine groups is toward a

flattened median lobe ( e.g . Figs. 54, 68), but only in a few Ceromya s.s. species is

its shape similar to that in Goniocera species. Considering this trend, and the

presumably derived phylogenetic position of these Ceromya s.s. species within that

genus, it is probable that the characteristic shape of sternum 5 among Goniocera

Systematics of the Genus Group Taxa of the Siphonini

43

species is a synapotypy, and its similar appearance in Ceromya s.s. is the result of

convergence.

Shape of the distiphallus is similar among examined Goniocera species (Fig.

107). In particular the posterolateral margins are curved toward the midline, and

incised to form two short sclerotized projections (one per side) - as these projections

are only seen in posterior view they are not visible in Fig. 107. The only other

siphonines known to share this feature are two species of the Ceromya palloris

group, C. flaviseta and C. Ontario , which are not otherwise similar to Goniocera

species. The male genitalia of G. montium were not examined, but it is predicted

that they have the same states here suggested as synapotypies of the genus.

The pregonite of G. io differs from those of G. schistacea and G. versicolor by

the presence of tiny spinules on the membranous portion (Fig. 71). This feature is in

most Ceromya s.s. species, and is considered synapotypic of that group. The

presence of spinules on the pregonite of G. io can be interpreted in several ways, as

discussed in the Evolution chapter. These different interpretations of the pregonite in

G. io lead to different phylogenetic senarios regarding Goniocera (Figs. 166-169),

but none challenges the well established monophyly of the genus.

Adult females of G. io and G. schistacea have a small sternum 8 (Fig. 145),

which is a derived state among siphonines. The size of this sclerite is unknown in G.

montium and G. versicolor , but if also small, then would represent another

synapotypy of the genus.

Tent caterpillars ( Malacosoma species, Lasiocampidae) are the only known hosts

of Goniocera species (Table 2). Since G. montium is the only Goniocera species for

which a host is unrecorded, and Malacosoma species are not known hosts for any

other siphonines, I hypothesize that this parasitic specialization is a synapotypy of

Goniocera species.

Geographic distribution

This genus includes three sympatric species in the Palearctic region, all western

in distribution (Mesnil 1963a, Herting 1984): G. montium (a rarely collected species

only known from France), G. schistacea (Denmark and Middle Europe) and G.

versicolor (ranging from southern Sweden to England, France, Germany, Austria

and Poland). The single Nearctic species, G. io, is recorded from eastern Canada and

northeastern USA (Sabrosky and Amaud 1965).

List of described species included in Goniocera

N io (Aldrich), 1929: 10 ( Cartocometes ). Holotype female, USA: New York,

Riverhead (USNM).

P montium (Villeneuve), 1921: 47 ( Actia ( Euchaetactia )). Holotype male, France:

[Col du] Lautaret (CNC). Holotype examined.

P schistacea Brauer and Bergenstamm, 1891: 354. Holotype female, Austria (not

Quaest. Ent., 1989, 25 (1,2)

44

O’Hara

located).

syn. enigmatica Villeneuve and Nielsen in Nielsen, 1917: 32. Holotype

female, Denmark: Tisvilde (not located). — Mesnil, 1962: 800.

P versicolor (Fallen), 1820: 19 ( Tachina ). Syntypes, Sweden: Sk&ne (NRS).

syn. latifrons (Meigen), 1824: 365 (Tachina). Holotype female, Austria

(lost).— Herting, 1984: 120.

hartigii (Ratzeburg), 1844: 172 ( Musca (Tachina)). Type, Germany

(lost). — Herting, 1982: 8.

ludibunda (Robineau-Desvoidy), 1850: 195 (Ceromya). Syntypes,

France (lost). — Herting, 1974: 18.

Genus Proceromyia Mesnil

Figs. 2-3, 48-49, 73-74, 108-109, 146.

Proceromyia Mesnil, 1957: 35 (as subgenus of Ceromya). Type-species, Ceromya ( Proceromyia )

macronychia Mesnil, 1957 (monotypy).

Nipponoceromyia Mesnil and Shima, 1978: 324. Type-species, N. pubioculata Mesnil and Shima, 1978

(original designation). New synonymy.

Recognition

The two described species of Proceromyia are only recorded from Japan and the

Kuril Islands. One, P. pubioculata , is markedly autapotypic in external head

features, differing from other siphonines in having a densely haired eye and 8-12

frontal setae (Fig. 3). The other species, P. macronychia , is less distinctive and

easily mistaken externally for a Ceromya species (head shown in Fig. 2). Both

species share externally the following unique combination of character states (cf.

Table 1): narrow vertex (apparently unique, but not quantitatively assessed in this

study), prostemum bare (rare among siphonines), fore tibia with preapical ad seta

subequal in length or longer than d seta (as in Entomophaga species and several

other species), large tarsal claws (only as large in some Siphona s.s. species), wing

vein R4+5 setulose between base and crossvein r-m and other veins bare, and anal

vein not extended to wing margin.

Features of the male genitalia, particularly shape of sternum 5 (Figs. 48-49),

pregonite (Figs. 73-74) and distiphallus (Figs. 108-109) are also diagnostic for the

genus, though the latter two structures closely resemble those in the sister genus

Entomophaga (Figs. 75-76 and 110-111). Proceromyia species are distinguished

externally from those of Entomophaga by their short aristomere 1 (cf. Figs. 2-3 and

4-5), bare prostemum and larger tarsal claws.

Key to adults of Proceromyia species

(See section entitled “Review of major keys to genera and subgenera of the

Siphonini” for information about how other authors have keyed (and classified) the

following species.)

Systematics of the Genus Group Taxa of the Siphonini

45

1. Eye densely haired; 8-12 frontal setae (Fig. 3, eye hairs not

shown) (Japan) Proceromyia pubioculata Mesnil & Shima

1' Eye almost bare; five frontal setae (Fig. 2) (Japan and Kuril

Islands) Proceromyia macr onychia Mesnil

Description

(Note: female of P. pubioculata unknown.)

Length: 3.0-5. 5mm.

Head (Figs. 2-3). — (Head of P. pubioculata also shown in Mesnil and Shima 1978, fig. 10.) Generally

five frontal setae in P. macr onychia, 8-12 in P. pubioculata, rather fine in both. Proclinate orbital setae thin

but average length, subequal in most specimens. Eye almost bare ( P . macronychia) or densely haired (P.

pubioculata). Eye of male and female subequal, medium-large, 0.78-0.84 head height. Flagellomere 1 of

male short to medium-short, 0.34-0.43 head height; shape average in width; not bifid. Flagellomere 1 of

female usually slightly smaller than in male. Aristomere 1 short. Aristomere two 1.5-3X longer than wide.

Aristomere 3 almost bare, slightly short to normal length, evenly tapered to tip or abruptly narrowed on

apical two-thirds. Clypeus U-shaped in P. macronychia, not examined in P. pubioculata. Palpus short to

medium, clavate. Proboscis with prementum short, labella padlike.

Thorax. — Prostemum bare. Lower proepimeral seta weak, not directed downward. Katepistemum bare

anterior to mid coxa. Lower katepistemal seta shorter than upper anterior seta. Three postsutural

dorsocentral setae. Upper part of anepistemum with single setula in P. macronychia, with three setulae in

single examined specimen of P. pubioculata. Fore tibia with preapical ad seta subequal to or longer than d

seta. Mid tibia with one ad seta. Tarsomeres normal in size; claws large. Wing: CuAl with distal portion

0.24-0.32 length of proximal portion (i.e. dm-cu near wing margin; mean 0.27); anal vein not extended to

wing margin. Wing setulae: R] bare dorsally and ventrally; /?4+5 setulose between base and r-m\ CuA] bare.

Abdominal terga 1-5. — Abdomen ovoid in shape. T]+2 without median marginal setae; lateral marginal

setae strong. T3-T5 average or with lateral discal setae on one or more segments.

Male genitalia (Figs. 48-49, 73-74, 108-109). — S5 (Figs. 48-49) little varied, posterior margins of

processes obtusely angled; apical lobe slightly differentiated; median lobe rounded, relatively unmodified;

processes moderately setulose. T6 forming single, broad, dorsally continuous sclerite. Ejaculatory apodeme

with fan-shaped portion slightly wider to about 1.5X wider than hypandrial apodeme. Pregonite (Figs.

73-74) in profile smoothly curved anteriorly and more or less pointed apically, with small spines along

anterolateral ridge. Epiphallus small in P. pubioculata, absent from P. macronychia. Distiphallus (Figs.

108-109) with posterior margin partially reduced, laterally incised, anterior margin reduced except for long,

spined anterolateral arm; apex broader than long in ventral view. Postgonite apically broad and turned

outward. Surstylus average length, broadened at midlength, straight; basally free from epandrium. Cerci

rather short (P. macronychia) to average length (P. pubioculata), smoothly curved at midlength; moderately

setose on basal half.

Examined male genitalia of: P. macronychia and P. pubioculata (latter shown in Mesnil and Shima 1978,

fig. 11).

Female genitalia (Fig. 146). — (Only P. macronychia examined.) Short. S6 with short hairs. T6 distinctly

developed, enclosing spiracles of segment 6; narrowly discontinuous dorsally. S7 with long anterior

apodeme; without posteromedial keel. T7 present as two lateral sclerites; spiracles of segment 7 displaced

anteriorly and enclosed in posterior margin of T6. S8 distinctly developed, haired. Tx0 present as small

median sclerite.

Examined female genitalia of: P. macronychia.

Hosts. Unknown.

Phylogenetics

Mesnil described P. macronychia in 1957 (p. 35) as a new species in new

subgenus Ceromya ( Proceromyia ), basing the new taxon primarily on the presence

of large claws and narrow vertex. At that time Ceromya s.l. was recognized as a

Quaest. Ent., 1989, 25 (1,2)

46

O’Hara

siphonine genus in which the anal vein did not extend to the wing margin and from

which the derived features of Actia and Goniocera were lacking.

Mesnil later completely revised the Palearctic Ceromya species (1963a: 829),

and recognized two primary divisions within the genus: Ceromya (Ceromya)

characterized by wing vein R4+5 setulose from base to beyond crossvein r-m, and

both C. (Proceromyia) and C. (Stenoparia) characterized by vein R4+5 not setulose

beyond r-m. I discuss here the monophyly of Mesnil’ s C. (Proceromyia) - status of

C. (Stenoparia) and placement of its type species C. monstrosicornis are discussed

in the Phylogenetics section of Ceromya s.s., while the other C. (Stenoparia) species

recognized by Mesnil, C. nigrohalterata, is placed in Entomophaga and discussed in

the Phylogenetics section of that genus.

The second species here included in Proceromyia is a siphonine of unusual

appearance, P. pubioculata. This species was described in its own genus,

Nipponoceromyia Mesnil and Shima (1978), primarily because of its uniquely

haired eye and numerous frontal setae (Fig. 3). The authors commented that P.

pubioculata seemed related to Proceromyia macronychia (1978: 325), citing as

evidence the shared possession of weak proclinate orbital setae, narrow vertex and

short distal section of wing vein CuAx. I doubt that the first character state is

uniquely shared by these two species, the last certainly is not, but the narrow vertex

might be a synapotypy of these species. Both species also have unusually large

tarsal claws. The male genitalia of Proceromyia macronychia and Nipponoceromyia

pubioculata were apparently not compared by Mesnil and Shima, but corroborate

their phylogenetic hypothesis and as discussed below provide better evidence for a

sister species relationship between these species.

Similarities in the male genitalia of Proceromyia macronychia and

Nipponoceromyia pubioculata belie the external differences between these species,

leading to the conclusion that the autapotypic features of N. pubioculata are the

result of divergence from a more P. macronychia-like ancestor. In support of this

are the remarkable similarities in male sternum 5 (Figs. 48-49), pregonite (Figs.

73-74) and distiphallus (Figs. 108-109) - similarities which also attest to a close

relationship with Entomophaga (see the Phylogenetics section of that genus and the

Evolution chapter for a discussion of these intergeneric relationships). Given the

general external similarities between P. macronychia and P. pubioculata (listed in

Recognition section) and diverse shapes of male genitalia among siphonines in

general, I regard the shared similarities in male genitalia of these two species as

synapotypies. I propose the following classification to reflect this hypothesized

relationship between P. macronychia and N. pubioculata : N. pubioculata is moved

to Proceromyia , and Proceromyia is ranked at the generic level because of its sister

group relationship with Entomophaga , and its lack of known synapotypies with

Ceromya. Both Proceromyia and Nipponoceromyia were ranked as monobasic

genera by Herting (1984: 121-2) before the male genitalia of the two included

species were compared (in contrast, Andersen 1983 placed Proceromyia as a

Systematics of the Genus Group Taxa of the Siphonini

47

synonym of Ceromya and did not study Nipponoceromyia pubioculata). It seems

appropriate to modify Herting’s classification of these two species by combining

them under one generic name now that their genitalic features are known to be so

similar.

Geographic distribution

The known ranges of the two included species are very limited: P. macronychia

was described from Hokkaido, Japan, and has since been recorded from the Kuril

Islands, USSR (Richter 1976b). P. pubioculata is only known from male specimens

collected from Honshu Island, Japan.

List of described species included in Proceromyia

P macronychia (Mesnil), 1957: 35 ( Ceromya {Proceromyia)). Holotype male,

Japan: Hokkaido, Obihiro (CNC). Holotype examined.

P pubioculata (Mesnil and Shima), 1978: 325 {Nipponoceromyia). Holotype

male, Japan: Honshu, Kawaragoya (BLKU). Paratype examined. New

combination.

Genus Entomophaga Lioy

Figs. 4-5,50-51,75-76, 110-111, 147.

Entomophaga Lioy, 1864: 1332. Type-species, Tachina exoleta Meigen, 1824 (by designation of Coquillett,

1910: 538).

Recognition

Entomophaga comprises two described European species, E. nigrohalterata and

E. exoleta. The former is commonly collected while the latter is known from very

few specimens. A diagnostic combination for these species is the possession of an

elongate aristomere 1 (Figs. 4-5), fore tibia with preapical ad seta subequal in length

or longer than d seta, wing vein R4+5 setulose between base and crossvein r-m and

other veins bare, and anal vein not extended to wing margin {cf. Table 1). In

addition, features of the male genitalia are distinctive, and are similar only to the

male genitalia of the externally very different appearing Proceromyia species {cf.

head profiles in Figs. 2-5, sternum 5 in Figs. 48-51, pregonite in Figs. 73-76 and

distiphallus in Figs. 108-111).

Entomophaga exoleta has been considered an Actia species by some authors

because of its row of hairs on the katepistemum anterior to the mid coxa (as in Fig.

33), but is clearly misplaced there based on other features (see Phylogenetics section

and Evolution chapter). Likewise, E. nigrohalterata has been mistaken for a

Ceromya species because its phylogenetically important character states have been

misinterpreted.

Quaest. Ent., 1989, 25 (1,2)

48

O’Hara

Key to adults of Entomophaga species

(See section entitled “Review of major keys to genera and subgenera of the

Siphonini” for information about how other authors have keyed (and classified) the

following species.)

1 . Three postsutural dorsocentral setae; katepistemum with row of

hairs directly anterior to mid coxa, extended upward almost to

lower katepistemal seta (as in Actia spp., Fig. 33); male

sternum 5 U-shaped, apical lobe distinctly differentiated (Fig.

50); pregonite spined along anterolateral margin (Fig. 75);

distiphallus with lateral margin markedly reduced (Fig. 110)

(Europe; very rarely collected) . Entomophaga exoleta (Meigen)

1'. Four postsutural dorsocentral setae; katepistemum almost bare

directly anterior to mid coxa, except for several hairs in

posteroventral comer (Fig. 34); male sternum 5 with apical lobe

slightly differentiated (Fig. 51); pregonite bare along

anterolateral margin (Fig. 76); distiphallus with lateral margin

partially reduced (Fig. Ill) (Europe; commonly

collected) Entomophaga nigrohalterata (Vill.)

Description

Length: 3.0-5. 0mm.

Head (Figs. 4-5). — Four or five frontal setae, in normal arrangement. Anterior proclinate orbital seta

longer than posterior one in E. exoleta, setae subequal in length in E. nigrohalterata. Eye bare. Eye of male

small to medium, 0.63-0.77 head height; eye of female slightly larger than in male. Flagellomere 1 of male

medium to medium-long, 0.58-0.66 head height; shape broad to subquadrangular; not bifid. Flagellomere 1

of female slightly smaller than in male. Aristomere 1 elongate, 1.5-5X longer than wide. Aristomere two

3-4X longer than wide. Aristomere 3 almost bare, slightly shorter than average, evenly tapered to tip.

Clypeus U-shaped. Palpus long, clavate. Proboscis with prementum short, labella padlike.

Thorax. — Prostemum setulose. Lower proepimeral seta weak, not directed downward. Katepistemum

bare {E. nigrohalterata) or with row of hairs anterior to mid coxa (E. exoleta). Lower katepistemal seta

shorter than upper anterior seta. Three (E. exoleta) or four (E. nigrohalterata) postsutural dorsocentral setae.

Upper part of anepistemum with single setula. Fore tibia with preapical ad seta subequal to or longer than d

seta. Mid tibia with one ad seta. Tarsomeres normal in size, claws small to medium-large. Wing: CuAx with

distal portion 0.25-0.43 length of proximal portion (i.e. dm-cu near wing margin; mean 0.33); anal vein not

extended to wing margin. Wing setulae: Ri bare dorsally and ventrally; R4+5 setulose between base and r-m;

CuA] bare.

Abdominal terga 1-5. — Abdomen ovoid in shape. 7j+2 without median marginal setae; lateral marginal

setae strong. T3-T5 average or with lateral discal setae on 1 or more segments.

Male genitalia (Figs. 50-51, 75-76, 110-111). — S5 with apical lobe of processes slightly differentiated

(posterior margins almost transverse, Fig. 51) in E. nigrohalterata, distinctly differentiated and posterior

margin approximately U-shaped in E. exoleta (Fig. 50); median lobe rounded, relatively unmodified;

processes moderately setulose. T6 forming single, broad, dorsally continuous sclerite. Ejaculatory apodeme

with fan-shaped portion slightly wider to almost 2.0X wider than hypandrial apodeme. Pregonite in profile

smoothly curved anteriorly, pointed apically; either bare (E. nigrohalterata. Fig. 76) or spined along

anterolateral margin (£. exoleta. Fig. 75). Epiphallus present: very short in E. nigrohalterata, long and

narrow in E. exoleta. Distiphallus (Figs. 110-111) with posterior margin partially reduced, laterally incised,

anterior margin reduced except for long, spined anterolateral arm; broader than long in ventral view.

Postgonite apically broad and turned outward. Surstylus average length, straight (E. nigrohalterata) or

Systematics of the Genus Group Taxa of the Siphonini

49

curved posteriorly ( E . exoleta); basally free from epandrium. Cerci rather short, smoothly curved at

midlength; moderately setose on basal half.

Examined male genitalia of: E. exoleta and E. nigrohalterata (latter shown in Andersen 1983, fig. 24).

Female genitalia (Fig. 147). — (Only E. nigrohalterata examined.) Short. S6 with very short hairs. T6

distinctly developed, enclosing spiracles of segment 6; narrowly discontinuous dorsally. S7 with long

anterior apodeme; slightly keeled posteromedially. f7 present as two lateral sclerites; spiracles of segment 7

displaced anteriorly to very near or associated with T6. S8 absent. Tl0 absent or present as two small sclerites.

Examined female genitalia of: E. nigrohalterata (also see Andersen 1983, fig. 9).

Hosts. Unknown.

Phylogenetics

Herting (1975: 4) revised the standard classification of E. exoleta and E.

nigrohalterata by removing the former from Actia and the latter from Ceromya, and

uniting them under a newly defined Entomophaga Lioy (cf. Mesnil 1963a). Herting

based his concept of Entomophaga primarily on two characteristics shared by E.

exoleta and E. nigrohalterata : an elongate aristomere 1 and long preapical ad seta

on fore tibia. The second feature is shown in this study to be shared with related taxa

(see Evolution chapter), but the first is accepted here as a synapotypy of

Entomophaga, though not unique to the genus (Table 1).

Recently Andersen (1983: 12) reviewed the Old World Siphonini and

re-assigned E. exoleta to Actia and E. nigrohalterata to Ceromya. Andersen returned

E. exoleta to Actia because he considered its row of katepistemal hairs to be clear

evidence of its membership in Actia. He interpreted the elongate aristomere 1 of E.

exoleta and E. nigrohalterata as autapotypies of each species, consequently

returning the latter to Ceromya because “it does not differ significantly from other

Ceromya species in external characters” (1983: 12). However, Andersen noted that

the male and female genitalia of E. nigrohalterata were different from the

Ceromya- type.

Andersen’s classification of the two species here placed in Entomophaga and

Proceromyia was based on external characters, since only the genitalia of E.

nigrohalterata were examined. I hypothesize that characteristics of the male

genitalia of the four Entomophaga and Proceromyia species indicate that these

species form a monophyletic lineage, based on shape of the pregonite (Figs. 73-76)

and distiphallus (Figs. 108-111), which are derivable from a common groundplan

and lack synapotypies with the Actia and Ceromya lineages. The two Proceromyia

species are not only remarkably similar in male genitalic features (Figs. 48-49,

73-74, 108-109) but share unique external similarities as well, so are certainly sister

species. The male genitalia of Entomophaga species are more equivocal: they

clearly indicate a close relationship with Proceromyia, but not a sister species

relationship between E. exoleta and E. nigrohalterata. However, given the general

external similarities between E. exoleta and E. nigrohalterata, particularly head

habitus (Figs. 4-5) and derived state of aristomere 1, these species most probably

form a monophyletic group. If these external similarities are not synapotypies then

Quaest. Ent., 1989, 25 (1,2)

50

O’Hara

Entomophaga might be paraphyletic with respect to Proceromyia (i.e. either E.

exoleta or E. nigrohalterata being more closely related to the Proceromyia lineage

than to its congener).

Features of the male genitalia of E. exoleta not only corroborate its placement in

the Entomophaga+Proceromyia lineage, but provide the best evidence for removing

this species from Actia. Actia species share the synapotypies of a more or less

V-shaped sternum 5 (Figs. 58-59) and apically spinose pregonite (Figs. 85-86), and

these states are lacking from E. exoleta. It is concluded that the row of katepistemal

hairs that are found (among siphonines) only in E. exoleta and most Actia species is

the result of convergence; convergence certainly accounts for the presence of this

row of hairs in Actia and some non-siphonine tachinids. E. exoleta also has one seta

on the upper portion of the anepistemum while almost all Actia species have two (cf.

Figs. 31 and 32).

E. nigrohalterata has not been shown to share any synapotypies with Ceromya,

either here or by previous authors. Spinules on the membranous portion of the

pregonite in most Ceromya s.s. species, here considered a synapotypy of Ceromya

s.s., are absent from E. nigrohalterata. Further research is needed to resolve

relationships within Ceromya s.l., and will not only provide information relevant to

the placement of E. nigrohalterata , but to other species of Entomophaga and

Proceromyia as well.

Adult females of E. nigrohalterata are atypical in their lack of a sclerotized

abdominal sternum 8 (Fig. 147). Study of the female genitalia of E. exoleta has not

yet been possible, but if sternum 8 is lacking then a sister species relationship

between E. exoleta and E. nigrohalterata would be corroborated (otherwise this loss

might be autapotypic of E. nigrohalterata).

Similar in some respects to the species of Entomophaga is Ceromya

monstrosicornis. Its placement is discussed in the Phylogenetics section of Ceromya

s.s.

Geographic distribution

Entomophaga exoleta is a rarely collected species, recorded from a few localities

in France (type locality, and a female in SMNS), Hungary (Andersen, pers. comm.)

and England (Crosskey 1976b). E. nigrohalterata is a relatively common species in

Europe, with records from England, Belgium, Denmark, Germany, Poland, Austria

and Switzerland (Mesnil 1963a, Herting 1967b, 1984, Draber-Moriko 1978, 1981).

E. exoleta and E. nigrohalterata are at least narrowly sympatric, with the former

slightly more southern in distribution.

List of described species included in Entomophaga

P exoleta (Meigen), 1824: 353 ( Tachina ). Syntypes, France: Provence (MNHN).

syn. anicula (Meigen), 1824: 409 (Tachina). Holotype female, Europe

Systematics of the Genus Group Taxa of the Siphonini

51

(MNHN).— Herting, 1975: 2.

P nigrohalterata (Villeneuve), 1921: 45 ( Actia ). Holotype male, Denmark (CNC).

Holotype examined.

syn. articulata (Stein), 1924: 131 {Actia). Syntypes, Copenhagen,

Denmark and Lobauer Berg, German Democratic Republic (ZMUC;

not located, possibly lost). — Lundbeck, 1927: 465.

sufferta (Villeneuve), 1942b: 133 {Actia). Holotype male, Germany:

no locality given (CNC). — Herting, 1981: 8. Holotype examined.

Genus Ceromya Robineau-Desvoidy sensu lato

Figs. 6-7, 35, 37, 39-40, 52-57, 77-84, 112-116, 138, 148-150, 157, 159.

Note about classification of Ceromya s.l. species

No known features are clearly interpretable as synapotypic of Ceromya s.l., and

as a result the monophyly of this diverse and cosmopolitan genus has not been

positively established. However, features of the male genitalia suggest that Ceromya

s.l. comprises two monophyletic lineages. These are here called Ceromya s.s.

(including type species C. bicolor) and the C. silacea species group (an informal

species group in which C. silacea is the most derived species and has the oldest

name), and are treated individually throughout this paper.

The unusual division of Ceromya s.l. into two categories of different rank is

adopted for practical and nomenclatural reasons. Firstly, male genitalia were not

studied in all Ceromya s.l. species, so characterization of the lineages may be

incomplete, as well as assignment of described species to the C. silacea species

group. Secondly, recognition of the C. silacea species group as a subgenus or genus

would require erection of a new genus-group name, which is premature on the basis

of available evidence. Ceromya s.s. could equally well be called the C. bicolor

species group, but the former designation was chosen to clearly identify the group

that would be called Ceromya if the C. silacea species group is later removed from

Ceromya s.l.

Recognition

Ceromya s.l. belongs to the group of siphonines in which the anal vein is not

extended to the wing margin (Fig. 21). This characteristic separates all known

Ceromya s.l. species from species of Peribaea and Siphona s.l. Additionally, the

lower katepistemal seta is shorter than the upper anterior one in all but a very few

Ceromya s.l. species (Fig. 31) while subequal in length or longer in most Peribaea

and Siphona s.l. species (Fig. 32; see Table 1).

New World species of Ceromya s.l. are easily separable from other New World

siphonines by the following combination of states: anal vein not extended to wing

margin (Fig. 21), absence of row of hairs on katepistemum (Fig. 34; row present in

all New World Actia species, Fig. 33) and only one ad seta on mid tibia (Fig. 37;

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row of ad setae in Goniocera io, the only New World Goniocera species).

Four Old World genera - Goniocera, Actia, Entomophaga and Proceromyia - are

characterized along with Ceromya s.l. by an anal vein not extended to the wing

margin. All Goniocera species have a row of ad setae on the mid tibia (0-1 seta in

Ceromya s.l. species, as in Figs. 37-38) and are separable from Ceromya s.l. species

by this feature. The diagnosis given above for New World Ceromya s.l. species will

not separate Old World members of this taxon from species of Entomophaga or

Proceromyia, or several Actia species. It is therefore necessary to diagnose Old

World Ceromya s.l. species by the presence of at least six of the following seven

states: 0-1 anepistemal setula, lower katepistemal seta shorter than upper anterior

one, katepistemum lacking row of hairs, fore tibia with short preapical ad seta, mid

tibia with 0-1 ad seta, wing vein R4+5 setulose beyond crossvein r-m and anal vein

not extended to wing margin. Similarities and differences among Old World

Ceromya s.l. species and Old World Actia, Entomophaga and Proceromyia species

are discussed below.

Most Old World Actia species are distinguishable from Ceromya s.l. species by

presence of a row of katepistemal hairs (Fig. 33), but a very few Old World Actia

species lack this row of hairs and must be recognized by other features (externally

by presence of two anepistemal setulae, though characteristics of the male genitalia

provide the best means by which to recognize Actia species - see Recognition

section of Actia).

The four species of Entomophaga and Proceromyia share two states which in

combination distinguish them from Ceromya s.l. species: fore tibia with preapical ad

seta subequal in length or longer than d seta (only shared with two Ceromya species:

Nepal sp. 1 and Australia sp. 3) and wing vein R4+5 not setulose beyond crossvein

r-m (same in three Ceromya species: C. cephalotes, C. monstrosicornis and C.

natalensis ). Features of pregonite and distiphallus are also unique to the

Entomophaga+Proceromyia lineage.

Male genitalic features of Ceromya s.l. species are discussed in the Recognition

sections of Ceromya s.s. and C. silacea species group. The possible monophyly of

Ceromya s.l. is discussed in the Evolution chapter.

Ceromya Robineau-Desvoidy sensu stricto

Figs. 6-7, 39-40, 53-57, 77-83, 113-116, 138, 149-150, 157.

Ceromya Robineau-Desvoidy, 1830: 86. Type-species, C. testacea Robineau-Desvoidy, 1830 (by

designation of Coquillett, 1910: 520) = Tachina bicolor Meigen, 1824.

Ceromyia. Variant spelling of Ceromya.

Polychaetoneura Walton, 1914: 90. Type-species, P. elyii Walton, 1914 (original designation) =

Thryptocera americana Townsend, 1892. Recent synonymy by Wood (1987: 1239) in key to Nearctic

tachinid genera.

Xanthoactia Townsend, 1919: 585. Type-species, Lasioneura palloris Coquillett, 1895 (original

designation). Recent synonymy by Wood (1987: 1239) in key to Nearctic tachinid genera.

Stenoparia Stein, 1924:5'. (Siphona) 128. Type-species, 5. monstrosicornis Stein, 1924 (monotypy).

Systematics of the Genus Group Taxa of the Siphonini

53

Schizoceromyia Townsend, 1926b: 542. Type-species, Schizotachina fergusoni Bezzi, 1923 (original

designation).

Actinactia Townsend, 1927: 248. Type-species, A. lutea Townsend, 1927 (original designation). New

synonymy.

Schizactiana Curran, 1927b: 356 (as subgenus of Actia). Type-species, Actia ( Schizactiana ) valida Curran,

1927 (original designation).

Pseudactia Malloch, 1930b: 124 (as subgenus of Actia). Type-species Actia ( Pseudactia ) hirticeps Malloch,

1930 (monotypy).

Recognition

Features of the male genitalia provide the only means for placing Ceromya s.l.

species into Ceromya s.s. or the Ceromya silacea species group. The following

characteristics of the male genitalia distinguish species of Ceromya s.s. from those

of the C. silacea species group: pregonite with enlarged membranous area

anteriorly, with tiny spinules in most species (Figs. 39-40, 77-83; similar in

Goniocera io (Fig. 71), otherwise unique to Ceromya s.s. species); and distiphallus

without infolded, sclerotized structure posteriorly (Figs. 113-116; cf. C. silacea , Fig.

112).

External features of specimens of most described Ceromya s.l. species were

examined during this study, but male genitalia were examined in specimens of only

a portion of these species (as listed in descriptions of male genitalia of Ceromya s.s.

and C. silacea sp. grp.). Possibly a complete study of the male genitalia of all

species listed in Ceromya s.s. will reveal several species that should be reassigned to

the C. silacea species group. The evidence available at this time is insufficient to

permit a thorough reclassification of Ceromya s.l. species into Ceromya s.s. and C.

silacea species group.

Description

Length: 3.0-6.0mm.

Head (Figs. 6-7). — Five frontal setae in most species, three or four in a very few; normal arrangement.

Anterior proclinate orbital seta longer than posterior one in most species. Eye size of male medium-small to

large, 0.68-0.86 head height; eye of female slightly smaller to slightly larger than in male, in most species

eye size of sexes subequal. Flagellomere 1 of male markedly varied in length, 0.38-0.80 head height; shape

from linear to subquadrangular, bifid in C. fergusoni, C. invalida and C. valida, normal in others.

Flagellomere 1 of female smaller than in male; not bifid. Aristomere 1 short. Aristomere two 1.5-6X longer

than wide in species with normal antenna, up to 10X in species with bifid flagellomere 1. Aristomere 3 short

and thickened to near tip, to long and evenly tapered; almost bare, micropubescent, or rarely short plumose.

Clypeus U-shaped in most species, narrow and enclosed in membrane in a few. Palpus short in most species,

medium to long in a few; enlarged apically in females of a few species; clavate. Proboscis with prementum

short, labella padlike (labella very slightly lengthened in C. luteicornis).

Thorax. — Prostemum setulose in most species, bare in a very few. Lower proepimeral seta weak, not

directed downward except in a very few species (though not long as in Peribaea species). Katepistemum

bare anterior to mid coxa. Lower katepistemal seta much shorter than upper anterior seta in most species,

slightly shorter or subequal to it in two closely related species: C. flaviseta and C. Ontario. Three or four

postsutural dorsocentral setae. Upper part of anepistemum with single setula. Fore tibia with preapical ad

seta much shorter than d seta in almost all species, known to be as long as d seta only in Nepal sp. 1 and

Australia sp. 3. Mid tibia without ad seta in C. fergusoni, one ad seta in other species. Tarsomeres normal in

size in most species, with tarsomere 5 of fore leg elongate and broadened in female of several species;

tarsomere 5 of all legs elongate and dilated in female of Australia sp. 3; claws short in most species, medium

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in a few. Wing: CuAx with distal portion 0.24-0.67X length of proximal portion (mean 0.40); anal vein not

extended to wing margin. Wing setulae: /?, dorsally bare, or distally or entirely setulose, ventrally bare or

distally setulose; R4+5 setulose from base to beyond r-m in most species, not beyond r-m in C. cephalotes, C.

natalensis and C. monstrosicornis. CuAx bare or setulose, with basal section setulose in C. americana

complex, C. languidula, Brazil spp. 1 and 4 (bare in other siphonines).

Abdominal terga 1-5. — Abdomen ovoid in shape in almost all species, slightly elongate in a very few.

ri+2 without median marginal setae; lateral marginal setae absent to strong. T3-T5 average in most species,

with weak lateral discal setae (T3-T5) and/or extra pair of lateral marginal setae (T3 and T4 only) in a few.

Male genitalia (Figs. 39-40, 53-57, 77-83, 113-116, 138). — S5 markedly varied, inner (or posterior)

margins of processes obtusely angled (Fig. 55) to transverse (Fig. 57), in most species U-shaped (Figs.

53-54, 56); apical lobe usually undifferentiated (Fig. 57) to narrowly pointed, in a very few species distinctly

differentiated and apically curved inward (Fig. 56); median lobe markedly varied, from rounded (Figs.

55-56) or elongate (Fig. 53) to flattened plate of varied forms (Fig. 54; rarely similar to shape in Goniocera),

and with or without accessory lobe; processes sparsely to densely setulose. T6 varied from two very small

lateral sclerites to a single, broad, dorsally continuous sclerite. Ejaculatory apodeme with fan-shaped portion

0.5-1.5X width of hypandrial apodeme. Pregonite in profile extremely varied: membranous anterior portion

of pregonite enlarged in most species and usually with spinules anterolaterally (Figs. 39-40, 79-83; these

often tiny and visible only at higher magnifications, 100-400X); in one species (C. lutea. Fig. 83) pregonite

elongate and curved posteriorly (unique within the Siphonini); in one Old World group pregonite

ring-shaped and spinules borne anteriorly on sclerotized portion (Fig. 78). Epiphallus present or absent.

Distiphallus (Figs. 113-116) extremely varied: in most species anterior margin incised, posterior margin

entire in C. lutea (Fig. 116) and Nepal sp. 1, at least partially incised in other species, in profile distiphallus

laterally incised to varied degrees in most species (Figs. 114-116), lateral margin entire apically in a few

(Fig. 113). Postgonite large, apically rounded, curved ventrally, or bilobed. Surstylus short to long, thin to

broad (Fig. 138), straight or curved posteriorly; with long hairs along length in C. languidula (and in C.

varichaeta of Ceromya silacea sp. grp.); basally fused with epandrium in most species (Fig. 138), free in

others. Cerci short to average length, deeply inflexed at midlength in some species (Fig. 138), smoothly

curved in others; moderately to densely (Fig. 138) setose on basal half.

Examined male genitalia of: C. amblycera, C. americana complex, C. cornuta, C. flaviceps, C. flaviseta, C.

invalida, C. languidula, C. lavinia, C. lutea, C. natalensis, C. Ontario, C. palloris, C. nr. punctipennis,

Australia spp. 1-2,5, Brazil spp. 1,3-4, Chile spp. 1-2, Mexico spp. 2-5, Nepal sp. 1, New Guinea spp.

[numbered by Shima] 4,6,11,15,18, Peru sp. 1 and U.S. sp. 1. Examined published figures of: C. bicolor

(Andersen 1983, fig. 23) and C. pruinosa (Shima 1970c, figs. 3f, 6).

Female genitalia (Figs. 149-150). — Short to relatively long and extensible. S6 with short to average

length hairs, though species with short hairs usually sparsely haired on most of stemite with longer row of

hairs along posterior margin. T6 varied from two small lateral sclerites to distinctly developed and narrowly

discontinuous dorsally; spiracles of segment 6 near or enclosed within anteroventral portion of T6. S1 without

anterior apodeme in most species, with apodeme in a few; a few species slightly to (rarely) sharply keeled

posteromedially. T1 absent from a few species, present as two small to large lateral sclerites in most species;

spiracles of segment 7 in most species in membrane between segments 6 and 7, in a few species enclosed

within T6 or near T1. S8 distinctly developed, haired. F10 absent to distinctly developed as a median plate or

two sclerites.

Examined female genitalia of: C. americana complex, C. bicolor (Andersen 1983, fig. 10), C. cornuta, C.

lavinia, C. lutea, C. Ontario, Australia sp. 3 and Nepal sp. 4.

Taxonomic changes

Lectotype designation for Ceromya cibdela (Vill.). — Ceromya cibdela was

described in Actia by Villeneuve in 1913 from an unspecified number of specimens

collected from Oshogbo, Nigeria, during October and November of 1910. While

studying siphonine collections in the BMNH and CNC, I found a specimen in each

labelled as the type of C. cibdela and bearing appropriate locality data. Each

specimen bears a type label characteristic of the institution: a round, red-bordered

“Type” label on the BMNH specimen, and a red, rectangular “TYPE” label on the

Systematics of the Genus Group Taxa of the Siphonini

55

CNC specimen (of the sort used in the CNC to denote types in the Mesnil

collection). Both additionally have attached a Villeneuve determination label with

the designation “Typ.”. Both specimens fit Villeneuve’s brief description of the

species and by all indications are syntypes (no holotype designation was published).

They are not, however, conspecific. The CNC specimen belongs to Ceromya and

agrees with the current interpretation of C. cibdela (Mesnil 1954, 1963a, Crosskey

1976a), while the BMNH specimen belongs to Siphona (Aphantorhaphopsis). To

retain C. cibdela in its current usage I hereby designate the CNC specimen as

lectotype. The condition of the CNC specimen also favors its selection as lectotype

because it is a male in good condition while the BMNH specimen lacks its

abdomen.

Notes about Ceromya fergusoni Bezzi and two related nominal species. — Three

named species of Ceromya from eastern Australia have flagellomere 1 bilobed: C.

fergusoni (Bezzi 1923b; type-species of Schizoceromyia ), C. valida (Curran 1927b;

type-species of Schizactiana) and C. invalida (Malloch 1930a). These are

unquestionably at least closely related, and as discussed below, perhaps conspecific.

I dissected a male paratype (USNM) of C. invalida , and the presence of spinules

distally on the membranous portion of the pregonite confirms its placement in

Ceromya s.s.

As noted by Crosskey (1973: 137), the type of C. fergusoni is missing. Amaud

(1982) cites a paratype in MCSN, but this is in error. The MCSN specimen

(examined by me in 1986) bears a collection label of “Sydney, 3.12.23 [December

3, 1923], Health Dept.”, and stands in the MCSN collection under the label

“ Schizotachina fergusoni parat. Bezzi”. Bezzi did not name a paratype, and his

paper describing the species was read on November 28, 1923, about a week before

the “paratype” was collected. The specimen is nonetheless important, for reasons

given below.

Two characters given by previous authors separate C. fergusoni, C. valida and C.

invalida. One is length of aristomere 2: in the description of C . fergusoni it is much

longer than length of aristomere 3, in holotype of C. valida the articles are subequal

in length, and in holotype of C. invalida it is slightly more than half length of

aristomere 3. The second diagnostic character is presence or absence of an ad seta

on mid tibia (character 24). C . fergusoni is cited as lacking this seta, it is long in the

type of C. valida and short though stout in type of C. invalida. In all other respects

the three named species seem not to differ significantly.

Malloch (1930a) knew about both Bezzi’s C. fergusoni and Curran’s C. valida

when he described C. invalida, but finding Curran’s description lacking information

about the mid tibial ad seta, he divided all his material between C. fergusoni and C.

invalida. Malloch apparently did not see any specimens with an aristomere 2

subequal in length to aristomere 3, as decribed for Curran’s C. valida 7

The labelled holotype of C. valida agrees in all respects with its description, but was collected from

Quae st. Ent., 1989, 25 (1,2)

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I strongly suspect that the character states used to separate the three nominal

species are unreliable within this species complex. Length of aristomere 2 is subject

to some variation in some other siphonines, particularly those in which it is

elongate. Under different circumstances, I would consider the mid tibial ad seta as

reliable, but several factors suggest it may not be here. Firstly, Malloch’s (1930a)

specimen’s of “C. fergusoni ” and “C. invalida ” were collected from the same

locality (Sydney), and between the same months (Sept, to Dec.). Secondly, the

MCSN specimen identified (by Bezzi?) as C. fergusoni lacks the mid tibial ad seta

as described for that species, but has an aristomere 2 as described for C. invalida

(also note that this specimen is labelled similarly to the specimens of “C. fergusoni ”

and “C. invalida ” studied by Malloch). Thirdly, the types of the three nominal

species vary in relative length of the mid tibial ad seta, suggesting a pattern of

intraspecific variability. Fourthly, a BMNH specimen from South Australia has an

aristomere two 0.8X length of aristomere 3 and a long mid tibial ad seta, placing it

closest to C. valida, or by Malloch’s criteria in C. invalida.

I have not examined enough material to firmly establish the conspecificity of the

three nominal species discussed above. Neither have I examined the male genitalia

of specimens of all three forms to determine if there are genitalic differences among

them. Also the correlation between aristomere 2 length and presence/absence of a

mid tibial ad seta needs to be studied to determine if these characteristics are

distributed as uniformly as suggested by Malloch. The evidence now available casts

doubt on the correctness of recognizing all three named species as valid species, but

I reserve any change in nomenclature until more specimens can be examined and

my above concerns addressed.

Hosts

Ceromya s.s. species mostly parasitize Macrolepidoptera, without apparent

preference for a particular family (Table 2). A single record for a rearing from a

tenthredinid for Ceromya bicolor (a common European species also recorded from

the Arctiidae and Lasiocampidae) is, if accurate, the only known record of a

siphonine parasitizing a hymenopteran.

Phylogenetics

Species of Ceromya s.s. are not known to possess any synapotypic character

states externally. The group, in addition to members of the C. silacea species group,

is defined externally by the lack of derived features found in other siphonine

lineages (see Recognition section of Ceromya s.l.), and Ceromya s.s. species are

only separable from species of the C. silacea species group by features of the male

'(cont’d) Palmerston on Sept. 1908 by Lichtwardt, and not (as cited by Curran) collected from Palmerston in

1910 by Fred P. Dodd. There is no other indication that this specimen is not the one selected by Curran as

holotype of this nominal species.

Systematics of the Genus Group Taxa of the Siphonini

57

genitalia.

Though members of Ceromya s.s. do not share known synapotypies externally,

one derived characteristic of the male genitalia suggests the group is monophyletic,

i.e. the more or less enlarged membranous area on the anterior surface of the

pregonite, which in most Ceromya s.s. species is partially covered with tiny spinules

(Figs. 39-40 and 77-83; in some species these spinules are only visible at

magnifications of 100X-400X). These spinules are a derived state within the

Siphonini, and so far as known are only present in Ceromya s.s. species and

Goniocera io. The phylogenetic significance of this apotypic state in G. io is

discussed in the Evolution chapter, though for the purposes of this discussion the

state is interpreted as independently derived in G. io and Ceromya s.s.

The size and number of spinules on the pregonite of Ceromya s.s. species varies

from species to species, and they are entirely absent from a few (cf. Figs. 77-83).

Species which lack spinules are, with rare exceptions (as discussed below),

assignable to species groups which possess them (for example, the pregonite is bare

in C. flaviceps and spinulose in several closely related species), so presence of

spinules is here interpreted as the groundplan condition, or is an underlying

synapotypy (as defined in Evolution chapter), of Ceromya s.s. Several Ceromya s.s.

species show modification from the simple spinulose condition, such as New Guinea

sp. 6 (Fig. 78) and Brazil sp. 3 (Fig. 82), but the pregonite of these species is

traceable to the primitive condition through species with intermediate states. More

specifically, the pregonite of certain species suggests that the ring-shaped pregonite

of New Guinea sp. 6 is derivable from a bilobed pregonite of a C. flaviceps-Wkt

ancestor, and the enlarged spines on the pregonite of Brazil sp. 3 are derivable from

an ancestor with a spinulose condition similar to Mexico sp. 5 (Fig. 81; note that the

spined condition of the pregonite in Brazil sp. 3 is distinctly different from the

Actia- type, Figs. 85-86).

The pregonite of C. lutea (Fig. 83) is unique in curving posteriorly. No other

siphonine is known to share this condition, and along with the derived shapes of its

male sternum 5 (Fig. 57) and distiphallus (Fig. 116), this seems to indicate that this

species is not closely related to other known Ceromya s.s. species. C. lutea is

interpreted as a member of Ceromya s.s. because it has tiny spinules on the anterior

(membranous) portion of the pregonite.

Setulation of wing vein R4+5 is a markedly labile character within siphonine

lineages (Table 1), though all Ceromya s.l. species except C. cephalotes, C.

natalensis and C. monstrosicornis have R4+5 setulose beyond r-m. The placement of

these three species in Ceromya was carefully evaluated because the monophyly of

Ceromya s.l. is not well established, and Entomophaga and Proceromyia species are

also characterized by R4+5 not setulose beyond r-m. The male genitalia of C.

cephalotes and C. natalensis were examined, and though spinules are absent from

the pregonite, other features of the male genitalia indicate that the former belongs to

the C. bicolor group and the latter is closely related to C. languidula. Both species

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are therefore retained in Ceromya s.s. The placement of C. monstrosicornis in

Ceromya s.s. is more equivocal because the male genitalia were unavailable for

study and for other reasons discussed below.

Stein described C. monstrosicornis in 1924 in a new monobasic genus,

Stenoparia. Mesnil (1963a) classified Stenoparia as a subgenus of Ceromya , adding

C. nigrohalterata to the taxon. Species of C. (Stenoparia) and C. (Proceromyia)

were separated from those of Ceromya s.s. by not having wing vein R4+5 setulose

beyond crossvein r-m , and the latter (with single species C. macronychia) was

separated from the former by its larger tarsal claws and several other minor

differences (1963a: 829). In the classification adopted here, C. macronychia and

Nipponoceromyia pubioculata (described after Mesnil’ s Palearctic revision) are

included in the genus Proceromyia , and C. nigrohalterata is included in the genus

Entomophaga with E. exoleta (see Phylogenetics sections of Proceromyia and

Entomophaga).

There is insufficient evidence to place C. monstrosicornis in Entomophaga or

Proceromyia (it certainly is not a Goniocera species). This species lacks the

elongate aristomere 1 of Entomophaga or large tarsal claws of Proceromyia , and

does not have a long preapical ad seta on the fore tibia which is common to both.

Yet C. monstrosicornis is the only Ceromya s.l. species with a bare prostemum (as

in Proceromyia ), one of only three Ceromya s.l. species with R4+5 not setulose

beyond r-m , and has a dark, uniformly pruinose abdomen as in Entomophaga and

Proceromyia. Unfortunately, a male specimen of C. monstrosicornis could not be

located in North American or European collections (and the male type could not be

located in the ZMHU), so the phylogenetically important characters of the male

genitalia could not be studied. C. monstrosicornis is left in Ceromya s.s. (not in

Ceromya s.l., as it is certainly not a member of the C. silacea sp. grp.) until its male

genitalia are studied and its relationship to Ceromya , Entomophaga and

Proceromyia established.

The placement of two undescribed species, Australia sp. 3 and Nepal sp. 1, in

Ceromya s.s. also requires explanation. These species differ from other Ceromya s.l.

species in having a long preapical ad seta on the fore tibia. They share this state with

Entomophaga and Proceromyia species and some species of Goniocera and Actia,

so the phylogenetic interpretation of this state in these Ceromya species is relevant

to their placement. These species are certainly not close to Actia, so the discussion

which follows focuses on their possible affinities with the other three genera.

Australia sp. 3 is known only from adult females and first instars, so pertinent

characters of the male genitalia are unknown. Females of this species are unusual

among Ceromya species in having enlarged tarsomere 5 on all legs and a sharply

keeled sternum 7, while the first instar has an uncharacteristically broad labrum.

These states are evidently autapotypies as they are not shared with the non -Ceromya

genera mentioned above. External characteristics of the female favour the placement

of Australia sp. 3 in Ceromya s.s., though it is noted that examination of the male

Systematics of the Genus Group Taxa of the Siphonini

59

genitalia would provide valuable information about this species’ affinities.

The placement of Nepal sp. 1 is enigmatic, as it appears to belong in Goniocera,

Entomophaga, Proceromyia or Ceromya, but lacks the synapotypies of any of these.

Its placement is complicated further by its unusual combination of character states:

(1) haired parafacial (with several larger setulae) in common with all but one

Goniocera species (though the parafacial is more heavily setulose in those species

than in Nepal sp. 1; Fig. 1), Proceromyia pubioculata and Ceromya monstrosicornis

(both with hairs only on parafacial; Fig. 3), (2) long preapical ad seta on fore tibia

(as mentioned above), (3) haired prostemum (shared with most siphonines, but not

Proceromyia and some Goniocera species), (4) R4+5 setulose beyond r-m (as in

Goniocera species and almost all Ceromya species, but not Entomophaga and

Proceromyia species), and (5) uniformly pruinose abdomen (as in Goniocera,

Proceromyia and Entomophaga species, and C. monstrosicornis but not other

Ceromya species). I have been unable to polarize these states with confidence, so

cannot interpret their phylogenetic significance (see also Evolution chapter under

Phylogenetics of non -Siphona s.l. siphonine lineages). The male genitalia of Nepal

sp. 1 do not closely resemble those of the aforementioned non -Ceromya genera, but

similarly lack the typical spinules on the pregonite possessed by most Ceromya s.s.

species (male genitalia of C. monstrosicornis unavailable for study) and has an

aedeagus which cannot be placed into a Ceromya s.s. species group (though very

different from the typical shape in the Ceromya silacea species group).

Such an array of character states in Nepal sp. 1 is not easily interpretable, and

any placement at this time is tentative at best. This species seems to share more

states with species of Ceromya s.s. (see description of Ceromya s.s., which includes

Nepal sp. 1) than with species of the other genera discussed, so is placed here for the

present. A possible relationship with C. monstrosicornis needs to be investigated.

Geographic distribution

Thirty-nine described species are assigned to Ceromya s.s., of which only eight

are New World in distribution (see Sabrosky and Amaud 1965 for ranges of species

north of Mexico, Guimaraes 1971, Cortes 1967, and Cortes and Hichins 1969 for

ranges of species south of the United States). However, preliminary study of

specimens of New World Ceromya s.s. indicates that the number of described

species is a significant underestimate of the true diversity of the group in the New

World. I estimate from examined material that there are about 15 undescribed New

World species, almost all Neotropical in distribution.

Most accurately known are the British (Crosskey 1976b) and European

(Draber-Moriko 1981, Herting 1984) faunas, consisting of five described species. A

key for recognition of all of these species has not been published, though they are

treated in part in Mesnil (1963a) and Herting (1977). The fauna of Asia is less well

documented, and appears to be depauperate. C. bicolor is the only Ceromya s.s.

species recorded from central Asia (Richter 1971, 1975, 1980), and the only other

Quaest. Ent., 1989, 25 (1,2)

60

O’Hara

described eastern Palearctic species, C. pruinosa, is apparently restricted to Japan

(though Herting 1984 cites this species as a possible synonym of C. bicolor).

However, I have seen specimens of several undescribed species of Ceromya s.s.

collected from Nepal.

Greatest diversity of described Ceromya s.s. species in the Old World is recorded

from the tropics: eight species in the Afrotropical region and 13 in the Oriental

region (distributions given in Crosskey 1980, 1976a, respectively; key to Philippine

species in Dear and Crosskey 1982, though these authors include S.

(Aphantorhaphopsis) in their concept of Ceromya). Only three species are described

from Australia (Crosskey 1973). Still, the described portion of the Old World fauna

of Ceromya s.s. belies its true diversity: Shima (pers. comm.) has tentatively

recognized close to 20 new species in the Oriental region, and I have seen

specimens of numerous undescribed species from Australia and Africa.

From a cosmopolitan perspective Ceromya s.s. can best be considered a tropical

group, with relatively few species in the Nearctic and Palearctic regions. The genus

still requires a great deal of descriptive work, especially with respect to its tropical

elements.

List of described species included in Ceromya sensu stricto

S amblycera (Aldrich), 1934: 132 ( Actia ). Holotype male, Argentina: Bariloche

(USNM). Holotype examined. New combination, moved from Actia.

N americana (Townsend), 1892: 69 ( Thryptocera ). Holotype male (not female),

USA: D.C., Washington (UKL). Holotype examined,

syn. elyii (Walton), 1914: 91 ( Polychaetoneura ). Three female syntypes,

USA: Connecticut, East River (USNM). — Curran, 1933a: 5.

Syntypes examined.

A amicula Mesnil, 1954: 40. Holotype male, Zaire: Bambesa (MRAC). Holotype

examined.

O apicipunctata (Malloch), 1926: 510 {Actia). Holotype male, Philippines: Luzon,

Benquet (USNM). Holotype examined.

O bellina Mesnil, 1957: 44. Holotype male, Burma: Kambaiti (ZMU). Holotype

examined.

P bicolor (Meigen), 1824: 354 ( Tachina ). Holotype male, no locality data

(MNHN).

syn. testacea Robineau-Desvoidy, 1830: 88. Type(s), France: Lille

(lost). — Herting, 1974: 18.

rufina (Zetterstedt), 1838: 641 {Tachina). Holotype female, Sweden:

Dalecarlia (UZI). — Herting, 1984: 121.

fasciata (Stein), 1924: 132 {Actia). Lectotype female (by designation

of Herting, 1977: 10), Yugoslavia: Sarajevo (NMBA). — Considered

a possible color variant of bicolor Mg. by Herting, 1977: 10.

Systematics of the Genus Group Taxa of the Siphonini

61

A buccalis (Curran), 1933c: 163 ( Actia ). Holotype male, Zimbabwe: Gatooma

(AMNH). Holotype examined.

O capitata Mesnil, 1957: 42. Holotype male, Burma: Kambaiti (ZMU). Holotype

examined.

O cephalotes Mesnil, 1957: 40. Holotype male, Burma: Kambaiti (ZMU).

Holotype examined.

A cibdela (Villeneuve), 1913: 35 (Actia). Lectotype male (by designation in

text), Nigeria: Oshogbo (CNC). Lectotype examined,

syn. cibdella. Incorrect subsequent spelling of cibdela Villeneuve

(Curran, 1927a: 323).

S cornuta (Aldrich), 1934: 131 (Actia). Holotype male, Chile: Angol (USNM).

Holotype examined. New combination, moved from Actia.

P dilecta Herting, 1977: 10. Holotype male, Switzerland: Gordola (SMNS).

Holotype examined.

O dubia (Malloch), 1930b: 146 (Actia). Holotype female, Malaysia: Selangor

(BMNH). Holotype examined.

A femorata Mesnil, 1954: 38. Holotype male, Zaire: Bambesa (MRAC). Holotype

examined.

U fergusoni (Bezzi), 1923b: 657 (Schizotachina). Holotype male, Australia: New

South Wales, Sydney (type missing according to Crosskey, 1973: 137).

syn. fergussoni. Incorrect subsequent spelling of fergusoni Bezzi (Curran,

1927b: 355-356).

P flaviceps (Ratzeburg), 1844: 172 (Musca (Tachina)). Type, Germany (lost).

syn. flaviceps (Stein), 1924: 134 (Actia). — Objective synonym; see

Herting, 1982: 8 and 1984: 190 (note 94), cf. Herting, 1977: 9.

P flaviseta (Villeneuve), 1921: 45 (Actia). Male syntype from Berlin, Germany,

female syntype from Samara, USSR (CNC). Syntypes examined.

O hirticeps (Malloch), 1930b: 146 (Actia (Pseudactia)). Holotype male, Malaysia:

Kedah Peak (BMNH). Holotype examined.

U invalida (Malloch), 1930a: 305 (Actia (Schizoceromyia)). Holotype male,

Australia: New South Wales, Sydney (SPHTM). Holotype examined.

A languidula (Villeneuve), 1913: 36 (Actia). Two male syntypes, Nigeria:

Oshogbo (BMNH and CNC). Syntypes examined.

A languidulina Mesnil, 1977b: 178. Holotype female, Madagascar:

Ambohitantely (MNHN).

O latipalpis (Malloch), 1930b: 145 (Actia). Holotype female, Malaysia: Kedah

Peak (BMNH). Holotype examined.

A lavinia (Curran), 1927a: 324 (Actia). Holotype female, South Africa: Natal,

Clan Syndicate (PPRI). Holotype examined.

O longimana Mesnil, 1957: 38. Holotype female, Burma: Kambaiti (ZMU).

Holotype examined.

S lutea (Townsend), 1927: 283 (Actinactia). Holotype male, Brazil: Sao Paulo

Quaest. Ent., 1989, 25 (1,2)

62

O’Hara

(USNM). Holotype examined. New combination.

A luteicornis (Curran), 1933c: 162 ( Actia ). Holotype male, southern Zimbabwe

(BMNH). Holotype examined.

O maculipennis (Malloch), 1930b: 141 {Actia). Holotype male, Malaysia:

Selangor (BMNH). Holotype examined.

P monstrosicornis (Stein), 1924: 128 ( Stenoparia ). Holotype male, German

Democratic Republic: Mecklenburg (ZMHU; not located, possibly lost),

syn. monstruosicornis. Incorrect subsequent spelling of monstrosicornis

Stein (Mesnil, 1963a: 829, 831).

A natalensis (Curran), 1927a: 325 {Actia). Holotype male, South Africa: Natal,

Cramond (PPRI). Holotype examined.

N Ontario (Curran), 1933a: 4 {Actia). Holotype female, Canada: Ontario, Lake of

Bays, Norway Point (CNC). Holotype examined.

N palloris (Coquillett), 1895b: 50 {Lasioneura). Lectotype male (by designation

of Coquillett, 1897: 58), USA: New Hampshire (USNM). Lectotype examined.

O portentosa Mesnil, 1957: 43. Holotype female, Burma: Kambaiti (ZMU).

Holotype examined.

P pruinosa Shima, 1970c: 188. Holotype male, Japan: Hokkaido, Berabonai

(BLKU). Paratype examined.

O punctipennis (Malloch), 1930b: 140 {Actia). Holotype male, Malaysia: Kedah

Peak (BMNH). Holotype examined.

O punctum (Mesnil), 1953: 107 {Actia). Holotype male, China: Canton (BMNH).

Holotype examined.

O rotundicornis (Malloch), 1930b: 145 {Actia). Holotype male, Malaysia: Pahang,

Fraser’s Hill (BMNH). Holotype examined.

S subopaca (Aldrich), 1934: 133 {Actia). Holotype male, Argentina: Bariloche

(BMNH). Holotype examined.

S unicolor (Aldrich), 1934: 133 {Actia). Holotype male, Argentina: Bariloche

(BMNH). Holotype examined.

U valida (Curran), 1927b: 356 {Actia {Schizactiana)). Holotype male, Australia:

Queensland, Palmerston (DEI). Holotype examined.

Nomen dubium

P erythrocera Robineau-Desvoidy, 1830: 87. Type(s), France (lost).

List of examined, undescribed, species included in Ceromya sensu stricto

Ceromya nr. punctipennis: Two males from se. Popondetta, New Guinea (BLKU).

Ceromya Australia sp. 1: Two males from Queensland (CNC, DPI).

Ceromya Australia sp. 2: Three males, one female from Queensland (DPI).

Ceromya Australia sp. 3: One female from Mt. Glorious, Queensland (DPI).

Ceromya Australia sp. 5: One male, two females from Mt. Glorious, Queensland (DPI).

Systematics of the Genus Group Taxa of the Siphonini

63

Ceromya Brazil sp. 1: Males and females from Nova Teutonia (CNC, USP).

Ceromya Brazil sp. 3: One male from Amazonas (INPA).

Ceromya Brazil sp. 4: One male from Nova Teutonia (CNC).

Ceromya Chile sp. 1: One male and one female from Magellanes (CNC).

Ceromya Chile sp. 2: One male from Isla de Chiloe (CNC).

Ceromya Mexico sp. 1: Three males and two females from Durango (CNC).

Ceromya Mexico sp. 2: One male each from Colima (UCB) and Chiapas (CNC).

Ceromya Mexico sp. 3: One male from Chiapas, one female from Veracruz (CNC). Two possibly

conspecific males from SE Brazil (CNC).

Ceromya Mexico sp. 4: One male from Chiapas (CNC).

Ceromya Mexico sp. 5: One male from Veracruz (CNC). One male and several females possibly conspecific

from SE Brazil (CNC, USP).

Ceromya Nepal sp. 1 : Males and females from Nepal (CNC).

Ceromya Nepal sp. 3: Two males, one female from 28°00’N 85°00’E (CNC).

Ceromya Nepal sp. 4: One female from 28°00’N 85°00’E (CNC).

Ceromya Nepal sp. 5: One male, one female from Kathmandu (CNC).

Ceromya New Guinea sp. 4: Two males from se. Popondetta (BLKU).

Ceromya New Guinea sp. 5: One male each from se. Popondetta and Nabire (BLKU).

Ceromya New Guinea sp. 6: Two males from Wau (BLKU).

Ceromya New Guinea sp. 11: Two males from Mt. Kaindi (BLKU).

Ceromya New Guinea sp. 15: One male from Mt. Kaindi (BLKU).

Ceromya New Guinea sp. 18: One male each from Mt. Kaindi and se. Mt. Giluwe (BLKU).

Ceromya Peru sp. 1: One male from Quincemil, Cuzco (CNC).

Ceromya U.S. sp. 1: One male from Adair Co., Missouri (MSU)

Ceromya silacea (Meigen) species group

Figs. 35, 37, 52, 84, 112, 148, 159.

Recognition

Male genitalic features provide the only characteristics by which members of the

Ceromya silacea species group (a strictly Old World taxon) can be distinguished

from Ceromya s.s. species. Species of C. silacea species group are unique within the

Siphonini in possessing posteriorly on the distiphallus an infolded and sclerotized

structure (Fig. 112) - in other siphonines the posterior margin is sclerotized or

membranous (the latter in all but a very few species), but not infolded.

Ceromya silacea is one of the species with greatest development of the infolded

structure on the distiphallus. As shown in Fig. 112, this structure is extended

anteriorly to near the anterior margin of the distiphallus (note too that the

distiphallus is extensively membranous anteriorly and laterally in this species) and

projects posteriorly from between the posterolateral margins. In some other species

of this species group the infolded structure is smaller.

The pregonite of species of the C. silacea species group is long and sickle-like in

most species (Fig. 84) and shorter and apically rounded in others; the membranous

portion is not expanded and spinules are absent (cf. Ceromya s.s. section and Figs.

39-40, 77-83).

Quaest. Ent., 1989, 25 (1,2)

64

O’Hara

Description

Length: 3. 0-5. 5mm.

Head. — Five frontal setae in most species, three (C. silacea) or four in a very few; normal arrangement.

Anterior proclinate orbital seta longer than posterior one. Eye size of male medium-large to large, 0.77-0.86

head height; size in female subequal to that of male. Flagellomere 1 of male medium-short to medium

length, 0.47-0.56 head height; shape from linear to large and almost triangular, not bifid. Flagellomere 1 of

female smaller than in male. Aristomere 1 short. Aristomere two 1.5-5X longer than wide. Aristomere 3

rather short to long and evenly tapered; micropubescent to pubescent. Clypeus U-shaped. Palpus short,

clavate. Proboscis with prementum short, labella padlike.

Thorax (Figs. 35, 37). — Prostemum setulose. Lower proepimeral seta weak, not directed downward.

Katepistemum bare anterior to mid coxa. Lower katepistemal seta much shorter than upper anterior seta in

most species, subequal in length to it in three closely related species: C. normula, C. similata and C.

varichaeta. Three or four postsutural dorsocentral setae. Upper part of anepistemum with single setula. Fore

tibia with preapical ad seta much shorter than d seta. Mid tibia with one ad seta. Mid femur with pilose patch

on anterior surface from about midlength to distal end in males of C. normula and C. varichaeta (Fig. 35).

Tarsomeres normal in size, claws small. Wing: CuA] with distal portion 0.29-0.47X length of proximal

portion (mean 0.36); anal vein not extended to wing margin. Wing setulae: R t dorsally bare, or distally or

entirely setulose, ventrally bare or distally setulose; /?4+5 setulose from base to beyond r-m\ CuAx bare or

setulose.

Abdominal terga 1-5. — Abdomen ovoid in shape. T]+2 without median marginal setae; lateral marginal

setae absent to strong. T3-T5 average or with weak lateral discal setae.

Male genitalia (Figs. 52, 84, 112). — S5 with, inner (or posterior) margins of processes obtusely angled

to almost transverse (Fig. 52 and Shima 1970c, figs. 3d,e); apical lobe undifferentiated to rounded; median

lobe rounded or pointed in most species, slightly flattened medially in C. varichaeta (but not to degree found

in Goniocera)-, without accessory lobe; processes very sparsely setulose (most species) to densely setulose

(C. varichaeta). T6 present as pair of small lateral sclerites. Ejaculatory apodeme with fan-shaped portion

half to subequal width of hypandrial apodeme. Pregonite in profile long and sickle-like in most species (Fig.

84), apically rounded in C. varichaeta-, bare. Epiphallus absent from most species, present in C. varichaeta.

Distiphallus (Fig. 112) broad in profile, with short to long posterolateral arm except in C. varichaeta, and

unique infolded and sclerotized structure formed from posterior surface (large in most species, very small in

C. varichaeta ) which is deeply U-shaped in posterior view. Postgonite large, apically bilobed. Surstylus

average length to long, straight in a few species {e.g. Shima 1970c, fig. 5b), thin and curved posteriorly in

most (e.g., op. cit., fig. 5a); with long hairs along length in C. varichaeta (also found in C. languidula of

Ceromya s.s .); basally fused with epandrium or free (C. varichaeta). Cerci rather short to average length,

posteriorly deeply inflexed at midlength (e.g., op. cit., fig. 5b) to gently curved or almost straight (e.g., op.

cit., fig. 5a); moderately to densely setose on basal half.

Examined male genitalia of: C. mellini, C. silacea, C. varichaeta, Australia sp. 4 and Nepal sp. 2. Examined

published figures of: C. dorsigera and C. silacea (both in Shima 1970c, figs. 3d,e, 5).

Female genitalia (Fig. 148). — (Only C. silacea examined.) Medium length (slightly extensible). S6 with

average length hairs. T6 present as two lateral sclerites; enclosing spiracles of segment 6 in anteroventral

portion. S7 with anterior apodeme; without posteromedial keel. T7 present as two lateral sclerites; spiracles of

segment 7 enclosed within anterior portion of T7. S8 distinctly developed, haired. 7j0 present as two distinct

sclerites.

Examined female genitalia of: C. silacea.

Taxonomic changes

Status of Ceromya similata Mesnil. — Mesnil described Ceromya similata in

1954 as a sympatric subspecies of C. varichaeta (Curran), noting that it differs from

the nominal subspecies primarily in having Rx distally rather than entirely setulose

dorsally. As explained elsewhere (e.g. Taxonomic changes section of Siphona

(Siphona)), I do not accept the concept of sympatric subspecies; therefore, C.

varichaeta similata must either be elevated to species status or declared conspecific

with C. varichaeta.

Systematics of the Genus Group Taxa of the Siphonini

65

I have examined the holotypes of C. varichaeta and C. similata , and a closely

related species C. normula (the latter placed in Peribaea by Crosskey, 1980). C.

varichaeta and C. normula are based on males which share a striking synapotypy: a

pilose patch anteriorly on the mid femur (Fig. 35). The types differ little except that

/?! is dorsally setulose entirely in C. varichaeta and only distally in C. normula. This

difference in Rx setulation is very reliable as a species specific character (in contrast

to the states Rx distally bare or setulose), so I accept C. varichaeta and C. normula

as valid species. Setulation of R{ in the holotype of C. similata matches that of C.

normula , not C. varichaeta , and might be conspecific with that species. However,

the type of C. similata is female (thus lacking the male-linked pilose patch on mid

femur), so is difficult to compare critically with the male types of the other two

nominal species. Though C. similata has been associated with C. varichaeta in

publications, it is more likely conspecific with C. normula or a valid species. I treat

C. similata here as a valid species pending examination of additional male and

female specimens of the three nominal species.

Hosts

Hosts have only been recorded for two species, and both belong to the Noctuidae

(Table 2).

Phylogenetics

Adults of the C. silacea species group are similar to those of Ceromya s.s.

externally, but differ in two important male genitalic characteristics. First, they lack

spinules on the membranous anterior portion of the pregonite, which are present in

most Ceromya s.s. species and are interpreted as synapotypic of that group. Second,

they have a uniquely infolded posterior margin on the distiphallus (Fig. 112), which

is interpreted as synapotypic of the C. silacea species group (monophyly of

Ceromya s.l. discussed in Evolution chapter).

In C. silacea, C. mellini, Australia sp. 4 and Nepal sp. 2, the infolded region of

the distiphallus is very large (Fig. 1 12). In C. varichaeta the infolded region is small

and nearer the apex, and perhaps represents a more primitive condition. The male

genitalia of the other species placed in this group were not examined, but these

species are thought to belong here because they appear very similar externally to

certain well established members of this group. These species should be removed

from the C. silacea group if they are found to lack the synapotypy (of the

distiphallus) of this group.

The labrum is hook-like in the first instar of C. silacea (Fig. 159) and

hatchet-like or intermediate in other examined Ceromya s.l. species (see O’Hara in

press “a”). It remains to be determined whether this feature is another synapotypy of

the C. silacea species group.

Quae st. Ent., 1989, 25 (1,2)

66

O’Hara

Geographic distribution

The eight described species of the Ceromya silacea species group are exclusively

Old World in distribution. The centre of diversity is the Old World tropics (see

Crosskey 1976a and 1980 for ranges of Oriental and Afrotropical species; none

described from Australia), with only three species recorded from the Palearctic

region: C. dorsigera (described from Switzerland and recorded from Japan by

Shima 1970c), C. silacea (widespread, with records from England (Crosskey

1976b), Eurasia (Mesnil 1963a, Shima 1970c, Richter 1971, 1976b, 1980,

Draber-Moriko 1981), and southern India (examined specimens from CNC)), and C.

pendleburyi (SE Asia and Japan, Mesnil 1963a and Shima 1970c). The three

Palearctic species are keyed in Shima (1970c). In addition to the described species I

have seen specimens of a new species from Nepal (Nepal sp. 2) and another from

Australia (Australia sp. 4) [numbers used here for new species refer to Ceromya

s.ll

List of described species included in the Ceromya silacea species group

P dorsigera Herting, 1967a: 8. Holotype male, Switzerland: Gordola (SMNS).

Holotype examined.

O mellina (Mesnil), 1953: 109 ( Actia ). Holotype male, Burma: Kambaiti (ZMU).

Holotype examined.

A normula (Curran), 1927a: 322 {Actia). Holotype male, South Africa: East

London (PPRI). Holotype examined. New combination, moved from Actia.

O patellicornis Mesnil, 1957: 40. Holotype male, India: Darjeeling (BMNH).

Holotype examined.

0,P pendleburyi (Malloch) 1930b: 144 {Actia). Holotype male, Malaysia: Pahang

(BMNH). Holotype examined.

0,P silacea (Meigen), 1824: 355 {Tachina). Holotype male, no locality data

(MNHN).

syn. siebeckii (Sintenis), 1897: 151 {Thryptocera). Holotype female,

Estonia: Pamu (not located).— Herting, 1984: 121.

A similata Mesnil, 1954: 39 (as subspecies of Ceromya varichaeta (Curran)).

Holotype female, Zaire: Tshumba (MRAC). Holotype examined. New status.

A varichaeta (Curran), 1927c: 6 {Actia). Holotype male, Zaire: Faradje (AMNH).

Holotype examined.

List of examined, undescribed, species included in the Ceromya silacea species

group

Ceromya Australia sp. 4: One male, one female from Queensland (DPI).

Ceromya Nepal sp. 2: Males from 28°00’N 85°00’E (CNC).

Systematics of the Genus Group Taxa of the Siphonini

67

Genus Actia Robineau-Desvoidy

Figs. 8-10, 21, 23, 29, 31, 33, 41-42, 58-59, 85-86, 1 17-119, 151-152, 160.

Actia Robineau-Desvoidy, 1830: 85. Type-species, Roeselia lamia Meigen, 1838, by designation of

I.C.Z.N., 1987: 71 (Opinion 1432).

Thryptocera Macquart, 1834: 310. Type-species, T. bicolor Macquart, by designation of Townsend, 1916:

624) = Tachina crassicornis Meigen, 1824. — Herting, 1976: 3.

Tryptocera. Variant spelling of Thryptocera.

Gymnophthalma 1838, Lioy, 1864: 1341. Type-species, Tachina crassicornis Meigen, 1824 (monotypy).

Gymnopareia Brauer and Bergenstamm, 1889: 103 (35). Type-species, Tachina crassicornis Meigen, 1824

(monotypy).

Gymnoparia. Variant spelling of Gymnopareia.

Actiopsis Townsend, 1917: 121. Type-species, A autumnalis Townsend, 1917 (original designation).

Setasiphona Townsend, 1934: 248. Type-species, Actia siphonosoma Malloch, 1930 (original designation).

Recognition

Actia is a diverse genus of cosmopolitan distribution. With few exceptions,

members are recognized externally by the presence of a row of hairs on the

katepistemum anterior to the mid coxa {cf. Figs. 33 and 34). Among other

siphonines, only the European species Entomophaga exoleta shares this state, and is

distinguished from Actia by its elongate aristomere 1 (Fig. 4), long preapical ad seta

on fore tibia, and features of the male genitalia (especially sternum 5 - cf. Figs. 50

and 58-59). Five Actia species, A. completa and A. magnicornis from Malaysia and

A. parvis eta, A. nr. parviseta and Australia sp. 5 from eastern Australia, either lack

this row of katepistemal hairs or the number of hairs is reduced. These species have

two setulae on the upper portion of the anepistemum (see below), and males are

additionally recognized as members of Actia by their V-shaped sternum 5 (similar to

Figs. 58-59) and spined pregonite (similar to Figs. 85-86).

The upper portion of the anepistemum has two setulae in almost all Actia species

(Fig. 31; one setula in most other siphonines. Fig. 32). Two setulae are present in a

few Peribaea species and a few species in different Siphona s.l. lineages, but this

state seems to be restricted to Actia among siphonines in which the anal vein does

not extend to the wing margin (Table 1).

Three Actia species, A. completa and A. fulvicauda from Malaysia and A.

chrysocera from the Seychelles Islands, are the only known siphonines other than

Peribaea and Siphona s.l. species to have the anal vein extended to the wing margin.

A. fulvicauda and A. chrysocera possess a row of katepistemal hairs as in most other

Actia species, but A. completa has an incomplete row. A. completa is recognized as

an Actia species by its V-shaped sternum 5, spined pregonite and two anepistemal

setae (as mentioned above).

Males of most Actia species have a more or less V-shaped sternum 5, with little

or no constriction of the median lobes above the median cleft (Figs. 58-59). Sternum

5 departs slightly from this shape in a few species, but even in these it more closely

resembles the sternum 5 of other Actia species than non-congeneric siphonines.

Quaest. Ent., 1989, 25 (1,2)

68

O’Hara

The only feature apparently universal among (and unique to) Actia species is a

J-shaped, spinose pregonite (Figs. 41-42 and 85-86). All examined male genitalia of

Actia species were of this type, including species mentioned above as having

atypical external features.

Description

Length: 2.5-6.0mm.

Head (Figs. 8-10). — Five frontal setae (rarely four), normal arrangement. Anterior proclinate orbital

seta longer than posterior one in almost all species. Eye of male small to large, 0.65-0.89 head height; eye of

female subequal to or slightly smaller than in male. Flagellomere 1 of male markedly varied in length,

0.43-0.75 head height; linear to broad; bifid in only one known species, A. yasumatsui (Shima 1970b, fig. 1).

Flagellomere 1 of female smaller than in male or subequal in size; not bifid. Aristomere 1 short. Aristomere

2 varied from 1.5-5X longer than wide, relatively short (2-3X) in most species. Aristomere 3 short and

thickened to near tip, to long and evenly tapered; almost bare to short plumose. Clypeus varied from narrow

and enclosed in membrane to broadened or U-shaped. Palpus short in most species, long in some species

with elongate proboscis; enlarged apically in females of a few species; clavate. Prementum short to long, in a

few Old World species of latter slender and elongate like typical Siphona species (Fig. 9). Labella also

markedly varied, padlike or slightly lengthened (like in Siphonopsis) in most species, quite elongate (half

head height or longer) and with numerous pseudotracheae and flexible in life in a few species (e.g. A.fallax,

A. jocularis, A. longilingua ); in a very few Old World species labella as in Siphona species (Fig. 9): about

head height in length, basal portion inflexible in life, with reduced number of pseudotracheae apically (e.g.

A. malaisei).

Thorax (Figs. 21, 23, 29, 31, 33). — Prostemum setulose in almost all species (apparently bare only in A.

nigra). Lower proepimeral seta weak or absent. Most species with row of hairs on katepistemum directly

anterior to mid coxa extended upward almost to lower katepistemal seta (Fig. 33); several hairs in lower

comer only (i.e. groundplan condition of Siphonini) in A. parviseta and A. nr. parviseta, and several hairs in

lower comer to row extended halfway to lower katepistemal in A. completa, A. magnicornis and Australia

sp. 5. Lower katepistemal seta shorter (in most species much shorter) than upper anterior seta (Fig. 31). Four

postsutural dorsocentral setae in most species, three in a few. Upper part of anepistemum with two setulae in

most species (Fig. 31), with single setula in a very few. Fore tibia with preapical ad seta much shorter than d

seta in most species, ranging to about 0.75 length of d seta in a few species (known in A. infantula, A. lamia

and A. nudibasis), and subequal to d seta in a few species (known in A. eucosmae, A. nigriventris, A.

parviseta , and several undescribed Australian species). Mid tibia with one ad seta in almost all species

(markedly reduced or absent from A. eucosmae, A. parviseta, A. perdita and Australia sp. 3). Tarsomeres

normal in size, or tarsomere 5 of fore leg broadened (known only in female of A. tarsata, though slightly

larger than average in females of a few other species), tarsomere 5 of all legs slightly enlarged in female of

A. nigriventris ; claws short. Wing (Fig. 21) with vein M complete in most species, slightly developed or

absent after bend in some (e.g. A eucosmae, A. exsecta, A. lamia, A. munroi, A. nigriventris, A. perdita, A.

pulex, A. rufescens and A. takanoi)', vein CuAx with distal portion 0.25- 1.6X length of proximal portion (in

most species short, 0.3-0.7; mean 0.56); anal vein not extended to wing margin, except in A. completa, A.

fulvicauda and faintly in A. chysocera. Wing setulae: /?, dorsally bare, or distally or entirely setulose,

ventrally bare or distally setulose; R4+5 setulose from base to or beyond r-m\ CuAl bare or setulose; unusual

patterns in a few species with additional veins setulose: A. ciligera (R2+ 3, R4+5 and M setulose dorsally and

ventrally) A.fallax (M setulose ventrally) and A. gratiosa (Sc setulose dorsally).

Abdominal terga 1-5. — Abdomen ovoid in shape in most species, slightly elongate in a very few. Tx+2

without median marginal setae; lateral marginal setae absent from most species, strong in a few. TyTs

average in most species, with weak lateral discal setae in a few (particularly among species in Holarctic

region).

Male genitalia (Figs. 41-42, 58-59, 85-86, 117-119). — S5 little varied, processes elongate, rounded or

pointed posteriorly, inner margins approximately V-shaped in most species (Fig. 58), obtusely angled in a

very few (Fig. 59; rarely as much as in other siphonines); median lobe undifferentiated (Fig. 58) or scarcely

differentiated (Fig. 59); median cleft scarcely constricted posteriorly by median lobes in most species (Fig.

58), distinctly constricted by rounded (Fig. 59) to elongate median lobes in a few; processes moderately

setulose with several pair of large setae. T6 slightly sclerotized, generally continuous dorsally. Ejaculatory

Systematics of the Genus Group Taxa of the Siphonini

69

apodeme varied from small to very large (0.5-2.0X width of hypandrial apodeme), in most species width of

fan-shaped portion subequal to width of hypandrial apodeme. Pregonite (Figs. 85-86) broad subapically and

approximately J-shaped, outer surface short spinose on apical half or less; asetose. Epiphallus absent.

Distiphallus (Figs. 117-119) with posterior margin partially developed in some species, absent from others;

not laterally broadened as in Peribaea species; in profile from broad and truncate apically to very narrow

and sharply pointed apically (Fig. 119), with numerous forms between. Postgonite in most species apically

broad, rounded or truncate, in a few species narrow or intermediate in width. Surstylus short to long, shape

varied relatively little, narrow or broad in a few species, intermediate in most; medium to long hairs basally

in some species of an Old World group; basally free from epandrium. Cerci varied little, short to average

length, sharply inflexed at midlength in some species, smoothly curved posteriorly in most; moderately

setose on basal half.

Examined male genitalia of: A. autumnalis, A. brevis, A. completa, A. diffidens, A. infantula, A. interrupta,

A. lamia, A. longilingua, A. magnicornis, A. malaisei, A. nitidella, A. nudibasis, A. parviseta, A. pilipennis,

A. rufescens, Australia spp. l-2,4-5, Jamaica sp. 1, Liberia sp. 1, Mexico sp. 1, Nepal spp. 1-2, New World

spp. 1-19 and Uganda sp. 1. Examined published figures of: A. lamia (Andersen 1983, fig. 25), A. darwini,

A. painei (both in Crosskey 1962, figs. 1-6), A.jocularis, A. nigra (both in Shima 1970c, figs. 2, 3b,c, 4), A.

pokharana and A. yasumatsui (both in Shima 1970b, fig. 2).

Female genitalia (Figs. 151-152). — Short and markedly uniform and unspecialized, with only minor

differences among species. S6 with average length hairs. Spiracles of segment 6 enclosed in T6; in some

species T6 forming a broad sclerite narrowly discontinuous dorsally, in others represented by two small

lateral sclerites, in a few species unsclerotized. S7 without anterior apodeme except for short projection in A.

crassicornis (Andersen 1983, fig. 12); without posteromedial keel. T7 absent or present only as two small

lateral sclerites; spiracles of segment 7 in membrane between segments 6 and 7. S8 distinctly developed,

haired, with lateral edges curved inward. T10 distinctly developed as median sclerite, in most species

subequal in size to S8.

Examined female genitalia of: A. diffidens, A. interrupta, A. lamia, A. malaisei, A. nr. parviseta, Australia sp.

1, New World spp. 4-5 and Zaire sp. 1. Examined published figures of: A. crassicornis and A. lamia (both in

Andersen 1983, figs. 11, 12).

Taxonomic changes

Synonymy of Actia brevis Malloch with Actia darwini Malloch. — The male

holotypes of these nominal species were examined and compared. They differ

primarily in abdominal coloration and setulation of vein CuA{, but these

characteristics vary within some species and the degree of difference present here is

not unusual among conspecifics. I also examined other specimens from several

localities in eastern Australia, and these exhibit states intermediate between those in

the A. brevis and A. darwini types. I conclude that these types belong to the same

species, and here synonymize Actia brevis Malloch (1930a) with Actia darwini

Malloch (1929b).

Actia parviseta Malloch (Fig. 10). — Malloch described this species in 1930 and

placed it in his widely defined Actia. Later Crosskey (1966: 109) shifted it to

Ceromya, presumably because it lacks a row of hairs on the katepistemum anterior

to the mid coxa - a characteristic usually diagnostic for Actia. Dissection of the male

genitalia of the type revealed that they possess two Actia synapomorphies: a

V-shaped S5 and spinose pregonite. Furthermore, the female reproductive system of

a specimen of a closely related species (A. nr. parviseta) was examined and

contained first instars of the Actia type (i.e. dorsal cornu lacking from the

cephalopharyngeal skeleton; Fig. 160 and O’Hara in press “a”). It thus appears that

Actia parviseta and a few other species (A. completa, A. magnicornis, A. nr.

Quae st. Ent., 1989, 25 (1,2)

70

O’Hara

parviseta and Australia sp. 5) have a bare or sparsely haired katepistemum anterior

to the mid coxa, while still possessing all other known synapomorphies of Actia.

Hosts

The hosts of Actia are better known than those of other siphonines, with records

for about 20 Actia species (Table 2). Hosts belong predominantly to the Tortricidae,

but a variety of other Microlepidoptera and a number of Macrolepidoptera are also

parasitized. Host larvae vary in habit from small leaf miners and rollers to large,

bare to hairy caterpillars which feed openly on vegetation. There is one record of an

Actia species parasitic on a pyrrhocorid bug, Dysdercus sp. This record is doubtful,

as Hemiptera are almost exclusively parasitized by phasiines among the Tachinidae.

Phylogenetics

A row of hairs on the katepistemum anterior to the mid coxa has long been

considered a diagnostic feature of Actia. It is now apparent that several species

belonging to the Actia lineage lack this row of hairs (see Recognition and

Description sections). The relationships of these species to other Actia species is

unknown, so the absence of a row of katepistemal hairs from these species cannot

be positively polarized as primitive or derived. If derived, then these species must be

descended from ancestors possessing a row of katepistemal hairs, and I consider this

explanation the more reasonable at this time. This explanation is hypothesized

because: 1) no other character states support the primitive position of these species

within Actia, and 2) several of these species have a variable number of katepistemal

hairs, ranging from a few to a nearly complete row; the varied extent of this row is

more parsimoniously interpreted as a reversal from a complete to incomplete row

rather than as an independent gain from an incomplete to complete row. Thus a row

of hairs on the katepistemum is here considered a synapotypy of Actia species, with

absence of this row in several Actia species interpreted as resulting from reversal(s)

from the apotypic state.

Almost all Actia species have two setulae on the upper portion of the

anepistemum (Fig. 31). Most other siphonines have one (Fig. 32), though two are

present in a few species of Peribaea and a few species of most supraspecific taxa of

Siphona s.l. (Table 1). This distribution suggests that two anepistemal setulae is the

derived state within the tribe, though its many independent acquistions weaken its

value as a synapotypy of Actia.

Several species of Actia have the anal vein extended to the wing margin (see

Recognition and Description sections), but possess the derived genitalic states of

Actia, and all but one of these (A. completa ) have a katepistemal row of hairs.

Extension of the anal vein to the wing margin probably arose independently in Actia

and the Peribaea-Siphona s.l. lineage.

Sternum 5 is approximately V-shaped in most Actia species, and derivable from

that shape in the others. The pregonite is J-shaped and spinose, and these states

Systematics of the Genus Group Taxa of the Siphonini

71

appear to be universal among Actia species. It is hypothesized that these

characteristics of the male sternum 5 and pregonite are synapotypies of Actia (the

spinose pregonite proposed as an Actia synapotypy by Andersen 1983), and in

combination are better diagnostic features of the genus than the row of katepistemal

hairs discussed above and previously used to define the genus.

Another possible synapotypy of Actia species is lack of a dorsal cornu from the

cephalopharyngeal skeleton of first instars (Fig. 160). First instars of all nine

examined Actia species share this state, though the illustration of Actia dubitata by

Farinets (1980) seems to indicate that a distinct dorsal cornu is present in that

species (O’Hara in press “a”). Lack of a dorsal cornu is therefore either a

synapotypy of Actia or a derived subgroup of Actia.

Previous placement of Entomophaga exoleta in Actia is discussed in the

Phylogenetics section of Entomophaga Lioy.

Geographic distribution

Compared to 56 described species in the Old World, the Actia fauna of the New

World (four described species in the Nearctic region and one in the Neotropical

region) at first seems but a minor component of the world fauna. However, this

paucity of describ9d Actia species gives a false impression of the true diversity of

this genus in the New World. I have examined specimens of several new Nearctic

Actia species, and from relatively meagre collections of Neotropical Actia , have

recognized close to 20 undescribed species.

The Actia fauna of the Old World is much better known than its New World

counterpart, though undescribed species undoubtedly live in all regions, particularly

the Australian. Distributions are listed in Crosskey (1973, 1976a, 1980) for species

of Australia and the Oriental and Afrotropical regions, respectively. There are no

adequate keys to aid in the identification of the numerous species of these three

regions except for Dear and Crosskey’s (1982) key to Philippine species. Malloch’s

(1930b) key to the Oriental species is long outdated and Mesnil’s (1954) key to

Actia (as Entomophaga ) species of the upper Congo area only includes about half of

the described Afrotropical species.

Palearctic Actia are the best studied: most European species can be identified

using Mesnil’s keys (1963a, 1975: 1399), though species described later by Shima

(1970c) and Richter (1974, 1976a, 1980) from Japan and central Asia must be

identified from descriptions. Known distributions of several Palearctic Actia species

have been extended beyond Europe to Israel by Kugler (1979) and to central or

eastern Asia by Herting (1968b, 1973), Richter (1971, 1975, 1976a,b, 1980, 1981,

1986) and Richter and Khitsova (1982). The ranges of all 13 Palearctic species are

summarized in Herting (1984) and Norwegian Actia are listed in Rognes (1986).

Quaest. Ent., 1989, 25 (1,2)

72

O’Hara

List of described species included in Actia

A antiqua (Mesnil), 1954: 31 ( Entomophaga ). Holotype male, Zaire: Bambesa

(MRAC). Holotype examined.

N autumnalis (Townsend), 1917: 122 (Actiopsis). Holotype female, USA:

Maryland, Grove Hill (USNM). Holotype examined.

O brunnea Malloch, 1930b: 136. Holotype female, Malaysia: Malaya, Kedah

Peak (BMNH). Holotype examined.

A chrysocera Bezzi, 1923a: 96. Holotype male, Seychelles: Long Island

(BMNH). Holotype examined.

A ciligera (Mesnil), 1954: 29 ( Entomophaga ). Holotype female, Zaire: L. Kivu

(MRAC). Holotype examined.

O completa Malloch, 1930b: 139. Holotype male, Malaysia: Malaya, Selangor

(BMNH). Holotype examined.

P crassicornis (Meigen), 1824: 351 ( Tachina ). Holotype male, no locality data

(MNHN).

syn. bicolor (Macquart), 1834: 312 ( Thryptocera ). Type, France: Lille

(lost). — Herting, 1976: 3.

flavipalpis (Macquart), 1848: 135 ( Thryptocera ). Holotype female,

Switzerland: near Zurich (ETH). — Herting, 1976: 5.

nigripalpis (Robineau-Desvoidy), 1851: 182 ( Thryptocera ). Holotype

female (not male), France (MNHN). — Herting, 1974: 19.

palpalis (Rondani), 1859: 14 ( Thryptocera ). Lectotype female (by

designation of Herting, 1969a: 198), Italy: Parma (MZF). — Herting,

1969a: 198.

claripennis (Robineau-Desvoidy), 1863: 716 ( Thryptocera ).

Holotype female, France (MNHN). — Herting, 1974: 19.

scutellaris (Rondani), 1865: 195 ( Thryptocera ). Holotype male, Italy:

Parma (MZF). — Herting, 1969a: 200.

A cuthbertsoni Curran, 1933c: 162. Holotype male, Zimbabwe: Gatooma

(AMNH).

syn. cuthbersoni. Incorrect subsequent spelling of cuthbertsoni Curran

(Mesnil, 1977a: 83).

U darwini Malloch, 1929b: 334. Holotype male, Australia: Northern Territory,

Darwin (SPHTM). Holotype examined.

syn. brevis Malloch, 1930a: 309. Holotype male, Australia: New South

Wales, Sydney (SPHTM). Holotype examined. New synonymy.

O deferens Malloch, 1930b: 130. Holotype female (head lost), Malaysia: Malaya,

Kedah Peak (BMNH). Holotype examined.

N diffidens Curran, 1933a: 5. Holotype male, Canada: Nova Scotia, Kentville

(CNC). Holotype examined.

P dubitata Herting, 1971: 12. Holotype female, Switzerland: Delemont (SMNS;

Systematics of the Genus Group Taxa of the Siphonini

73

not located, possibly lost). Paratype examined.

U eucosmae Bezzi, 1926: 239. Holotype female, Australia: Queensland, Milton

Farm (published as “Brisbane”) (BMNH). Holotype examined.

A exsecta Villeneuve, 1936: 416. Two male syntypes, Uganda: Kampala (1 in

BMNH). Syntype examined.

A fallax (Mesnil), 1954: 29 ( Entomophaga ). Holotype female, Zaire: near Rweru

(MRAC). Holotype examined.

O fulvicauda Malloch, 1935: 680. Holotype male, Malaysia: Malaya, Selangor

(BMNH). Holotype examined.

A gratiosa (Mesnil), 1954: 34 ( Entomophaga ). Holotype male, Zaire: L. Kivu

(MRAC). Holotype examined.

A hargreavesi Curran, 1933c: 160. Holotype female (head missing), Uganda:

Kampala (BMNH). Holotype examined,

syn. comitata Villeneuve, 1936: 416. Four male and 2 female syntypes,

Uganda: Kampala (BMNH). — Crosskey, 1980: 852. Syntypes

examined.

P infantula (Zetterstedt), 1844: 1047 ( Tachina ). Six syntypes, Sweden: Sk&ne

(UZI).

syn. antennalis (Rondani), 1859: 16 ( Thryptocera ). Four syntypes, Italy:

Appennines (MZF). — Herting, 1969a: 190.

aristalis (Rondani), 1865: 194 ( Thryptocera ). Two syntypes, Italy:

Appennines (MZF). — Herting, 1969a: 190.

villeneuvii (Strobl in Czemy and Strobl), 1909: 221 ( Thryptocera

(. Actia )). Syntypes, Spain: Elche (NMBA).- — Mesnil, 1963a: 817.

villeneuvei. Incorrect subsequent spelling of villeneuvii Strobl

(Mesnil, 1963a: 817).

N interrupta Curran, 1933a: 6. Holotype male, USA: New York, Tuxedo

(AMNH). Holotype examined.

P jocularis Mesnil, 1957: 47. Holotype male, Japan: Tokura (CNC). Holotype

examined.

P lamia (Meigen), 1838: 254 ( Roeselia ). Neotype male (by designation of

O’Hara, 1985: 95), France: Saint-Sauveur (MNHN). Neotype examined,

syn. pilipennis Robineau-Desvoidy, 1830: 86 (junior homonym of A.

pilipennis (Fallen); replaced by A. lamia (Meigen) under Article 60

of the I.C.Z.N.). Lectotype male (by designation of O’Hara, 1985:

95), France: Saint-Sauveur (MNHN). Lectotype examined.

frontalis (Macquart), 1845: 289 ( Thryptocera ). Replacement name

for A. pilipennis Robineau-Desvoidy, 1830 (objective synonym).

obscurella Robineau-Desvoidy, 1851: 187. Syntypes, France

(MNHN).— Herting, 1974: 19.

vitripennis Rondani, 1859: 19. Replacement name for A. pilipennis

Robineau-Desvoidy, 1830 (objective synonym).

Quaest. Ent., 1989, 25 (1,2)

74

O’Hara

lamina. Incorrect subsequent spelling of lamia Meigen (Schiner,

1862:518).

U lata Malloch, 1930a: 307. Holotype male, Australia: New South Wales, Sydney

(SPHTM). Holotype examined.

A linguata Mesnil, 1968: 10. Holotype male. South Africa: Cape Town (BMNH).

Holotype examined.

A longilingua (Mesnil), 1954: 36 (. Entomophaga ). Holotype male, Zaire: Rutshuru

(MRAC). Holotype examined.

O magnicornis Malloch, 1930b: 133. Holotype male, Malaysia: Selangor

(BMNH). Holotype examined.

P maksymovi Mesnil, 1952b: 153. Holotype male, Switzerland: Engadine (not

located; a female paratype from same locality is mislabelled as holotype in

CNC). Paratype examined.

syn. maxymovi. Incorrect subsequent spelling of maksymovi Mesnil

(Richter, 1975: 644 and Richter, 1976a: 572).

O malaisei (Mesnil), 1953: 110 ( Crocuta ( Siphona )). Holotype male, Burma:

Kambaiti (ZMU). Holotype examined.

O mimetica Malloch, 1930b: 143. Holotype female, Malaysia: Malaya, Kedah

Peak (BMNH). Holotype examined.

P mongolica Richter, 1976a: 572. Holotype male, Mongolia: Eastern aimak

(ZIL). Holotype examined.

A munroi Curran, 1927a: 322. Holotype female, South Africa: Barberton (PPRI).

Holotype examined.

P nigra Shima, 1970c: 184. Holotype male, Japan: Hokkaido, Mt. Satsunai,

Pirikapetanu (BLKU). Paratype examined.

A nigrapex Mesnil, 1977a: 83. Holotype male, Madagascar: Amber Mtn.

(MNHN).

O nigriventris Malloch, 1935: 680 (as variety of Actia eucosmae Bezzi). Holotype

female (abdomen lost), Malaysia: Malaya, Selangor (BMNH). Holotype

examined.

P nigroscutellata Lundbeck, 1927: 462. Two male and 2 female syntypes,

Denmark: Tisvilde (ZMUC). Syntypes examined.

A nitidella Villeneuve, 1936: 417. Holotype female, Uganda: Kampala (BMNH).

Holotype examined.

S nitidiventris Curran, 1933a: 4. Holotype female, Panama: Barro Colorado Is.

(AMNH). Holotype examined.

P nudibasis Stein, 1924: 135. Four syntypes, German Democratic Republic:

Crimmitschau (ZMHU).

syn. ? resinellae (Schrank), 1781: 478 ( Musca ). Type(s), Austria (lost). —

Mesnil, 1963a: 825.

O oblimata Mesnil, 1957: 45. Holotype male, Burma: Kambaiti (ZMU). Holotype

examined.

Systematics of the Genus Group Taxa of the Siphonini

75

U painei Crosskey, 1962: 173. Holotype male. New Britain: Rabaul (BMNH).

Holotype examined.

A pallens Curran, 1927a: 322. Holotype female, South Africa: Natal, Durban

(PPRI). Holotype examined.

P pamirica Richter, 1974: 1268. Holotype male, USSR: Pamir, Debastia (ZIL).

Holotype examined.

U parviseta Malloch, 1930a: 308. Holotype male, Australia: New South Wales,

Sydney (SPHTM). Holotype examined.

O pellex (Mesnil), 1953: 111 ( Crocuta ( Siphona )). Holotype female, Burma:

Kambaiti (ZMU). Holotype examined.

O perdita Malloch, 1930c: 333. Holotype male (head lost), Malaysia: Malaya,

Selangor (BMNH). Holotype examined.

O philippinensis Malloch, 1930b: 134. Holotype female (head and abdomen lost),

Philippines: Luzon, Benguet (USNM). Holotype examined.

A picipalpis (Mesnil), 1954: 33 ( Entomophaga ). Holotype female (not male),

Zaire: Rutshuru (MRAC). Holotype examined.

P pilipennis (Fallen), 1810: 273 (Tachina). Lectotype male (by designation of

Crosskey, 1974: 302), Sweden (NRS).

syn. broteas (Walker), 1849: 763 {Tachina). Holotype male, England

(BMNH). — Crosskey, 1974: 277. Holotype examined.

flavisquamis (Robineau-Desvoidy), 1851: 181 {Thryptocera).

Syntypes, France (MNHN). — Herring, 1974: 19.

humeralis (Robineau-Desvoidy), 1851: 183 ( Thryptocera ). Holotype

female, France (lost). — Herring, 1984: 123.

exscensa (Walker), 1853: 66 {Tachina). Holotype female, England

(BMNH). — Crosskey, 1974: 283. Holotype examined.

nigrifrons (Robineau-Desvoidy), 1863: 714 {Thryptocera). Holotype

male, France (MNHN). — Herring, 1974: 19.

bigoti (Milliere), 1864: 385 {Morinia). Type(s), France (lost). —

Herring, 1984: 123.

reducta Villeneuve, 1920: 66 (as variety of A. pilipennis (Fallen),

though Villeneuve’ s description of A. pilipennis actually refers to A.

nudibasis (Herring, in litt.)). Type(s), no locality given (not located).

O pokharana Shima, 1970b: 275. Holotype male, Nepal: Pokhara (BPBM).

U pule x Baranov, 1938: 410. Lectotype male (by designation of Sabrosky and

Crosskey, 1969: 35), Solomon Islands: Tulagi (BMNH). Lectotype examined.

U quadriseta Malloch, 1936: 20. Holotype female, Australia: New South Wales,

Nyngan (SPHTM). Holotype examined.

A rejecta Bezzi in Bezzi and Lamb, 1926: 569. Holotype male (not female),

Rodriguez Is. (BMNH). Holotype examined.

A rubiginosa (Mesnil), 1954: 35 {Entomophaga). Holotype male, Zaire: Mokoto

(MRAC). Holotype examined.

Quaest. Ent., 1989, 25 (1,2)

76

O’Hara

N rufescens (Greene), 1934: 34 ( Actiopsis ). Holotype female, USA: South Dakota

(USNM). Holotype examined.

A russula Mesnil, 1977a: 84. Holotype male, Madagascar: Joffreville (MNHN).

O siphonosoma Malloch, 1930b: 136. Holotype male, Malaysia: Malaya, Selangor

(BMNH). Holotype examined.

O takanoi Baranov, 1935: 557. Lectotype female (by designation of Sabrosky and

Crosskey, 1969: 35), Philippines: Luzon, Los Banos (USNM). Lectotype

examined.

P tarsata Richter, 1980: 541. Holotype female, USSR: Chitinskaya Oblast,

Adriano vka (ZIL). Holotype examined.

A triseta (Mesnil), 1954: 32 ( Entomophaga ). Holotype male, Rwanda: near

Rweru (MR AC). Holotype examined.

A vulpina (Mesnil), 1954: 34 (, Entomophaga ). Holotype male, Zaire: Bambesa

(MRAC). Holotype examined.

O yasumatsui Shima, 1970b: 273. Holotype male, Hong Kong: Kowloon,

Taipokau (BPBM).

Nomen dubium

P rubrifrons (Robineau-Desvoidy), 1830: 87 ( Ceromya ). Type(s), France (lost). —

see Herting, 1974: 18.

Nomina nuda

N labellata Kamran, 1980: 52.

N pauciseta Kamran, 1980: 52.

List of examined, undescribed, species included in Actia

Actia nr. parviseta : One female from SE Queensland, Australia (DPI).

Actia Australia sp. 1 (=A. darwinil ): Males and females from Queensland (DPI).

Actia Australia sp. 2: One male from Mt. Lewis, Queensland (DPI).

Actia Australia sp. 3: One male, one female from Rockhampton, Queensland (DPI).

Actia Australia sp. 4: Two males, one female from Mt. Tamborine, Queensland (DPI).

Actia Australia sp. 5: Males and females from Queensland (CNC, DPI).

Actia Jamaica sp. 1: Males and females from St. Catherine (USNM).

Actia Liberia sp. 1: One male from Liberia (USNM).

Actia Mexico sp. 1 : One male from San Cristobal, Chiapas (CNC).

Actia Nepal sp. 1: Males from Nepal (CNC).

Actia Nepal sp. 2: One male from Kathmandu (CNC).

Actia New World spp. 1-19: These species are presently under revision by O’Hara and Shima.

Actia Uganda sp. 1 : One male from Ankole (USNM).

Systematics of the Genus Group Taxa of the Siphonini

77

Genus Peribaea Robineau-Desvoidy

Figs. 11-12, 27, 60-61,87-88, 120-121, 139, 153-156, 161.

Herbstia Robineau-Desvoidy, 1851: 184 (junior homonym, preoccupied by Herbstia Edwards, 1834).

Type-species,//, tibialis Robineau-Desvoidy, 1851 (monotypy).

Peribaea Robineau-Desvoidy, 1863: 720. Type-species, P. apicalis Robineau-Desvoidy, 1863 (by

designation of Coquillett, 1910: 587).

Strobliomyia Townsend,, 1926c:31. Type-species, Thryptocera fissicornis Strobl, 1910 (original

designation).

Eogymnophthalma Townsend, 1926a: 35. Type-species, E. orientalis Townsend, 1926 (original designation)

= Tachina orbata Wiedemann, 1830.

Talaractia Malloch, 1930a: 305 (as subgenus of Actia). Type-species, Actia ( Talaractia ) baldwini Malloch,

1930 (original designation).

Tarar actia. Incorrect subsequent spelling of Talaractia Malloch (Malloch, 1930a: 305).

Uschizactia Townsend, 1934: 248. Type-species, Actia uniseta Malloch, 1930 (original designation).

Recognition

Species of the genus Peribaea are widely distributed throughout the Old World,

and are easily recognized among siphonines by their possession of a strong,

downwardly directed, proepimeral seta (Fig. 27). Species of Chaetostigmoptera

Townsend and the Neaerini, and several other tachinids, also have a distinctly

developed, downwardly directed, proepimeral seta (varied from weak to strong),

though other differences indicate that this similarity is due to convergence.

Together, Peribaea and Siphona s.l. are distinguishable from all but a very few other

siphonines (and almost all other tachinids) by having the anal vein extended to the

wing margin.

Description

Length: 2.5-5.0mm.

Head (Figs. 11-12). — Five frontal setae, normal arrangement. Anterior proclinate orbital seta longer

than posterior one in most species. Eye of male and female subequal in size, medium to large, 0.73-0.86

head height; size in female subequal to that of male. Flagellomere 1 of male markedly varied in length,

0.39-0.75 head height; shape from linear or broad to bifid ( e.g . figs. 3a, 4a and 5a in Shima 1970a), trifid (P.

jepsoni and P. trifurcata; e.g. fig. 2a, op. cit.), or pectinate (P. baldwini, P. cervina and P. pectinata; Fig. 1 1

and fig. la, op. cit.). Flagellomere 1 of female simple, not bifid. Aristomere 1 short. Aristomere two 1.5-12X

longer than wide (2-4X in most species). Aristomere 3 long and evenly tapered in most species, short and

thickened to near tip in a few; almost bare to short plumose, in most species micropubescent. Clypeus

U-shaped in most species, only slightly broadened in a few. Palpus short, clavate. Proboscis with prementum

short to medium (not elongate), labella padlike.

Thorax (Fig. 27). — Prostemum setulose. Lower proepimeral seta strong and directed downward (Fig.

27). Katepistemum bare anterior to mid coxa. Lower katepistemal seta slightly shorter than or subequal in

length to upper anterior seta in most species, slightly longer in a very few. Four postsutural dorsocentral

setae (apparently never three). Upper part of anepistemum with single setula. Fore tibia with preapical ad

seta much shorter than d seta. Mid tibia with one ad seta. Tarsomeres normal in size, or tarsomere 5 of fore

leg elongate and broadened (only known in female of P. lobata)\ claws short in most species, medium in a

very few. Wing: CuAl with distal portion 0.63- 1.3X length of proximal portion (mean 0.84); anal vein

extended to wing margin. Wing setulae: /?, dorsally bare, or distally or entirely setulose, ventrally bare or

distally setulose; R2+3 bare except setulose dorsally and ventrally in P. modesta; R4+5 setulose from base to

beyond r-m; CuA} bare in almost all species (a few sparsely setulose).

Abdominal terga 1-5. — Abdomen ovoid in most species, markedly elongate in a few, and very elongate

in P. ugandana.

Quaest. Ent., 1989, 25 (1,2)

78

O’Hara

Tl+2 without median marginal setae; lateral marginal setae absent from most species, strong in a few. T3-T5

average in most species, with extra pair of lateral marginal setae on T3 in a few, weak lateral discal setae on

T4-T5 in P. discicornis.

Male genitalia (Figs. 60-61, 87-88, 120-121, 139). — S5 markedly varied, inner (or posterior) margins of

processes obtusely angled (Fig. 60) to almost transverse in most species, in a very few species V-shaped

(Fig. 61; resembling typical Actia shape except median cleft more distinct); apical lobe undifferentiated to

slightly differentiated; median lobe slightly to broadly rounded in most species, rather elongate and/or

flattened medially or posteromedially in a few; processes moderately setulose, with three to six pair of large

setae in most species. T6 varied from small pair of lateral sclerites to single, broad, dorsally continuous

sclerite. Ejaculatory apodeme with fan-shaped portion wider than hypandrial apodeme in most species,

subequal in width in a very few; some species with weak ring-like sclerotized sheath basally (e.g. P.

apicalis). Pregonite (Figs. 87-88) in profile approximately C-shaped, broad at mid length; bare. Epiphallus

present or absent, where present generally narrow. Distiphallus (Figs. 120-121) large, posterior margin

complete, variously incised laterally, distinctly broader than long in ventral view. Postgonite apically

rounded, pointed, or turned outward. Surstylus (Fig. 139) average length to long, broad in a few species,

straight to curved posteriorly; basally free from epandrium in most species (Fig. 139), fused with epandrium

in a few (e.g. P. trifurcata)', apically enlarged in some species, greatly swollen in a few, in one

(?undescribed) species from Australia subapically with small medially-directed lobe. Cerci short in most

species (Fig. 139), average length in a few, in profile straight along posterior margin or smoothly curved at

midlength; moderately setose on basal half.

Examined male genitalia of: P. apicalis, P. fissicornis, P. repanda, P. ?similata, P. tibialis, P. ugandana ,

Africa spp. 1-4, Australia spp. 1-3, Nepal sp. 1 and Sri Lanka sp. 1. Examined published figures of: P.

fissicornis (Andersen 1983, fig. 18), P. hirsuta, P. illugiana, P. pectinata, P. sedlaceki, P. trifurcata (all five

in Shima 1970a, figs. 1-5), P. insularis (Shima 1970c, figs. 1, 3a), P. alternata and P. orbata (both in Shima

1981, figs. 1 and 3).

Female genitalia (Figs. 153-156). — Varied from short and unmodified (Fig. 153) to extremely elongate

(Figs. 155-156). S6 bare or with very short hairs; in primitive forms S6 and S7 flat and wider than long, in

derived forms both sterna slightly to sharply keeled medially and pointed posteriorly in most species, in very

specialized forms both sterna elongate and tip of S7 extended over cerci, with intersegmental membrane

similarly elongate (Figs. 155-156); S7 without anterior apodeme. T6 distinctly developed and enclosing

spiracles of segment 6; dorsally continuous or discontinuous. T7 present as lateral sclerites in unmodified

forms, fused with S7 in some derived forms with S7 elongate (Figs. 154-156); enclosing spiracles of segment

7. 58 bare, absent (particularly in elongate forms) to average-sized, flat sclerite. 7j0 absent from some

species, present as a small to medium median sclerite in others.

Examined female genitalia of: P. fissicornis, P. tibialis, P. ugandana , Africa spp. 2-5 and Australia spp. 1,3.

Examined published figures of: P. alternata, P. orbata (both in Shima 1981, fig. 2) and P. tibialis (Andersen

1983, fig. 4).

Taxonomic changes

Identity of Tachina orbata Wiedemann. — The type of Tachina orbata

Wiedemann (1830: 336) is lost. Crosskey (1967) designated a neotype for this

species, believing T. orbata to be a widespread Old World species of Peribaea.

According to Mesnil (in litt.), the original description of T. orbata refers to a

neaerine. If one were to accept this opinion and invalidate Crosskey’s neotype (on

the basis of an incorrect neotype designation), then P. aegyptia would become the

valid name for this Peribaea species. However, in the interests of nomenclatural

stability, Crosskey and Shima plan to submit a proposal to the International

Commission on Zoological Nomenclature requesting that Crosskey’s (1967)

neotype be retained for the name Tachina orbata Wiedemann (Crosskey in litt.).

Current usage of orbata as a valid species of Peribaea is followed herein.

Systematics of the Genus Group Taxa of the Siphonini

79

Status ofPeribaea subaequalis (Malloch). — Malloch described P. monticola, P.

rotundipennis and P. subaequalis from specimens collected in the Cuemos Mts. on

Negros Island in the Philippines. The first two names were synonymized with P.

orbata by Crosskey (1966: 107), while the third has continued to be cited as valid

(Crosskey, 1976: 214). I examined all three of Malloch’s types along with

specimens of P. orbata , and found no substantial difference among them.

Unfortunately, Malloch’s type specimens are not ideal for comparison because all

three lack heads, and two are females while the third is a male. Nevertheless, the

fact that Malloch’s types were collected from the same locality (dates unknown) and

are similar to one another and to specimens of P. orbata in such important

characteristics as wing vein setulation and abdominal coloration strongly suggests

that they are all conspecific. (Even Malloch stated that his specimens “may

ultimately prove to be mere variations within a single species” (1930b: 142).) The

one difference, anal vein not extended to wing margin in the type of P. subaequalis,

is very rare in Peribaea and not typical of any known species, so probably

represents an aberration in this specimen (even the paratypes of P. subaequalis have

the anal vein reaching wing margin). For all these reasons I feel confident in adding

P. subaequalis to the list of synonyms of the widespread species P. orbata.

Hosts

Hosts of Peribaea species belong to several families of Microlepidoptera and

Macrolepidoptera (Table 2). The most commonly parasitized group is the

Noctuoidea, accounting for half the total records.

Adult females of Peribaea species are thought to larviposit directly on their hosts

(Herting 1957). Presumably the elongate ovipositor of some Peribaea species is an

adaptation to this behavior.

Phylogenetics

The monophyly of Peribaea is well established (Andersen 1983: 10). Adults

possess externally a strong, downwardly directed, proepimeral seta (Fig. 27). This

state is unique to Peribaea among siphonines, though is convergently, and often

more weakly, developed in species of Chaetostigmoptera Townsend and the

Neaerini, and several other tachinids. The distiphallus is varied in shape among

Peribaea species, though its sclerotized posterior surface is a shared characteristic

and is hypothesized as a synapotypy; the partially unsclerotized posterior surface in

all but a very few other (unrelated) siphonines (see Table 1) is interpreted as

plesiotypic. Sternum 8 in the female genitalia is bare in all examined Peribaea

species, though several derived species lack this sclerite entirely (Figs. 154-156). A

bare, unsclerotized sternum 8 is considered synapotypic of the genus, and absence of

sternum 8 is interpreted as a more derived condition (sternum 8 convergently lost in

a few other siphonines; see Table 1).

Quaest. Ent., 1989, 25 (1,2)

80

O’Hara

First instars of Peribaea species are characterized by an apically narrowed or

hook-like labrum (Fig. 161; also see O’Hara in press “a”). A hatchet-like labium

characterizes more primitive siphonines (with the known exception of Ceromya

silacea, Fig. 159), so the shape of the labrum in Peribaea species is interpreted as

derived.

Geographic distribution

Forty-five described species are recognized in Peribaea , all are restricted to the

Old World. Greatest diversity, accounting for almost half the described species, is

recorded from the Afrotropical region (particularly the upper Congo area, as a result

of Mesnil’s work on the siphonines of that area; distributions in Crosskey 1980).

More modest diversity is found in the three other Old World regions. Oriental

species are listed in Crosskey (1976a; Philippine species keyed in Dear and

Crosskey 1982) and Australian species in Crosskey (1973) and Shima (1970a, with

key to Papuan species; 1981). Most Palearctic Peribaea species are redescribed and

keyed in Mesnil (1963a; also key to separate P. apicalis and P. tibialis in Herting

1968a). Other records for Palearctic species are given in Mesnil and

Pschom-Walcher (1968), Herting (1968b, 1969b), Crosskey (1976b), Kugler (1979),

Karczewski (1983), Mihalyi and Weinberg (1984), Richter (1971, 1975, 1976a,

1980, 1981, 1986) and Rognes (1986), and summarized in Herting (1984).

I suspect that the preponderance of described species in the Afrotropical region

accurately reflects the true pattern of Peribaea diversity. Presence of ten species in

the Australian region (including one on the Solomon Islands), seven species on

Madagascar (Mesnil 1977a) and one each on Mauritius and Seychelles Islands,

indicates that the dispersal power of some members of this genus are as great as in

other siphonine genera that are distributed worldwide. It is therefore puzzling that

Peribaea has not reached the New World. Hosts do not seem to be the limiting

factor, as many host genera are Holarctic or cosmopolitan in distribution.

List of described species included in Peribaea

U alternata Shima, 1981: 445. Holotype male, New Guinea: Wau, Mt. Kaindi

(BPBM). Paratype examined.

A annulata (Mesnil), 1954: 21 ( Strobliomyia ). Holotype male, Zaire: Rwankwi

(MRAC). Holotype examined.

A anthracina Mesnil, 1977a: 81. Holotype male, Madagascar: Belazao (MNHN).

P apicalis Robineau-Desvoidy, 1863: 721. Holotype male (head and abdomen

lost), France: Saint Sauveur (MNHN).

U argentifrons (Malloch), 1930a: 309 ( Actia ). Holotype male, Australia: New

South Wales, Sydney (SPHTM). Holotype examined,

syn. angustifrons. Incorrect subsequent spelling of argentifrons Malloch

(Hardy, 1959: 213).

Systematics of the Genus Group Taxa of the Siphonini

81

U baldwini (Malloch), 1930a: 306 ( Actia { Talaractia )). Holotype male, Australia:

Queensland, Palm Is. (SPHTM). Holotype examined.

A cervina (Mesnil), 1954: 18 ( Strobliomyia ). Holotype male, Zaire: Rutshuru

(IRSN). Holotype examined.

A clara (Mesnil), 1954: 21 ( Strobliomyia ). Holotype male, Zaire: Katanga, Kalabi

(MRAC). Holotype examined.

A compacta (Curran), 1927a: 324 {Actia). Holotype male (head lost). South

Africa: East London (PPRI). Holotype examined.

P discicornis (Pandelle), 1894: 109 {Thryptocera). Holotype male, France:

Pyrenees, Tarbes (IRSN).

A ferina (Mesnil), 1954: 17 {Strobliomyia). Holotype male, Rwanda: Kibga

(MRAC). Holotype examined.

P fissicornis (Strobl), 1910: 139 {Thryptocera). Holotype male (not female),

Austria: “Styria” (NMBA).

A gibbicornis (Mesnil), 1954: 19 {Strobliomyia). Holotype male, Zaire: Rutshuru

(IRSN). Holotype examined.

U hirsuta (Shima), 1970a: 269 {Strobliomyia). Holotype male, New Guinea:

Popondetta (BPBM).

O hyalinata (Malloch), 1930b: 138 {Actia). Holotype female (not male), Malaysia:

Malaya, Selangor (BMNH). Holotype examined.

U illugiana (Shima), 1970a: 265 {Strobliomyia). Holotype male, New Britain:

Gazelle Peninsula, Illugi (BPBM).

O insularia (Shima), 1970c: 179 {Strobliomyia). Holotype male, Japan: Ryukyu

Islands, Amami Is., Tokunoshima (BLKU).

A jepsoni (Villeneuve), 1937: 2 {Strobliomyia). Holotype male, Mauritius (CNC).

Holotype examined.

P leucopheae (Mesnil), 1963b: 33 {Strobliomyia). Holotype female, USSR:

Tadjikistan, Varzoba (ZIL). Holotype examined.

A lobata Mesnil, 1977a: 80. Holotype male, Madagascar: Manjakatatompo

(MNHN).

A longiseta (Villeneuve), 1936: 417 {Actia). Holotype female, Uganda: Kampala

(BMNH). Holotype examined.

O malayana (Malloch), 1935: 678 {Actia). Holotype male (abdomen lost),

Malaysia: Malaya, Selangor (BMNH). Holotype examined.

A mitis (Curran), 1927a: 323 {Actia). Syntypes, South Africa: Barberton (PPRI).

Syntypes examined.

A modesta (Mesnil), 1954: 14 {Strobliomyia). Holotype male, Zaire: Rutshuru

(MRAC). Holotype examined.

0,P,A,U orbata (Wiedemann), 1830: 336 {Tachina). Neotype female (by

designation of Crosskey, 1967: 106), India: Assam, Azra (BMNH). Lectotype

examined.

syn. aegyptia (Villeneuve), 1912: 508 {Gymnopareia {Actia)). Lectotype

Quaest. Ent., 1989, 25 (1,2)

82

O’Hara

male (by designation of Crosskey, 1966: 108), Egypt: Qaliub

(BMNH). — Crosskey, 1976a: 214. Lectotype examined.

orientalis (Townsend), 1926a: 35 ( Eogymnophthalma ). Lectotype

male (by fixation of Townsend, 1940: 213), Indonesia: Sumatra, Fort

de Kock (ZMA). — Crosskey, 1966: 107. Lectotype examined.

nigripes (Curran), 1927c: 6 ( Actia ). Holotype male, Zaire: Boma

(AMNH). — Crosskey, 1980: 853. Holotype examined.

nigritula (Malloch), 1930a: 309 {Actia). Holotype female, Australia:

Queensland, Cairns (SPHTM). — Crosskey, 1966: 107. Holotype

examined.

subaequalis (Malloch), 1930b: 142 {Actia). Holotype male (head

lost), Philippines: Negros, Cuemos Mtns. (USNM). Holotype

examined. New synonymy.

monticola (Malloch), 1930b: 143 {Actia). Holotype male (head lost),

Philippines: Negros, Cuemos Mtns. (USNM). — Crosskey, 1966:

107. Holotype examined.

rotundipennis (Malloch), 1930b: 143 {Actia). Holotype female (head

lost), Philippines: Negros, Cuemos Mtns. (USNM). — Crosskey,

1966: 107. Holotype examined.

sororcula (Mesnil), 1954: 16 {Strobliomyia). Holotype female, Zaire:

Rutshuru (MRAC). — Crosskey, 1976a: 214. Holotype examined.

P palaestina (Villeneuve), 1934: 57 {Actia). Holotype female, Israel: Rehoboth

(SMNS). Holotype examined.

syn. alipes (Villeneuve), 1942b: 134 {Actia). Holotype female, Egypt:

Assuan (CNC). — Herting, 1982: 8. Holotype examined.

U pectinata (Shima), 1970a: 261 {Strobliomyia). Holotype male, New Britain:

Gazelle Penn., Illugi (BPBM).

U plebeia (Malloch), 1930a: 310 {Actia). Holotype male, Australia: New South

Wales, Coramba (SPHTM). Holotype examined,

syn. plebia. Incorrect subsequent spelling of plebeia Malloch (Hardy,

1959: 213).

A pulla Mesnil, 1977a: 82. Holotype male, Madagascar: Ambato-Boeni (MNHN).

A repanda (Mesnil), 1954: 16 {Strobliomyia). Holotype male, Zaire: nr. Rwindi,

Ndeko (MRAC). Holotype examined.

A rubea Mesnil, 1977a: 82. Holotype female, Madagascar: Amber Mtn. (MNHN).

U sedlaceki (Shima), 1970a: 267 {Strobliomyia). Holotype male, New Guinea:

Popondetta (BPBM).

O setinervis (Thomson), 1869: 519 {Thryptocera). Holotype female, China

(NRS).

0,P similata (Malloch), 1930b: 137 {Actia). Holotype male, Malaysia: Malaya,

Selangor (BMNH). Holotype examined.

A spoliata (Bezzi), 1923a: 95 {Actia). One male and 1 female syntype, Seychelles

Systematics of the Genus Group Taxa of the Siphonini

83

Islands (BMNH). Syntypes examined.

U stiglinae (Bezzi), 1928: 204 ( Actia ). Holotype male (not female), Fiji: Lautoka

(BMNH). Holotype examined. New combination, moved from Actia.

O, A suspecta (Malloch), 1924: 409 (Actia). Holotype male (not female), India:

Bihar, Pusa (BMNH). Holotype examined,

syn. nana (Curran), 1928: 237 (Actia). Holotype female, Uganda:

Kampala (BMNH). — Crosskey, 1976a: 214. Holotype examined.

P, A tibialis (Robineau-Desvoidy), 1851: 185 (Herbstia). Holotype male, France:

Saint Sauveur (lost).

syn. flavicornis Robineau-Desvoidy, 1863: 721. Holotype female, France:

Lozere (MNHN). — Herting, 1974: 19.

minuta Robineau-Desvoidy, 1863: 722. Holotype female, France

(MNHN).— Herting, 1974: 19.

A timida (Mesnil), 1954: 18 (Strobliomyia). Holotype male, Zaire: Rutshuru

(MRAC). Holotype examined.

U trifurcata (Shima), 1970a: 263 (Strobliomyia). Holotype male, New Guinea:

Popondetta (BPBM).

A ugandana (Curran), 1933c: 161 (Actia). Holotype male, Uganda: Kampala

(BMNH). Holotype examined.

O uniseta (Malloch), 1930b: 129 (Actia). Holotype male (head lost), Malaysia:

Malaya, Selangor (BMNH). Holotype examined.

P ussuriensis (Mesnil), 1963a: 807 (as subspecies of Strobliomyia hyalinata

(Malloch)). Holotype male, USSR: Ussuri, Sucan (ZIL). Holotype examined.

A vidua (Mesnil), 1954: 15 (Strobliomyia). Holotype male, Zaire: Rutshuru

(MRAC). Holotype examined.

List of examined, undescribed, species included in Peribaea

Peribaea Africa sp. 1: Two males from Ngong, Kenya (USNM).

Peribaea Africa sp. 2: One male and one female from Ankole, Uganda (USNM).

Peribaea Africa sp. 3: One male and one female from Archer’s Post, Kenya (CAS).

Peribaea Africa sp. 4: Males and females from Natal, South Africa (USNM).

Peribaea Africa sp. 5: One female from Kruger Nat. Pk., South Africa (USNM).

Peribaea Australia sp. 1 : Males and females from Queensland (DPI).

Peribaea Australia sp. 2: One male and one female from Kairi, Queensland (DPI).

Peribaea Australia sp. 3: Males and females from Queensland (DPI).

Peribaea Nepal sp. 1 : Three males from Lothar (CNC).

Peribaea Sri Lanka sp. 1 : One male from Colombo (CAS).

Quaest. Ent., 1989, 25 (1,2)

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O’Hara

Genus Siphona Meigen sensu lato

Figs. 13-20, 22, 24-26, 28, 30, 32, 34, 36, 38, 43-46, 62-70, 89-106, 122-136,

140-144, 162-163.

Recognition

Siphona Meigen, as considered here in the broad sense, is a diverse,

monophyletic group of cosmopolitan distribution. Adults are morphologically

varied, but all are characterized by the anal vein extended to the wing margin at least

as a sharply creased fold (Fig. 22). This state is also shared by Perihaea species, but

these are easily recognized by their two strong and opposed proepimeral setae (one

strong proepimeral seta in Siphona s.l. species and other siphonines, cf. Figs. 27 and

28).

Three known Old World Actia species are easily confused with Siphona s.l.

species because they have the anal vein extended to the wing margin. They differ

from members of Siphona s.l. in their possession of a partial to complete row of

katepistemal hairs and distinctive Actia- type male genitalia (see Recognition section

of Actia).

Though not very useful for general identification purposes, two features of first

instars are diagnostic (and synapotypic) of Siphona s.l. species; both on the ventral

surface of the abdomen on segments 6 and 7 (see O’Hara in press “a”). Segment 6 in

Siphona s.l. species is equipped posteriorly with from two, to a row of about 10,

large spinules or hooks (Figs. 162-163). Other siphonines are bare in this region or

have tiny spinules (Figs. 158-161) with the exception of Ceromya Australia sp. 3. S.

( Aphantorhaphopsis ) Uganda sp. 1 is the only known Siphona s.l. species without

the typical condition of the genus. The spinulose condition on the ventral surface of

segment 7 posteriorly also distinguishes almost all Siphona s.l. species from other

siphonines. With the exception of a few species (see Table 1), Siphona s.l. species

(Figs. 163) have a single dominant row of spinules in this position and other

siphonines (Figs. 158-161) have two or more even or uneven rows of spinules (S.

( Pseudosiphona ) species have the latter condition, as exemplified here by S. (P.) sp.

14, Fig. 162).

Key to adults of the subgenera of Siphona sensu lato

(See section entitled “Review of major keys to genera and subgenera of the

Siphonini” for information about how other authors have keyed (and classified) the

following taxa.)

1. Mid tibia lacking ad seta (Fig. 38); aristomere 1 at least 2X

longer than wide, subequal in length to aristomere 2 (Fig. 15);

2-3mm long (five species; western North America)

S. (Baeomyia O’Hara), p. 97

T. Mid tibia with one or two ad seta on lower half (Fig. 37);

Systematics of the Genus Group Taxa of the Siphonini

85

aristomere 1 shorter than aristomere 2, in most species shorter

than wide (Figs. 13-14, 16-20); average-sized specimens of all

but a very few species longer than 3mm 2

2. (T) Maxillary palpus cylindrical to tip, markedly reduced to

average length (Fig. 16) 3

2'. Maxillary palpus clavate apically, short to long (Figs. 1-15, 17-20) 4

3. (2) Distiphallus laterally incised and posterolateral arm clearly

developed (Fig. 127); pregonite bare in Mauritius sp. 1, with

long seta posteriorly in other species (Fig. 94) (Afrotropical,

Nearctic, Palearctic and Oriental regions)

S. (Ceranthia R.-D.), p. 99

3'. Distiphallus not laterally incised and without posterolateral

arm; pregonite bare (two Australian species, Australia sp. 1 and

New Guinea sp. 1)

two undescribed species of S. (Aphantorhaphopsis Tnsd.), p. 92

4. (2') Proboscis with prementum and labella elongate, labella slightly

longer than prementum and in most species longer than eye

height (Figs. 25-26); pregonite of male genitalia without seta

posteriorly and lacking spinules apically (Fig. 96); female

genitalia unmodified (Figs. 43, 45) except in S. melanura (over

80 described species; cosmopolitan) ....S, (Siphona Mg.), p. 108

4'. Proboscis with prementum and labella varied in length, labella

in most species padlike to slightly lengthened (Figs. 13-15,

17-19, 23-24), in a very few species as long as prementum (Fig.

20); male genitalia varied, in almost all species in which labella

are as long as prementum the pregonite has a seta posteriorly

and/or tiny spinules apically (Figs. 89-95, 97-98, 100,

102-106); female genitalia varied 5

5. (4') Old World in distribution (a polyphyletic group of 21 described

and many new species)

S. ( Aphantorhaphopsis Townsend), p. 92

5'. New World in distribution 6

6. (5') Distiphallus in profile with parallel anterior and posterior

margins, with or without spines apically (Figs. 122-123);

pregonite long and slender with tiny spinules apically (Fig. 89);

cerci broadened in posterior view (Fig. 142); head as in Fig. 13,

with labella varied from padlike to moderately lengthened;

body mostly yellow with sparse abdominal pruinosity; wing

vein R4+5 setulose from base to beyond crossvein r-m ( S .

singularis complex and one new species; Neotropical)

S. ( Actinocrocuta Townsend), p. 87

6'. Male genitalia varied, with not more than one of above states;

Quae st. Ent., 1989, 25 (1,2)

86

O’Hara

externally varied, but a very few species with above states 7

7. (6') Distiphallus in profile narrow on apical half and with or

without enlarged spines apically or anteriorly (Fig. 130);

pregonite without spinules anterolaterally (Figs. 97-98); male

sternum 5 with apex of apical lobe curved inward (similar to

Figs. 62-63); proboscis with labella elongate, 0.7-0.9

prementum length (Figs. 18, 24); wing vein R4+5 not setulose

beyond crossvein r-m (three described and at least 13 new

species; western United States and Neotropics)

S. (Siphonopsis Townsend), p. 120

7'. Distiphallus in profile broader on apical half than in Fig. 130

and with or without enlarged spines apically and/or anteriorly;

pregonite with or without tiny spinules anterolaterally; male

sternum 5 with or without apex of apical lobe curved inward;

externally varied, but a few species with above states 8

8. (7') Pregonite (Fig. 95) with tiny spinules anterolaterally, broad

with two or more tiny setae posteriorly in most species, a few

species with J-shaped pregonite or only one tiny seta

posteriorly; distiphallus triangular in profile in most species

(Fig. 128); cerci sharply inflexed at midlength in most species

(Fig. 140); head habitus varied, but most species with row of

short parafacial hairs extended to lower half of eye, arista

haired and proboscis with labella slightly more than half

prementum length (Fig. 17) (one described and at least 18 new

species; United States to northern Argentina)

S. (Pseudosiphona Townsend), p. 103

8'. Pregonite more slender than in Fig. 95, with not more than one

seta posteriorly and with or without spinules anterolaterally;

distiphallus varied, but not triangular as in Fig. 128; cerci

smoothly curved to sharply inflexed; externally varied, but a

very few species with above states 9

9. (8') Male genitalia with median lobe of sternum 5 flattened (Fig.

68), pregonite bare (no spinules or posterior seta; Fig. 99),

distiphallus smoothly tapered in profile (Fig. 131), and cerci

sharply inflexed at midlength; body relatively large, 4.0-5. 0mm

in length; also see Recognition section (one described and one

new species; Ecuador and Peru)

S. (Uruactia Townsend), p. 123

9'. Male genitalia varied, but with not more than two of the four

states above 10

10. (9') Male genitalia with pregonite lacking spinules anterolaterally

(Fig. 90; posterior seta present or absent), distiphallus tapered

Systematics of the Genus Group Taxa of the Siphonini

87

in profile (Fig. 124), and cerci not sharply inflexed at midlength

and of average shape (similar to Fig. 144) to slightly broadened

(Fig. 143) in posterior view; body relatively small, 2.0-3. 5mm

in length and dark-colored; also see Recognition section (two

described and at least four new species; SW United States to SE

Brazil) S. (Aphantorhapha Townsend), p. 89

10'. Male genitalia without above combination of states; externally

varied (two described and over 40 new species; New World,

mostly Neotropical)

...New World Siphona s.l. species unplaced to subgenus, p. 125

[See descriptive sections on three species groups and unplaced

species of New World Siphona 5./.]

Siphona (subgenus Actinocrocuta Townsend)

Figs. 13,62, 89, 122-123, 142.

Actinocrocuta Townsend, 1935: 228. Type-species, A. chaetosa Townsend, 1935 (original designation) =

Tachina singularis Wiedemann, 1830. New subgeneric status in Siphona Meigen.

Recognition

Adults of this Neotropical group of few species cannot be distinguished from all

other Siphona s.l. species except by features of the male genitalia, though the

following external characteristics are shared by the known species: yellow overall

except for varied amounts of reddish brown at femoral-tibial articulations,

surrounding marginal setae on abdomen, and along abdominal midline; abdomen

very sparsely pruinose; lower katepistemal seta at least length of upper anterior seta;

three postsutural dorsocentral setae; wing vein R4+5 setulose from base to beyond

crossvein r-m\ and anal vein extended to wing margin. The following combination

of male genitalic states is unique to S. (Actinocrocuta) species: pregonite slender,

apically rounded and covered with tiny spinules (Fig. 89; more curved than in Fig.

100); distiphallus in profile slender, nearly parallel-sided, varied apically (Figs.

122-123); and cerci in posterior view broadened to near tip (Fig. 142; more

broadened than in some S. ( Aphantorhapha ) species [Fig. 143], though similar in

shape in a very few other Siphona s.l. species).

Description

Length: 3.5-4.5mm.

Head (Fig. 13). — Anterior proclinate orbital seta longer than posterior one. Eye of male and female

subequal in size, medium to medium-large, 0.76-0.81 head height. Flagellomere 1 of male and female

subequal, medium-short to medium length, 0.41-0.52 head height; average width. Aristomere 1 short.

Aristomere 2 relatively short, about 1.5X longer than wide. Aristomere 3 long and evenly tapered, pubescent

to short plumose. Clypeus narrow and enclosed in membrane to slightly broadened. Palpus short, clavate.

Proboscis with prementum medium in length (slightly over half head height in length), labella padlike to

moderately lengthened.

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O’Hara

Thorax. — Prostemum setulose. Lower katepistemal seta longer than upper anterior seta. Three

postsutural dorsocentral setae. Upper part of anepistemum with single setula. Fore tibia with preapical ad

seta much shorter than d seta. Mid tibia with one ad seta. Claws short. Wing vein CuA] with distal portion

0.35-0.49 length of proximal portion (mean 0.42). Wing setulae: Sc ventrally setulose in some specimens; R]

dorsally distally or entirely setulose, ventrally bare; /?4+5 setulose from base to beyond r-m\ CuA{ bare or

setulose.

Abdominal terga 1-5. — Abdomen ovoid in shape. 7j+2 without median marginal setae, lateral marginal

setae absent to weak. TyT5 with average setation.

Male genitalia (Figs. 62, 89, 122-123, 142). — S5 (Fig. 62) with posterior margins of processes

approximately U-shaped; apical lobe distinctly differentiated, apex curved inward in some specimens of 5.

singularis complex; median lobe pointed to narrowly rounded, relatively unmodified; processes moderately

setulose. T6 apparently absent. Ejaculatory apodeme with fan-shaped portion subequal to 2.0X wider than

width of hypandrial apodeme. Pregonite (Fig. 89) in profile rather slender and elongate; outer surface short

spinose apically, tiny seta posteriorly in 5. singularis complex, seta absent from single specimen of S. (A.)

sp. 1. Epiphallus absent. Distiphallus reduced posteriorly, not incised laterally, in profile rather narrow

basally, nearly parallel-sided to tip, in S. singularis complex with or without (Fig. 122) apical hook

anteroventrally, in S. (A.) sp. 1 with several hooks radiated from apex ventrally and anteroventrally (Fig.

123). Postgonite rounded apically in S. singularis complex, short in S. (A.) sp. 1. Surstylus rather short to

average length, more or less straight; basally free from epandrium. Cerci average length, in profile rather

straight at midlength, in posterior view broadened to near apex (Fig. 142); moderately setose on basal half in

S. (A.) sp. 1, varied in members of 5. singularis complex from state in S. (A.) sp. 1 to densely setose along

length.

Examined male genitalia of: S. singularis complex, S. (A.) sp. 1.

Female genitalia — Moderately extensible. S6 bare to sparsely haired posteromedially; slightly keeled

posteromedially. T6 absent; spiracles of segment 6 in membrane dorsal to lateral margins of S6. S7 with long

anterior apodeme; slightly keeled posteromedially. T7 absent; (spiracles not located). S8 much broader than

average, haired. 7j0 present as two sclerites.

Examined female genitalia of: S. singularis.

Taxonomic changes

Synonymy of S. (Actinocrocuta) chaetosa (Townsend) with S. (Actinocrocuta)

singularis (Wiedemann). — The primary types (both male) of S. chaetosa (type

locality “Trinidad”) and S. singularis (type locality “Brasilia”) are externally very

similar, with no significant differences. The male genitalia are less similar, differing

particularly in features of the distiphallus and degree of setation on posterior surface

of cerci. In the absence of additional material I would be inclined towards retaining

both names, but additional material has been examined and includes not only

specimens with clearly intermediate states but also specimens with more extreme

states. I have not been able to sort these specimens into distinct morphological

groups because the apparent differences seem continuous and are not clinally

distributed. There is either one very morphologically varied species, or two or more

sibling species. Because of the range of observed variation and presence of

intermediates, I synonymize the name S. chaetosa with S. singularis , and refer to

this varied taxon as the S. singularis complex.

Hosts. Unknown.

Systematics of the Genus Group Taxa of the Siphonini

89

Phylogenetics

Siphona (Actinocrocuta) here comprises the S. singularis complex and S.

(Actinocrocuta) sp. 1. Adults of these taxa are similar in external appearance and

coloration (see Recognition section), but these characteristics alone are not good

indicators of monophyly because convergence in such features is common among

Siphona s.l. species. Monophyly of this subgenus is better supported by three

characters of the male genitalia: broadened cerci (Fig. 142), slender distiphallus

(Figs. 122-123) and slender and apically spined pregonite (Fig. 89). These states are

individually present in a few other Siphona s.l. species, in species unrelated to one

another. The states are therefore homoplastic, though the presence of all three in S.

( Actinocrocuta ) species is better explained by monophyly of the subgenus than by

independent development of these states in the two included species. The sister

group to S. ( Actinocrocuta ) cannot be established with confidence (it is certainly to

be sought among other Neotropical Siphona s.l. taxa), so S. ( Actinocrocuta ) is

retained as a narrowly defined subgenus of Siphona s.l.

Geographic distribution

The S. singularis complex is widely distributed throughout the Neotropics, with

records from Costa Rica, Colombia, Ecuador, Peru, Brazil (Manaus and

southeastern region) and Trinidad. A single male specimen of a new species,

referred to above as S. (A.) sp. 1, was collected at 400m from Avispas, Madre de

Dios, Peru (1-15.X.1962), along with several specimens of the S. singularis

complex.

List of described species included in Siphona {Actinocrocuta)

S singularis (Wiedemann), 1830: 335 ( Tachina ). Holotype male, Brazil (FSF).

Holotype examined.

syn. chaetosa (Townsend), 1935: 228 {Actinocrocuta). Lectotype male

(by fixation as “holotype” by Townsend, 1940: 275 [see explanation

under “Type designations of Coquillett and Townsend”]), Trinidad

(USNM). Lectotype examined. New synonymy.

List of examined, undescribed, species included in Siphona {Actinocrocuta)

S. ( Actinocrocuta ) sp. 1 : One male from Peru (CNC).

Siphona (subgenus Aphantorhapha Townsend)

Figs. 14, 63, 90, 124, 143.

Aphantorhapha Townsend, 1919: 586. Type-species, A. arizonica Townsend, 1919 (original designation).

New subgeneric status in Siphona Meigen.

Quaest. Ent., 1989, 25 (1,2)

90

O’Hara

Recognition

As presently conceptualized, S. ( Aphantorhapha ) comprises two described and

four undescribed New World species, and is one of the more difficult Siphona

subgenera to diagnose. Adults are rather small (2.0-3. 5mm in length) and

dark-colored, with more or less average-sized eye and flagellomere 1, three

postsutural dorsocentral setae, lower katepistemal seta subequal to or longer than

upper anterior seta, crossvein dm-cu slightly removed from wing margin, and anal

vein extended to wing margin. As these external characteristics are shared by some

other species of Siphona s.L, the male genitalia of specimens must be examined for

a reliable identification.

One of the two distinctive features of the male genitalia of S. ( Aphantorhapha )

species is the shape of the distiphallus (Fig. 124), which in profile has a

characteristically tapered appearance and rounded to pointed tip. The other

distinctive feature, though not present in all species (absent from single examined

male of S. atoma and some specimens of S. arizonica), is the slightly broadened

cerci (Fig. 143; broadened at midlength rather than to near tip like in S.

( Actinocrocuta ) species, Fig. 142). Like many other Siphona s.L species, the

pregonite (Fig. 90) is of average shape, apically bare, and with or without a seta

posteriorly.

Description

Length: 2.0-3. 5mm.

Head (Fig. 14). — Proclinate orbital setae subequal in length or anterior one longer. Eye of male medium

to medium-large, 0.71-0.83 head height; eye of female slightly smaller to slightly larger than in male.

Flagellomere 1 of male medium-short to medium length, 0.43-0.54 head height; shape from linear to

broadened. Flagellomere 1 of female shorter than in male. Aristomere 1 short. Aristomere two 2.0-4.0X

longer than wide. Aristomere 3 long and evenly tapered to short and thickened to near tip, micropubescent to

short plumose. Clypeus narrow and enclosed in membrane to slightly broadened. Palpus short, clavate.

Proboscis with prementum short to medium in length (about half head height in length), labella padlike in

most species, slightly lengthened in S. (A.) sp. 2.

Thorax. — Prostemum setulose. Lower katepistemal seta subequal in length or longer than upper

anterior seta. Three postsutural dorsocentral setae. Upper part of anepistemum with single setula. Fore tibia

with preapical ad seta much shorter than d seta. Mid tibia with one ad seta. Claws short. Wing vein CwA,

with distal portion 0.44-0.85 length of proximal portion (mean 0.64). Wing setulae: /?, dorsally bare or

entirely setulose, ventrally bare or distally setulose; /?4+5 setulose to or beyond r-m; CwA, bare.

Abdominal terga 1-5. — Abdomen ovoid in shape. 7j+2 without median marginal setae, lateral marginal

setae absent to weak. T3-T5 with average setation.

Male genitalia (Figs. 63, 90, 124, 143). — S5 (Fig. 63) with posterior margins of processes slightly

obtuse to approximately U-shaped; apical lobe distinctly differentiated, apex curved inward in some species;

median lobe rounded to broadly truncate, relatively unmodified; processes moderately setulose. T6 absent or

present as two lateral sclerites or broad dorsally continuous sclerite. Ejaculatory apodeme with fan-shaped

portion subequal to 1 .5X wider than width of hypandrial apodeme. Pregonite in profile smoothly curved

anteriorly (Fig. 90) to rather sharply bent anteriorly at midlength, and more or less pointed apically;

posteriorly bare in single specimen of S. (A.) sp. 2, with tiny to medium-sized seta in 5. arizonica , short seta

in single examined specimens of S. atoma and 5. (A.) spp. 3 and 4 and medium-sized seta in single examined

specimen of S. (A.) sp. 1. Epiphallus absent. Distiphallus reduced posteriorly, not incised to slightly incised

laterally, with enlarged teeth anterolaterally in S. arizonica (Fig. 124) and to lesser extent S. atoma , teeth not

enlarged beyond size of adjacent spinules in S. (A.) spp. 1 to 4, in profile tapered to rounded or pointed tip.

Postgonite apically rounded. Surstylus short to rather long and straight; basally free from epandrium. Cerci

Systematics of the Genus Group Taxa of the Siphonini

91

in profile short in S. arizonica and S. atoma, average length in S. (A.) spp. 1 to 4; straight to smoothly curved

along posterior margin; slightly broadened at midlength in posterior view in some specimens of S. arizonica

and single dissections of S. (A.) spp. 1-4 (Fig. 143), smoothly tapered to tip in some specimens of S.

arizonica and single dissection of S. atoma (similar to Fig. 144); moderately setose on basal half.

Examined male genitalia of: S. arizonica, S. atoma and S. (A.) spp. 1-4.

Female genitalia. — Short. S6 with average length hairs. T6 absent or present as median sclerite;

spiracles of segment 6 in membrane dorsal to lateral margins of S6. S7 with long anterior apodeme; not

keeled posteromedially. T7 absent or present as small lateral sclerites; spiracles of segment 7 in membrane

between segments 6 and 7. SH distinctly developed, haired. Tw present as two sclerites.

Examined female genitalia of: S. arizonica.

Hosts. Unknown.

Phylogenetics

Adults of this subgenus of six species (two described and four undescribed) share

a number of external characteristics (see Recognition and Description sections), but

none is considered synapotypic of the group.

Monophyly of S. (Aphantorhapha) is weakly supported by two slightly derived,

but apparently unique, states in the male genitalia. One is the tapered shape of the

distiphallus (Fig. 124). The other is the broadened cerci (Fig. 143), which were

observed in the single dissections of S. (A.) spp. 1-4 and some dissections of S.

arizonica specimens (i.e. S. arizonica with cerci varied from average to broadened).

Though the cerci of the single dissected male of S. atoma are of average shape, this

species is very similar in all other respects to other members of S. ( Aphantorhapha ).

In fact, the short cerci and toothed distiphallus of S. atoma suggests it is sister

species to S. arizonica.

Siphona pulla, described by Reinhard in Aphantorhapha , is known only from the

female holotype. The type does not agree in all respects with the present concept of

S. ( Aphantorhapha ). and cannot be placed into another supraspecific taxon of

Siphona s.l. without study of male genitalic characters. S. pulla is therefore left as an

unplaced species of Siphona s.l.

Siphona ( Aphantorhapha ) is admittedly not very distinctive, and is only

recognized as a subgenus because it is apparently monophyletic and because

synapotypies between it and other Siphona s.l. groups (which would permit a larger

and more broadly defined subgenus to be recognized) are unknown.

Geographic distribution

The two described species of S. ( Aphantorhapha ) and S. (A.) spp. 3 and 4 are

primarily southern Nearctic in distribution. S. arizonica is the most widely

distributed, ranging from Arizona and western New Mexico through Durango,

Chihuahua and Oaxaca to San Cristobal, Chiapas. S. atoma is only known from the

type series collected from College Station, Texas. S. (A.) sp. 3 is known from one

male collected from Grant Co., New Mexico, and S. (A.) sp. 4 from two males

collected near La Ciudad, Durango.

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Two new species of S. (Aphantorhapha) are recognized from the Neotropics.

Siphona (A.) sp. 1 is known from a single male collected from Ibarra, in the

Imbabura province of Ecuador (2200m, 2-4. VI. 1977). The other species, S. (A.) sp.

2, is known from several specimens collected over several years from Nova

Teutonia, SE Brazil.

List of described species included in Siphona (. Aphantorhapha )

N,S arizonica (Townsend), 1919: 586 (. Aphantorhapha ). Holotype male, USA:

Arizona, Chiricahua Mtns. (USNM). Holotype examined.

N atoma (Reinhard), 1947: 19 ( Aphantorhapha ). Holotype male, USA: Texas,

College Station (CNC). Holotype examined.

List of examined, undescribed, species included in Siphona (Aphantorhapha)

S. ( Aphantorhapha ) sp. 1: One male from Ecuador (BMNH).

5. ( Aphantorhapha ) sp. 2: Three males from SE Brazil (CNC, USP).

S. (Aphantorhapha) sp. 3: One male from New Mexico, USA (JEOH).

S. (Aphantorhapha) sp. 4: Two males from Durango, Mexico (CNC).

Siphona (subgenus Aphantorhaphopsis Townsend)

Figs. 64,91-92, 125.

Aphantorhaphopsis Townsend, 1926a: 34. Type-species, A. orientalis Townsend, 1926 (original

designation). New subgeneric status in Siphona Meigen.

Asiphona Mesnil, 1954: 9, 10 (as subgenus of Siphona). Type-species, Thryptocera selecta Pandelle, 1894

(original designation). New synonym of Aphantorhaphopsis Townsend.

Recognition

This taxon was called Asiphona Mesnil previously (e.g. Andersen 1983), but

Aphantorhaphopsis has priority and is used here. It comprises a probably

non-monophyletic assemblage of Old World Siphona s.l. species not belonging to

Siphona s.s. or S. (Ceranthia) [other Siphona subgenera are strictly New World in

known distribution]. The rationale for recognizing such an unsatisfactorily-defined

group in the formal classification of the Siphonini is explained below in the

Phylogenetics section.

Species of S. (Aphantorhaphopsis) have a short aristomere 1, lower proepimeral

seta undeveloped, lower katepistemal seta at least length of upper anterior one,

katepistemum without row of hairs anterior to mid coxa, and anal vein extended to

wing margin. These states distinguish S. (Aphantorhaphopsis) species from all Old

World non -Siphona s.l. species.

Siphona s.s. species have a characteristically elongate proboscis, with labella

rigid basally and at least as long as the prementum. Most S. (Aphantorhaphopsis)

species have padlike labella, and are thus easily separated from Siphona s.s. species

Systematics of the Genus Group Taxa of the Siphonini

93

by this state. The labella of a few S. (Aphantorhaphopsis) species are lengthened,

and in a very few as long as prementum, but only in Nepal sp. 1 are they also

inflexible along basal half (in life) and with reduced number of pseudotracheae as in

Siphona s.s. species. Nepal sp. 1 is distinguished from Siphona s.s. species by its

possession of a seta posteriorly on the pregonite (this seta is absent from Siphona

s.s. species and present in, or absent from, S. ( Aphantorhaphopsis ) species).

S. ( Ceranthia ) species are characterized by a cylindrical palpus (Fig. 16), long

seta posteriorly on the pregonite (Fig. 94) and distinctive shape of the distiphallus

(Fig. 127). Two S. (Aphantorhaphopsis) species, Australia sp. 1 and New Guinea sp.

1, have a cylindrical palpus like in S. ( Ceranthia ) species, but these do not have the

male genitalic states of that taxon (they have a bare pregonite and the distiphallus is

not laterally incised).

Description

Length: 3. 0-5. Omm.

Head. — Anterior proclinate orbital seta subequal to or longer than posterior one. Eye of male

medium-small to large, 0.69-0.89 head height; eye of female smaller than or subequal to eye of male.

Flagellomere 1 of male short to medium-long, 0.38-0.63 head height; shape linear to broad in most species,

subquadrangular in a few. Flagellomere 1 of female subequal or smaller than in male. Aristomere 1 short.

Aristomere two 1 .5-4X longer than wide. Aristomere 3 long and evenly tapered in most species, short and

thickened to near tip in a few; almost bare to short plumose. Clypeus narrow and enclosed in membrane to

U-shaped. Palpus short and clavate in most species, long in some species with an elongate proboscis,

reduced and cylindrical (as in S. ( Ceranthia ) spp.) in Australia sp. 1 and New Guinea sp. 1. Proboscis with

prementum short in most species, slightly lengthened to elongate in a few; labella padlike in most species,

slightly lengthened to subequal prementum length in a few with more than basal half flexible in life (e.g. S.

alticola, S. crassulata, S. fera, Nepal sp. 2 and New Guinea sp. 3), very long and Siphona- like (i.e. labella

inflexible over at least basal half and number of pseudotracheae reduced) in Nepal sp. 1 (proboscis 2.6X

head height).

Thorax. — Prostemum setulose, except bare in Nepal sp. 3. Lower katepistemal seta subequal in length

to, or longer than, upper anterior seta. Three or four postsutural dorsocentral setae. Upper part of

anepistemum with one setala in most species, two in a few. Fore tibia with preapical ad seta much shorter

than d seta in almost all species, known to be as long as d seta only in S. laticornis. Mid tibia with one ad

seta in most species (short in Australia sp. 1), two ad setae in Kenya sp. 1 and South Africa sp. 1. Claws

short. Wing vein CuAx with distal portion 0.26- 1.4X length of proximal portion (mean 0.47). Wing setulae:

R] dorsally bare or distally setulose, ventrally bare in most species, distally setulose in a few; R4+5 setulose

between base and r-m in most species, known to be setulose beyond r-m only in Australia sp. 2; CuA] bare.

Abdominal terga 1-5. — Abdomen ovoid in shape. 7j+2 without median marginal setae, lateral marginal

setae absent to strong. T3-T5 average or with weak lateral discal setae.

Male genitalia (Figs. 64, 91-92, 125). — S5 with posterior margins of processes approximately U-shaped

in most species, obtusely angled in a few, almost V-shaped in Nepal sp. 1 (Fig. 64, resembling typical Actia

shape except median cleft more distinct); apical lobe distinctly differentiated, apex slightly to markedly

curved inward in several species; median lobe narrowly to broadly rounded or truncate, relatively

unmodified; processes moderately setulose. T6 varied from two small lateral sclerites to single, narrow to

broad, dorsally continuous sclerite. Hypandrial apodeme lengthened in New Guinea sp. 1, very elongate in

Nepal sp. 4. Ejaculatory apodeme with fan-shaped portion 0.5-1.5X wider than hypandrial apodeme.

Pregonite (Figs. 91-92) in profile curved anteriorly and pointed apically in most species, in others varied

from short and broad to long and thin, in S. nigronitens expanded basally and fused with hypandrial

apodeme, in Kenya sp. 1 expanded ventrolaterally; bare or with small seta posteriorly in most species, large

seta present in S. nr. fera (absent from holotype of S. fera). Epiphallus present or absent. Distiphallus (Fig.

125) reduced posteriorly, markedly varied in profile, laterally incised or complete, apically pointed or

rounded or truncate, with or without recurved spines along anterior and/or lateral margin, in about half the

Quaest. Ent., 1989, 25 (1,2)

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species with long posterolateral arm. Postgonite long and pointed in Australia sp. 2, very reduced in S.

nigronitens and Nepal sp. 4, rounded or truncate in other species. Surstylus markedly varied, short to long,

thin to broad, straight to curved posteriorly; basally fused with epandrium in Nepal sp. 4, free from

epandrium in other species. Cerci average length and smoothly curved in most species, rather short in a few

species, slightly inflexed at midlength in a few species; moderately setose on basal half.

Examined male genitalia of: S. alticola, S. brunnescens, S. crassulata, S. fera, S. nr. fera, S. nigronitens, S.

orientalis, S. starkei, S. xanthosoma, Australia spp. 1-2, Kenya sp. 1, Nepal spp. 1-4, New Guinea spp. 1-3

and Uganda sp. 1. Examined published figure of: S. siphonoides (Andersen 1983, fig. 19).

Female genitalia. — Short. S6 with average length hairs. T6 absent (not examined in Australia sp. 1 and

Nepal sp. 4); spiracles of segment 6 in membrane dorsal to lateral margins of S6. S7 with long anterior

apodeme; not keeled posteromedially. Tn absent from most species, present as two small lateral sclerites in

Nepal sp. 1; spiracles of segment 7 in membrane between segments 6 and 7. S8 distinctly developed, haired,

broader than average in a few species, large and rather pointed in Australia sp. 1. T10 absent from Uganda sp.

1, present as lightly to distinctly sclerotized pair of sclerites or median sclerite in other species.

Examined female genitalia of: Australia sp. 1, Kenya sp. 1, Nepal spp. 1,4, South Africa sp. 1 and Uganda

sp. 1. Examined published figure of: S. samarensis (Andersen 1983, fig. 5).

Hosts

Known hosts of S. ( Aphantorhaphopsis ) comprise a diverse assemblage of

Macrolepidoptera, as might be expected of a grade-based taxon such as this (Table

2). Five host families are represented, of which two, the Noctuidae and

Geometridae, are commonly recorded among other siphonines. The three remaining

families, the Hesperiidae, Liparidae and Arctiidae, are virtually unreported as hosts

of other siphonines (a single record being that of an arctiid host for Ceromya

bicolor). One European species, S. samarensis , is recorded from an economically

important pest, the gypsy moth ( Porthetria dispar ), though its level of parasitism is

apparently very low.

Given the possibly polyphyletic nature of S. ( Aphantorhaphopsis ) and the

meagre extent to which both its species and hosts are known, the diversity of hosts

here recorded is probably far from representative for the taxon.

Phylogenetics

The present grouping of Old World species under S. ( Aphantorhaphopsis ) is one

of taxonomic convenience, as too little is known about these species to formulate a

phylogenetically-based classification. Adults have the derived features of Siphona

s.l., and are assignable within the genus to S. (Aphantorhaphopsis) by their lack of

the apotypic states defining the other Siphona subgenera. It is inferred that this

group lacks autapotypies because it is composed of several (numerous?) lineages.

This taxon is probably either paraphyletic or polyphyletic, depending upon the true

relationships of these species with those of other Siphona subgenera.

The diverse assemblage of species comprising this group can only be adequately

reclassified by a thorough study of at least external and male genitalic characters.

Even a revision of described species is apt to be inadequate for both classificatory

and identification needs, as many undescribed species are known.

The description above of S. (Aphantorhaphopsis) is based upon the study of

specimens of as many species as were available. Though this characterization of S.

Systematics of the Genus Group Taxa of the Siphonini

95

(Aphantorhaphopsis) is recognized as an interim measure until a thorough revision

is undertaken, it is nevertheless useful, as it is the first comprehensive review of the

included species. Its concept is the same as that of Andersen’s (1983) Asiphona

Mesnil, over which the name Aphantorhaphopsis has priority.

Andersen similarly diagnosed this taxon as a group of Old World siphonines

belonging to the Siphona lineage and lacking the derived states of the other taxa (in

his case Siphona s.s. and S. (Ceranthia) because his revision dealt strictly with Old

World siphonines). Andersen found no autapotypies of Asiphona, but retained the

taxon as a genus and hypothesized a sister group relationship between it and

Ceranthia. He based this hypothesis on the shared possession in these taxa of a seta

on the posterior surface of the pregonite, which is absent from Siphona s.s. species.

However, the present study has shown that some species of S. ( Aphantorhaphopsis )

lack this seta and many New World Siphona s.l. species possess it, so presence of a

seta is not synapotypic of this taxon and S. ( Ceranthia ). The significance of this

seta, with respect to the phylogeny of the supraspecific taxa of Siphona s.l., is

discussed in the Evolution chapter.

An attempt was made while revising the supraspecific taxa of Siphona s.l. to

discover species of S. ( Aphantorhaphopsis ) which might have close New World

relatives. None was found, perhaps because New World subgenera are mostly

Neotropical and southern Nearctic in distribution and are thus likely to be too old to

have easily recognized Old World members, if indeed any exist. It was not clearly

established whether or not some of the unplaced Siphona s.l. species of the northern

Nearctic region are closely related to any Old World S. (Aphantorhaphopsis)

species, though this is a possibility.

Geographic distribution

The 21 described species of S. ( Aphantorhaphopsis ) are fairly evenly distributed

throughout the Palearctic, Afrotropical and Oriental regions, with six species

described from each of the first two regions and eight from the third. Only one

species is described from Australia. Examination of material from the Old World

indicates that at least several, perhaps many, undescribed species exist in each

region, particularly in the Oriental and Australian regions.

The six described Palearctic species are apparently restricted to Europe (Herting

1984 [species included under broad concept of Ceranthia ]; no records in

publications of Richter) and are keyed and redescribed in Mesnil (1963a- 1964:

843-853). Species of the other regions are catalogued in Crosskey (1973, 1976a,

1980), mostly in the genus Ceromya. Dear and Crosskey (1982) key three species

occurring in the Philippines, S. angustifrons , S. orientalis and S. nr. selangor (all

keyed as Ceromya species), and species of central Africa and Madagascar are keyed

in Mesnil (1954 and 1977b, respectively).

The distribution of S. (Aphantorhaphopsis) species cannot be historically

interpreted until the phylogenetic relationships among these species are better

Quaest. Ent., 1989, 25 (1,2)

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O’Hara

known, as well as their relationships with other subgenera of Siphona.

List of described species included in Siphona {. Aphantorhaphopsis )

O alticola (Mesnil), 1953: 110 ( Crocuta {Siphona)). Holotype male, Burma:

Kambaiti (ZMU). Holotype examined. New combination.

O angustifrons (Malloch), 1930b: 131 {Actia). Holotype male, Malaysia: Kedah

Peak (BMNH). Holotype examined. New combination.

P brunnescens (Villeneuve), 1921: 46 {Actia). Holotype male, German

Democratic Republic: Oberlausitz, Niederoderwitz (CNC). Holotype examined.

New combination.

O crassulata (Mesnil), 1953: 112 {Crocuta {Siphona)). Holotype male, Burma:

Kambaiti (ZMU). Holotype examined. New combination.

A fera Mesnil, 1954: 26 (described in subgenus Asiphona). Holotype male, Zaire:

Nyongera (MRAC). Holotype examined. New combination.

O laboriosa Mesnil, 1957: 48 (described in subgenus Asiphona). Holotype male,

Burma: Kambaiti (ZMU). Holotype examined. New combination.

O laticornis (Malloch), 1930b: 131 {Actia). Holotype male (not female),

Malaysia: Selangor (BMNH). Holotype examined. New combination.

O mallochiana (Gardner), 1940: 178 {Actia). Type(s) puparia, India (?FRI). New

combination.

syn. perispoliata (Mesnil), 1953: 108 {Actia). Holotype male, China:

Canton (BMNH). — Crosskey, 1976a: 213. Holotype examined.

A nigronitens Mesnil, 1954: 25 (described in subgenus Asiphona). Holotype male,

Zaire: Rutshuru (MRAC). Holotype examined. New combination.

U norma (Malloch), 1929a: 116 {Actia). Holotype male, Australia: New South

Wales, Como (USNM). Holotype examined. New combination.

O orientalis (Townsend), 1926a: 35 {Aphantorhaphopsis). Holotype male,

Indonesia: Sumatra, Fort de Kock (ZMA). Holotype examined.

A picturata (Mesnil), 1977b: 179 {Asiphona). Holotype male, Madagascar:

Belazao (MNHN). New combination.

A pudica Mesnil, 1954: 27 (described in subgenus Asiphona). Holotype male,

Zaire: Eala (MRAC). Holotype examined. New combination.

P samar ensis (Villeneuve), 1921: 46 {Actia). Holotype female, Russia: Kujbysev

[Samara] (CNC). Holotype examined. New combination.

O selangor (Malloch), 1930b: 132 {Actia). Holotype male, Malaysia: Selangor

(BMNH). Holotype examined. New combination.

P selecta (Pandelle), 1894: 112 {Thryptocera). Syntypes, France: Var, Hyeres

(MNHN). New combination.

P siphonoides (Strobl), 1898: 235 {Gymnopareia). Holotype male, Austria:

Steiermark, Gesause (NMBA). New combination,

syn. brunneipalpis (Villeneuve), 1921: 45 {Actia). Holotype male. Federal

Systematics of the Genus Group Taxa of the Siphonini

97

Republic of Germany: Harz (CNC). — Mesnil, 1963a: 848. Holotype

examined.

A speciosa Mesnil, 1954: 28 (described in subgenus Asiphona). Holotype male,

Zaire: Rutshuru (MRAC). Holotype examined. New combination.

P starkei (Mesnil), 1952b: 155 ( Actia ). Holotype male, German Democratic

Republic: nr. Bautzen, Hennerstadt (CNC). Holotype examined. New

combination.

P verralli (Wainwright), 1928: 208 {Actia). Holotype male, Scotland:

Sutherlandshire, Mound (HDE). Holotype examined. New combination.

A xanthosoma Mesnil, 1954: 28 (described in subgenus Asiphona). Holotype

male, Zaire: Rwindi (MRAC). Holotype examined. New combination.

List of examined, undescribed, species included in Siphona

(Aphantorhaphopsis)

S. (Aphantorhaphopsis) Australia sp. 1: One male, three females from Queensland (DPI).

S. (Aphantorhaphopsis) Australia sp. 2: One male from Karumba, Queensland (DPI).

S. ( Aphantorhaphopsis ) Kenya sp. 1: One male and several females from Kenya (CAS, USNM).

S. (Aphantorhaphopsis) Nepal sp. 1: Males and one female from Nepal (CNC).

S. (Aphantorhaphopsis) Nepal sp. 2: One male, one female from Kathmandu (CNC).

S. (Aphantorhaphopsis) Nepal sp. 3: Two males from Nepal (CNC).

S. (Aphantorhaphopsis) Nepal sp. 4: One male, one female from Nepal (CNC).

S. (Aphantorhaphopsis) New Guinea sp. 1: Males and one female from New Guinea (BPBM).

S. (Aphantorhaphopsis) New Guinea sp. 2: Three males from ne. Mur Mur P. (BPBM).

S. (Aphantorhaphopsis) New Guinea sp. 3: One male from se. Mt. Saint Mary (BPBM).

S. (Aphantorhaphopsis) South Africa sp. 1: Two females from Cape Good Hope Nat. Res. (USNM).

S. (Aphantorhaphopsis) Uganda sp. 1: Males and females from Uganda, one female from Rhodesia (CNC,

USNM).

Siphona (subgenus Baeomyia O’Hara)

Figs. 15,22, 38, 93, 126.

Baeomyia O’Hara, 1984: 1390. Type-species, Aphantorhapha hurdi Reinhard, 1959 (original designation).

New subgeneric status in Siphona Meigen.

Recognition

Adults of S. ( Baeomyia ) species are among the smallest of siphonines, measuring

2-3mm in length. They are only recorded from western North America and are

easily recognized among New World siphonines by the absence of an ad seta on the

mid tibia (Fig. 38) - a state only shared with several Old World Actia and Ceromya

s.s. species. Also diagnostic is the elongate aristomere 1, which is subequal in length

to aristomere 2 (Fig. 15; only as long in a few Old World siphonines). S. ( Baeomyia )

species are also characterized by padlike labella, three postsutural dorsocentral

setae, wing vein R4+5 not setulose beyond crossvein r-m, anal vein extended to wing

margin, and crossvein dm-cu far removed from wing margin (Fig. 22).

Quaest. Ent., 1989, 25 (1,2)

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O’Hara

Description

Length: 2.0-3.0mm.

Head (Fig. 15; also see head profiles in O’Hara 1984, figs. 1-5.). — Anterior proclinate orbital seta

subequal in length to posterior one. Eye of male and female subequal, small to medium-large, 0.65-0.83 head

height. Flagellomere 1 of male short to medium length, 0.39-0.56 head height; shape average to broadly

subquadrangular. Flagellomere 1 of female smaller than in male or subequal in size. Aristomere 1 elongate,

subequal in length to aristomere 2, each 2.4-4.6X longer than wide. Aristomere 3 almost bare, very short,

evenly tapered or thickened to near tip. Clypeus U-shaped. Palpus short, clavate. Proboscis with prementum

short, labella padlike.

Thorax (Fig. 22). — Prosternum bare or sparsely setulose. Lower katepistemal seta shorter than or

subequal in length to upper anterior seta. Three postsutural dorsocentral setae. Upper part of anepistemum

with single setula. Fore tibia with preapical ad seta much shorter than d seta. Mid tibia lacking ad seta.

Claws short. Wing (Fig. 22 and O’Hara 1984, fig. 10); CwA, with distal portion 0.64- IX length of proximal

portion (i.e. dm-cu far removed from wing margin; mean 0.85). Wing setulae: /?, dorsally bare or with one

setula apically on bend, ventrally bare; R4+5 setulose between base and r-m\ CuAl bare.

Abdominal terga 1-5. — Abdomen ovoid in shape. T]+2 without median or lateral marginal setae. TyTs

with average setation, setae weakly to strongly developed.

Male genitalia (Figs. 93, 126 and O’Hara 1984, figs. 6-9). — S5 with posterior margins of processes

approximately U-shaped; apical lobe distinctly differentiated; median lobe rounded, relatively unmodified;

processes sparsely to moderately setulose. T6 narrow to broad, dorsally continuous. Ejaculatory apodeme

with fan-shaped portion 1.0-1.5X wider than hypandrial apodeme. Pregonite (Fig. 93) in profile curved

anteriorly, pointed apically; with tiny seta posteriorly (not shown in O’Hara 1984). Epiphallus absent.

Distiphallus (Fig. 126) reduced posteriorly, not incised laterally, without enlarged teeth anteriorly or

ventrally, in profile apically truncate. Postgonite short to average in size, apically rounded. Surstylus almost

straight to sharply curved posteriorly; basally free from epandrium. Cerci short to average length, smoothly

curved (O’Hara 1984, figs. 7-9) or sharply inflexed at midlength (O’Hara 1984, fig. 6); moderately setose on

basal half.

Examined male genitalia of: S. hurdi, S. juniperi, S. sonorensis and S. xanthogaster (all shown in O’Hara

1984, figs. 6-9).

Female genitalia. — Short. S6 with average length hairs. T6 absent; spiracles of segment 6 in membrane

dorsal to lateral margins of S6. S7 with long anterior apodeme; not keeled posteromedially. T7 absent;

spiracles of segment 7 in membrane between segments 6 and 7. S8 distinctly developed, haired. 7j0 absent.

Examined female genitalia of: S. antennata, S. hurdi, S. sonorensis and 5. xanthogaster (all shown in O’Hara

1984, figs. 1 1-14 [fig. 14, S. antennata, mislabelled as B. juniperi]).

Hosts

Hosts of two S. (Baeomyia) species are known, and are larvae belonging to the

geometrid genus Semiothisa (Table 2 and O’Hara 1984).

Phylogenetics

S. (Baeomyia) is a monophyletic taxon based on several synapotypies (see

O’Hara 1984: 1388-1389). These include absence of an ad seta on the mid tibia (this

seta independently lost in a few Old World Actia and Ceromya s.s. species),

aristomere 1 subequal in length to aristomere 2 (a rare state present in a few,

unrelated, Old World siphonines), very small-sized adults, very short aristomere 3

(figs. 1-5 in O’Hara 1984), and crossvein dm-cu far removed from wing margin

(Fig. 22). The last state is possibly not autapotypic of S. ( Baeomyia ) because it is

shared with some other Siphona s.l. species (see Table 1); the position of dm-cu is

such a labile character among siphonines that it is difficult to establish among which

lineages its derived state (i.e. far removed from wing margin) is synapotypic, and

Systematics of the Genus Group Taxa of the Siphonini

99

among which it is convergent (O’Hara 1984: 1388).

Geographic distribution

S. (Baeomyia) species are restricted in distribution to western North America.

Known distributions are shown and discussed in O’Hara (1984).

List of described species included in Siphona (. Baeomyia )

N antennata (O’Hara), 1984: 1393 {Baeomyia). Holotype male, USA: Arizona,

Graham Co., near Marijilda canyon (CAS). Holotype examined.

N hurdi (Reinhard), 1959: 161 ( Aphantorhapha ). Holotype male, USA:

California, Panamint Mtns. (CAS). Holotype examined.

N juniperi (O’Hara), 1984: 1395 {Baeomyia). Holotype male, Canada: British

Columbia, Williams Lake (CNC). Holotype examined.

N sonorensis (O’Hara), 1984: 1393 {Baeomyia). Holotype male, USA: Arizona,

Graham Co., 2.4km. west on Hwy. 366 from Hwy. 666 (CNC). Holotype

examined.

N xanthogaster (O’Hara), 1984: 1394 {Baeomyia). Holotype male, Canada:

British Columbia, Vermilion (CNC). Holotype examined.

Siphona (subgenus Ceranthia Robineau-Desvoidy)

Figs. 16, 44, 46, 65,94, 127.

Ceranthia Robineau-Desvoidy, 1830: 88. Type-species, C. fulvipes Robineau-Desvoidy, 1830 (by

designation of Robineau-Desvoidy, 1863: 685) = Ceromya abdominalis Robineau-Desvoidy, 1830. New

subgeneric status in Siphona Meigen.

Recognition

Members of this subgenus are widely distributed, and with few exceptions are

easily recognized. Adults are characterized externally by a cylindrical palpus (Fig.

16), a derived state within the Siphonini. This state is also present in S.

( Aphantorhaphopsis ) Australia sp. 1 and New Guinea sp. 1, but these species do not

have S. (Ceranthia)- like male genitalia (see below), and seem more closely related

to certain species of S. ( Aphantorhaphopsis ).

Two derived features of the male genitalia are also characteristic of S.

( Ceranthia ) species. The posterolateral margins of the distiphallus are incised to

varied degrees, with distinct posterolateral arm (one per side; Fig. 127). In addition,

a seta projects posteriorly from the pregonite (Fig. 94), and is larger than in all but a

very few other Siphona species. Atypical in this respect is S. ( Ceranthia ) Mauritius

sp. 1 , in which this seta is absent. This species possesses a cylindrical palpus and S.

(Ceranthia)- like distiphallus, and is recognized as a member of this subgenus by

these features.

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O’Hara

Adult females of S. (Ceranthia) species have a posteromedially keeled (Figs. 44,

46) sternum 7, and though this state is not unique to species of this subgenus, it is

apparently universal among its members.

Description

Length: 3.0-5.5mm.

Head (Fig. 16). — Anterior proclinate orbital seta subequal in length to posterior one in most species.

Eye of male medium-small to large, 0.69-0.88 head height; eye of female usually smaller than in male.

Flagellomere 1 of male medium-short to medium-long, 0.42-0.66 head height; shape broad to

subquadrangular. Flagellomere 1 of female shorter and narrower than in male. Aristomere 1 short.

Aristomere two 2.0-8. OX longer than wide, generally less than 4X. Aristomere 3 almost bare to

micropubescent, rather short and evenly tapered. Clypeus varied from narrow and enclosed in membrane to

U-shaped. Palpus short or reduced, cylindrical (not apically clavate). Proboscis with prementum short,

labella padlike.

Thorax. — Prostemum setulose. Lower katepistemal seta longer than upper anterior seta. Three or four

postsutural dorsocentral setae. Upper part of anepistemum with single setula in most species, with two in a

few. Fore tibia with preapical ad seta much shorter than d seta. Mid tibia with one ad seta. Claws short.

Wing vein CwA, with distal portion 0.30-0.51 length of proximal portion ( i.e . dm-cu near wing margin; mean

0.42). Wing setulae: /?, bare dorsally and ventrally in most species, distally setulose dorsally in a few,

distally setulose dorsally and ventrally in S. terrosa; R4+5 setulose between base and r-m in most species,

beyond r-m in S. pallida-, CuA] bare.

Abdominal terga 1-5. — Abdomen ovoid in shape. Ti+2 without median marginal setae; lateral marginal

setae strong. T3-T5 average in most species, with extra pair of lateral marginal setae on T3 in a few species.

Male genitalia (Figs. 65, 94, 127). — S5 (Fig. 65) little varied; posterior margins of processes

approximately U-shaped; apical lobe large and distinctly differentiated, in most species apex curved inward;

median lobe rounded, relatively unmodified; processes sparsely to moderately setulose. T6 narrow to broad,

dorsally continuous or narrowly discontinuous. Ejaculatory apodeme with fan-shaped portion 1.0-1.5X

wider than hypandrial apodeme. Pregonite (Fig. 94) in profile curved anteriorly, more or less pointed

apically; large seta posteriorly (except bare in Mauritius sp. 1). Epiphallus absent. Distiphallus (Fig. 127)

reduced posteriorly, laterally incised to form posterolateral arm extended slightly to markedly beyond

anterior margin; anterior margin spinulose or toothed. Postgonite apically rounded or truncate. Surstylus

straight, extended slightly beyond tip of cerci; basally free from epandrium. Cerci average length, smoothly

curved at midlength; moderately setose on basal half.

Examined male genitalia of: S. abdominalis (fig. 20 in Andersen 1983, as Ceranthia fulvipes ), S.flavipes, S.

plorans, S. scutellata, Ethiopia sp. 1, Mauritius sp. 1 and U.S. spp. 1-5.

Female genitalia (Figs. 44, 46). — Short to elongate (e.g. U.S. sp. 7). S6 with average length hairs,

ventrally flat or slightly keeled posteromedially. T6 absent; spiracles of segment 6 in membrane dorsal to

lateral, or anterolateral, margins of S6. S7 with long anterior apodeme; distinctly keeled posteromedially, in a

few species posterior margin narrow and elongate and covering S8. T7 absent; spiracles of segment 7 in

membrane dorsal to anterolateral margins of S7. S8 distinctly developed, haired. T]0 present as two sclerites.

Examined female genitalia of: S. abdominalis, S.flavipes and U.S. spp. 1-7. Examined published figure of:

S. tenuipalpis (Andersen 1983, fig. 6).

Hosts

The hosts of four S. ( Ceranthia ) species are known, and these all belong to the

Geometridae (Table 2).

Phylogenetics

S. ( Ceranthia ) is a clearly defined, monophyletic lineage of Siphona s.l. The

cylindrical palpus of adults (Fig. 16) has long been recognized as the distinguishing

feature (and synapotypy) of this group, and to this are added two genitalic states

Systematics of the Genus Group Taxa of the Siphonini

101

which further support monophyly of the subgenus: the distiphallus is

characteristically incised posterolaterally, with distinct posterolateral projection

(Figs. 127), and the pregonite possesses a large seta posteriorly (Fig. 94). Females

possess a posteromedially keeled (Figs. 44, 46) sternum 7, and larvae (so far as

known) are parasitic only on larval geometrids, but these characteristics may not be

autapotypic of S. (Ceranthia) species.

Two species assigned to S. ( Aphantorhaphopsis ) also have a cylindrical palpus -

Australia sp. 1 and New Guinea sp. 1 . These species do not possess other derived

states of S. ( Ceranthia ), and seem to have developed a cylindrical palpus

independently of S. ( Ceranthia ) species. Other characters suggest that they are

closely related to species with a normal palpus belonging to the broadly defined S.

( Aphantorhaphopsis ). With exclusion of these species from S. ( Ceranthia ), no

species of this subgenus are known from the Australian region.

S. ( Ceranthia ) Mauritius sp. 1 has the derived states of the palpus and

distiphallus found in other S. ( Ceranthia ) species, but lacks the large seta on the

pregonite which is present in other known species of this taxon. I hypothesize that

this seta was present in the groundplan of S. ( Ceranthia ) because at least a small seta

is widely distributed among other Siphona lineages (Table 1). Therefore lack of this

seta from Mauritius sp. 1 is interpreted as a secondary loss.

Geographic distribution

Eight described S. ( Ceranthia ) species are in the Palearctic region, five in the

Afrotropical region and one in the Nearctic region. Five of the eight Palearctic

species are apparently very limited in distribution, with one each in England ( S .

lichtwardtiana ), northern France ( S . tristella) and Algeria ( S . jocosa ), and two in

Japan ( S . japonica and S. sulfurea). More widespread are S. abdominalis (common

throughout Europe and collected from Mongolia and Chita region (Herting 1973 [as

“ Ceranthia sp. aff. anomala Zett.” - perhaps a new species], Richter 1975, 1980), S.

tenuipalpis (described from Berlin, and Zlatoust in the Ural Mtns.) and S. pallida

(described from Austria, and if correctly identified then also found on the Kuril

Islands (Richter 1976b)). A key to the described Palearctic species of S. ( Ceranthia )

is presented in Mesnil (1975: 1399-1400), and distributions are summarized in

Herting 1984 (with species of S. ( Aphantorhaphopsis ) included in Ceranthia ). I

have seen specimens of a new Palearctic species from Ethiopia.

Known ranges of the five described Afrotropical S. ( Ceranthia ) species are given

in Crosskey (1980). All were described by Mesnil, and their distributions reflect his

work on the fauna of the upper Congo area (four species) and Madagascar (one

species). I suspect that the S. ( Ceranthia ) fauna of the Afrotropical region is much

more diverse than presently recognized.

There are no described S. ( Ceranthia ) species in the Oriental and Australian

regions. I have not seen any specimens of S. ( Ceranthia ) species among borrowed

Australian siphonines, but examined one female belonging to the subgenus from

Quaest. Ent., 1989, 25 (1,2)

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O’Hara

Coimbatore in southern India (CNC specimen).

The New World S. (Ceranthia) are very inadequately known. The single

described species, S. flavipes, is recorded from eastern Canada and northeastern

USA. Approximately six undescribed S. ( Ceranthia ) species occur in North

America, and are mostly eastern or western in distribution. The most southern

record for S. ( Ceranthia ) in the New World is southwestern New Mexico, USA.

List of described species included in Siphona (I Ceranthia )

P abdominalis (Robineau-Desvoidy), 1830: 87. ( Ceromya ). Type(s), France

(lost).

syn. fulvipes (Robineau-Desvoidy), 1830: 88 ( Ceranthia ). Holotype male,

France: Saint Sauveur (lost). — Herting, 1974: 18.

microcera (Robineau-Desvoidy), 1830: 88 {Ceromya). Type(s),

France: near Paris (lost). — Mesnil, 1963a: 840.

anomala Zetterstedt, 1849: 3213, 3221. Two syntypes, Denmark

(UZI).— Herting, 1974: 18.

vivida (Robineau-Desvoidy), 1850: 196 {Ceromya). Types, France:

near Paris (lost). — Herting, 1974: 18.

grisea (Robineau-Desvoidy), 1850: 198 {Ceromya). Type(s) male,

France: near Paris (lost). — Bezzi and Stein, 1907: 389.

flavipes (Robineau-Desvoidy), 1850: 200 {Ceranthia). Unjustified

emendation of C. fulvipes Robineau-Desvoidy, 1830.

N flavipes (Coquillett), 1897: 58 {Thryptocera). Holotype female, USA: New

Hampshire, White Mtns. (USNM). Holotype examined.

P japonica (Mesnil), 1963b: 33 {Ceranthia). Holotype male, Japan: Kyushu

(CNC). Holotype examined.

P jocosa (Villeneuve), 1942b: 134 {Actia). Holotype male, Algeria: Algiers

(CNC). Holotype examined.

A lacrymans (Mesnil), 1954: 24 {Ceranthia). Holotype male, Zaire: Karisimbi

(MRAC). Holotype examined.

P lichtwardtiana (Villeneuve), 1931: 61 (as variety of Actia anomala

(Zetterstedt)). Type, locality not given (ZMHU; not located, possibly lost).

A livoricolor (Mesnil), 1977b: 178 {Ceranthia). Holotype female, Madagascar:

Anjavidilava (MNHN).

P pallida (Herting), 1959: 423 {Ceranthia {Actia)). Holotype female, Austria:

Aflenz, Dorfmeister (NMV). Holotype examined.

A plorans (Mesnil), 1954: 24 {Ceranthia). Holotype male, Rwanda: Sabinyo

(MRAC). Holotype examined.

A scutellata (Mesnil), 1954: 22 {Ceranthia). Holotype male, Zaire: Rweru

(MRAC). Holotype examined.

P sulfur ea (Mesnil), 1971: 72 {Ceranthia). Holotype female, Japan: Hokkaido,

Systematics of the Genus Group Taxa of the Siphonini

103

Nukabira (CNC).

P tenuipalpis (Villeneuve), 1921: 46 ( Actia ). Two male syntypes: Berlin, Federal

Republic of Germany and Zlatoust, USSR (CNC). Syntypes examined.

A terrosa (Mesnil), 1954: 23 ( Ceranthia ). Holotype male, Rwanda: Kundhuru

(MRAC). Holotype examined.

P tristella (Herting), 1966: 5 ( Ceranthia ). Holotype male, Switzerland: Wallis,

Tanay (ETH). Holotype examined.

List of examined, undescribed, species included in Siphona ( Ceranthia )

S. ( Ceranthia ) Ethiopia sp. 1: One male from Addis Abbaba (AMNH).

S. ( Ceranthia ) Mauritius sp. 1: Two males from Les Mares (CNC).

S. ( Ceranthia ) U.S. sp. 1: One male, females ranging from Washington state to California, USA (PHA,

WSUP).

S. ( Ceranthia ) U.S. sp. 2: Two males, females from northeastern North America (CAS, CNC, MSU).

S. (Ceranthia) U.S. sp. 3: One male, one female from New Mexico, USA (JEOH).

S. ( Ceranthia ) U.S. sp. 4: Males and females from northeastern North America (CNC, PHA, USNM, WLD,

WSUP).

S. ( Ceranthia ) U.S. sp. 5: Males and females from California, USA (CAS, JEOH, PHA, USNM).

S. ( Ceranthia ) U.S. sp. 6: Two females from British Columbia, Canada (CAS, CNC).

S. (Ceranthia) U.S. sp. 7: One female from Nevada, USA (USNM).

Siphona (subgenus Pseudosiphona Townsend)

Figs. 17, 66, 95, 128, 140, 162.

Pseudosiphona Townsend, 1916: 622. Type-species, Siphona brevirostris Coquillett, 1897 (original

designation). New subgeneric status in Siphona Meigen.

Recognition

This taxon is one of the more diverse subgenera of New World Siphona s.l.,

despite the fact that only its type species, S. brevirostris , is described. It comprises

two species north of Mexico and approximately 18 south of the United States.

Adults of most S. ( Pseudosiphona ) species have a characteristic habitus,

distinctive male genitalia, and unique larval cephalopharyngeal skeleton. However,

interspecific variation is common and certain species depart from the average

condition in one or more states. Because variation from the typical features is

particularly prevalent among external characters, and some other Siphona s.l.

species appear externally similar, examination of male genitalia is essential for the

recognition of members of this subgenus. The following diagnosis includes a suite

of characteristics shared by most species of S. (Pseudosiphona), though some

species deviate from it in one or more states: adults are light colored with mostly

yellow legs, light brown thoracic dorsum, and generally heavy pruinosity on the

anterior fourth of abdominal terga and very sparse pruinosity over rest of abdomen

(in contrast to the more evenly pruinose abdomens of most other New World

Siphona s.l. species). Head features are markedly varied, but the following

Quaest. Ent., 1989, 25 (1,2)

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O’Hara

combination of states is common to most S. (Pseudosiphona) species (Fig. 17): row

of short hairs extended from parafrontal to halfway down parafacial, medium-sized

flagellomere 1 , arista distinctly haired, eye medium-large, and proboscis with labella

slightly more than half length of prementum.

S. ( Siphonopsis ) is a very diverse Neotropical taxon and its members can

generally be distinguished externally from those of S. ( Pseudosiphona ) by their

more pruinose abdomen, shorter row of parafacial hairs, almost bare arista and

slightly longer labella. Some members of these taxa are rather similar despite these

differences between most species, but features of the male genitalia reliably separate

them.

S. (Pseudosiphona) sp. 15 is the least typical of the subgenus in head

characteristics, having an elongate prementum and labella like in Siphona s.s.

species. The male genitalia of this species deviate slightly from the average

condition (described below; not resembling the male genitalia of S. ( Siphona )

species), but features of the first instar confirm its placement here.

The male genitalia of most S. ( Pseudosiphona ) species are very distinctive. In

most species the pregonite (Fig. 95) is broadened, anterolaterally spinulose, and has

one to several tiny setae posteriorly; possession of all three of these states is

apparently unique to S. ( Pseudosiphona ). In a few species the pregonite is more

J-shaped. Spinules are present in all examined specimens, but this state is shared

with other taxa (though these other taxa have a more slender pregonite). The

possession of more than one posterior seta on the pregonite is apparently unique to

S. ( Pseudosiphona ) species, though at least one species has only one seta as in some

other Siphona s.l. species.

The distiphallus of most S. ( Pseudosiphona ) species is unusually uniform in

shape (Fig. 128). It is triangular and apically pointed in profile and anterolaterally

spined in all but a very few species, and this shape is unique to S. (Pseudosiphona).

Few species have an apically truncate distiphallus, but these have the typical states

of the pregonite.

The cerci of most S. (Pseudosiphona) species are sharply inflexed at midpoint

(Fig. 140). This state can be seen during external examination of most males and is

not present in many other New World Siphona s.l. species (see Table 1). This state

is not diagnostic of S. (Pseudosiphona) since some of its species have smoothly

curved cerci.

S. (Pseudosiphona) sp. 15, mentioned above as having a Siphona s.s. head

habitus, is easily recognized as a S. (Pseudosiphona) species by its male genitalia.

The pregonite is J-shaped rather than broadened, but has two tiny setae posteriorly

as in most other S. (Pseudosiphona) species. The distiphallus is of the typical S.

(Pseudosiphona) shape, though the cerci are smoothly curved in profile.

The cephalopharyngeal skeleton of first instars provides one of the best character

states by which to recognize S. (Pseudosiphona) species, though is not likely to be

routinely examined. The labrum is elongate and hook-like in most (but not all)

Systematics of the Genus Group Taxa of the Siphonini

105

species (Fig. 162), and this state is unique to S. ( Pseudosiphona ) species among

Siphona s.l. species (see O’Hara in press “a”).

Description

Length: 2.0-5.0mm.

Head (Fig. 17). — Proclinate orbital setae subequal in length or anterior one longer. Eye of male medium

to large, 0.73-0.86 head height; eye of female slightly smaller to slightly larger than in male. Flagellomere 1

of male short to medium-short, 0.38-0.48 head height; shape rather linear in most species, average to

subquadrangular in a few. Flagellomere 1 of female subequal or smaller than in male. Aristomere 1 short in

most species, as long as wide in at least one species. Aristomere 2 relatively short, 1.5-2X longer than wide.

Aristomere 3 short to long and evenly tapered, micropubescent to medium plumose. Clypeus narrow to

slightly broadened, enclosed in membrane. Palpus short, clavate. Proboscis with prementum short to

elongate (up to head height in length), between 0.5-0.7 head height in most species, labella slightly to

moderately lengthened, in most species between 0.6-0.7 prementum length, quite elongate (0.8- 1.0

prementum length) with basal half flexible or inflexible in life (latter state Siphona- like) in a few species.

Thorax. — Prostemum setulose. Lower katepistemal seta subequal in length or longer than upper

anterior seta. Three postsutural dorsocentral setae. Upper part of anepistemum with one or two setulae. Fore

tibia with preapical ad seta much shorter than d seta. Mid tibia with one ad seta. Claws short. Wing vein

CuAx with distal portion 0.36-0.74 length of proximal portion (mean 0.60). Wing setulae: /?, dorsally bare,

or distally or entirely setulose, ventrally bare. R4+5 setulose to or beyond r-m\ CuAx bare.

Abdominal terga 1-5. — Abdomen ovoid in shape. Tl+2 without median marginal setae, lateral marginal

setae absent from most species, weakly developed in a few. T3-T5 with average setation.

Male genitalia (Figs. 66, 95, 128, 140). — S5 (Fig. 66) with posterior margins of processes

approximately U-shaped (except almost V-shaped in 5. (P.) sp. 11, and resembling typical Actia shape

except median cleft more distinct); apical lobe distinctly differentiated in most species, rather short in S. (P.)

sp. 19, apex curved inward in a few species; median lobe broadly rounded to truncate and relatively

unmodified in most species, with accessory lobe slightly developed in a few species; processes moderately

setulose. T6 absent or present as two lateral sclerites. Ejaculatory apodeme with fan-shaped portion subequal

in width to width of hypandrial apodeme. Pregonite in profile broad along most of length and apically

rounded or pointed in most species (Fig. 95), rather J-shaped in a few; outer surface short spinose on apical

half or less and one to several (in most species several) tiny setae posteriorly. Epiphallus absent. Distiphallus

(Fig. 128) reduced posteriorly, not incised laterally, with enlarged spines anterolaterally in almost all

species, in profile triangular and apically pointed in most species, subquadrangular and apically truncate in

S. (P.) spp. 18 and 19, deeply incised anteriorly in most species. Postgonite apically rounded. Surstylus (Fig.

140) varied from average length to elongate, narrow to broad, in most species straight, in a few curved

posteriorly; basally free from epandrium. Cerci varied from short to elongate, in profile average width to

thick and smoothly curved (a few species) to sharply inflexed at midlength (most species, Fig. 140);

moderately (most species, Fig. 140) to densely (a few species) setose on basal half.

Examined male genitalia of: S. brevirostris, S. (P.) spp. 1-13,15,17-19.

Female genitalia. — Short to moderately extensible. S6 with short to long hairs apically, ventrally flat in

most species, slightly keeled posteromedially in S. (P.) sp. 15, sharply keeled posteromedially in S. (P.) sp.

17. T6 absent; spiracles of segment 6 in membrane dorsal to lateral margins of S6. S7 with anterior apodeme

rather short in 5. (P.) sp. 17, long in other species; flat or slightly to markedly keeled posteromedially,

average length to elongate, in S. (P.) sp. 1 elongate and posteromedially pointed and keeled. T7 absent or

present as two tiny to small lateral sclerites; spiracles of segment 7 in membrane between segments 6 and 7

in most species, in membrane dorsal to posterior margin of S6 in S. (P.) sp. 17. S8 distinctly developed,

almost bare in S. (P.) sp. 3, with thick setae in S. (P.) sp. 17, average setation in other species. ri0 present as

median sclerite or paired sclerites.

Examined female genitalia of: S. (P.) spp. 1-4,14-17.

Taxonomic changes

Lectotype designation for Siphona brevirostris Coquillett. — Siphona

( Pseudosiphona ) brevirostris was described in Siphona s.s. by Coquillett in his

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O’Hara

“Revision of the Tachinidae” (1897: 76). In that work Coquillett routinely published

USNM type numbers for his new species and appropriately labelled his holotypes in

the collection (see general discussion of Coquillett’s type designations near the

beginning of the Classification chapter). Coquillett’s type series of S. brevirostris

consists of three specimens (two females and one male - not three females as

published), two of which (both females) were attached to the same pin. The pin with

the male specimen bears a paratype label and the other pin, with the two females,

bears a type label and a handprinted Townsend label: “Pseudosiphona brevirostris

Coq. Gnt. mss.” [Gnt. = Genotype, referring to Townsend’s selection of S.

brevirostris as type species of Pseudosiphona Tnsd.]. It is evident that both

Coquillett and Townsend intended one of the females on the double-mounted pin to

be the holotype of S. brevirostris , but it is not clear which one. It seems appropriate

to consider all three specimens as syntypes, and to select one of the females as

lectotype. Accordingly, the two female specimens have been removed to separate

pins and one selected, and here designated, lectotype of Siphona brevirostris

Coquillett. A red bordered label has been attached to the pin bearing the lectotype,

and reads: “LECTOTYPE/ Siphona/ brevirostris Coq./ O’Hara designation” (the

diagonal slashes indicate separate lines on label). The lectotype also bears the

following labels: (1) From Miss/ Murtfeldt, (2) 6, (3) Type/ No. 3574/ U.S.N.M.

[red USNM label], (4) Siphona/ brevirostris/ Coq. [Coquillett’s handprinted label],

(5) Pseudosiphona/ brevirostris Coq./ Gnt. mss. [Townsend’s handprinted label].

The lectotype is pinned with a minuten dorso-ventrally through the thorax, and is in

good condition except for loss of the right mid leg and tarsi of the left mid and hind

legs. The remaining two specimens of the type series, a male and female, are

labelled as paralectotypes.

Hosts

Two new host records are known for S. (Pseudosiphona), based on label data

accompanying adult specimens in the USNM: one from caterpillars of a lycaenid

butterfly (collected in Costa Rica), and the other from larvae of Chloropteryx sp.

(collected in Trinidad) (Table 2).

Amaud (1978) records Siphona brevirostris from Oidaematophorus

homodactylus (Table 2), citing the work of Schaffner (1959). This record is

probably in error because specimens of S. brevirostris are extremely scarce in

collections, and their proper identification is very difficult (so few identified

specimens are known that comparison with type specimens is almost essential for

reliable determination - see discussion concerning the geographic distribution of S.

brevirostris below).

Phylogenetics

Most S. (Pseudosiphona) species have a characteristic habitus (see Recognition

section), but some species deviate from it and a few other Siphona s.l. species

Systematics of the Genus Group Taxa of the Siphonini

107

approximate it, so no external synapotypies of S. ( Pseudosiphona ) are proposed

here. Other character states, in the male genitalia and first instar, provide better

evidence for monophyly of this group.

Two structures of the male genitalia, the pregonite and distiphallus, have states

unique to S. ( Pseudosiphona ) species. Three characters are recognized on the

pregonite: shape, presence or absence of spinules anterolaterally, and presence or

absence of a seta(e) posteriorly. Though spinules are on the pregonite of all

examined S. ( Pseudosiphona ) species, their presence in some other New World

Siphona s.l. species indicates this state is not unique to this subgenus. States for the

other two characters are unique to S. ( Pseudosiphona ) and are interpreted as

synapotypies. The first is the characteristically broadened shape of the pregonite,

and the second is the presence of more than one seta posteriorly. A few S.

(Pseudosiphona) species lack one or the other of these states, but not both. A

triangular and anterolaterally spined distiphallus (Fig. 128) is hypothesized as

another synapotypy of S. (Pseudosiphona) species. The few species which depart

from this shape are interpreted as derived for this character as they have the derived

states of the pregonite.

First instars of five of the seven examined S. (Pseudosiphona) species have an

elongate, hook-like, labrum (Fig. 162). This state is interpreted as apotypic because

other Siphona s.l. species have a hatchet-like labrum. Two species of S.

(Pseudosiphona) have a slightly broadened labrum (in dorsal-ventral plane), but this

state is considered secondarily (and independently) derived in these species for

reasons given in O’Hara (in prep.).

Geographic distribution

Siphona (Pseudosiphona) is primarily a Neotropical group, for which I am aware

of only two species north of Mexico: S. brevirostris and undescribed species S. (P.)

sp. 3. The known range of the latter is from California to east Texas, and southward

into northcentral Mexico. The range of S. brevirostris is not well documented

because of a paucity of collected specimens and a problem with the location of the

type locality. The type locality is cited by Coquillett (1897) as Kirkwood, Missouri,

but Sabrosky and Amaud (1965) consider the type locality to be Rhode Island

because one of the syntypes bears a label inscribed with “In box with micro-larvae

from R.I.”. Perhaps Sabrosky and Amaud are correct, as the only unquestionably

conspecific specimen of S. brevirostris that I examined was collected in 1983 near

Durham in Strafford Co., New Hampshire (CNC). A possibly conspecific specimen

(unforunately a female and hence less easily identified) was collected from

southwestern New Mexico (USNM). From such information it is apparent that S.

brevirostris is at least present in northeast USA, and may range southwestward to

Missouri or even New Mexico.

Of the other approximately 18 undescribed species of S. (Pseudosiphona) known

to me, all are Neotropical, with greatest diversity in Chiapas (Mexico), southeastern

Quaest. Ent., 1989, 25 (1,2)

108

O’Hara

Brazil, and southern Peru (this pattern is due at least in part to collecting bias).

Several species seem to have very wide ranges, extending from Middle America to

southeastern Brazil. Unworked material includes specimens from Colombia,

Venezuela, British Guiana, Bolivia, and the Caribbean islands of Trinidad and

Jamaica. Records from southeastern Brazil and northern Argentina mark the known

southern distributional limit of S. ( Pseudosiphona ) species.

List of described species included in Siphona C Pseudosiphona )

N brevirostris Coquillett, 1897: 76. Lectotype female (by designation in text),

USA: Missouri, Kirkwood (USNM). Lectotype examined.

List of examined, undescribed, species included in Siphona {Pseudosiphona)

S. (Pseudosiphona) sp. 1: Males and females from Chiapas, Mexico (CNC).

S. ( Pseudosiphona ) sp. 2: Males and females from SE Brazil (CAS, CNC, USP), males from Ecuador

(CNC), Peru (CNC) and El Salvador (CAS). One possibly conspecific male from Chiapas, Mexico (CNC).

S. (Pseudosiphona) sp. 3: Males and females from California, USA (AMNH, CAS, CNC, MCZ, UCB, UKL,

USNM), single males from New Mexico, USA (JEOH) and Durango, Mexico (CNC), single female from

Texas, USA (CNC).

S. (Pseudosiphona) sp. 4: Males and females ranging from Sonora to Chiapas, Mexico (CAS, CNC).

S. (Pseudosiphona) sp. 5: Males from SE Brazil (CAS, CNC, USP) and Chiapas, Mexico (CNC).

S. (Pseudosiphona) sp. 6: Two males from Belize (CNC).

S. (Pseudosiphona) sp. 7: One male from SE Brazil (CNC).

S. (Pseudosiphona) sp. 8: Males from Ecuador and Peru (CNC).

S. (Pseudosiphona) sp. 9: One male from Peru (CNC).

S. (Pseudosiphona) sp. 10: One male from Chiapas, Mexico (CNC).

S. (Pseudosiphona) sp. 11: One male from Chiapas, Mexico (CNC).

S. (Pseudosiphona) sp. 12: Three males from Peru (CNC).

S. (Pseudosiphona) sp. 13: One male from SE Brazil (CNC).

S. (Pseudosiphona) sp. 14: One female from Chiapas, Mexico (CNC).

S. (Pseudosiphona) sp. 15: Males and one female from northern Argentina (CNC).

S. ( Pseudosiphona ) sp. 16: One female from Chiapas, Mexico (CNC).

S. (Pseudosiphona) sp. 17: One male, three females from SE Brazil (AMNH, BMNH, USP).

S. (Pseudosiphona) sp. 18: Males and females from SE Brazil (CNC, USP), single males from Ecuador

(CNC), Costa Rica (USNM) and Nicaragua (USNM).

S. (Pseudosiphona) sp. 19: Three males from Amazonas, Brazil (INPA).

Siphona (subgenus Siphona Meigen sensu stricto)

Figs. 25-26, 28, 32, 34, 36, 43, 45, 67, 96, 129, 141, 144, 163.

Crocuta Meigen, 1800: 39/ Type-species, Musca geniculata De Geer, 1776 (by designation of Coquillett,

1910: 528). Suppressed by I.C.Z.N., 1963: 339 (Opinion 678).

Siphona Meigen, 1803: 281. Type-species, Musca geniculata De Geer, 1776, by designation of I.C.Z.N.,

1974: 157 (Opinion 1008).

Bucentes Latreille, 1809: 339. Type-species, B. cinereus Latreille, 1809 (monotypy) = Stomoxys minuta

Fabricius, 1805.

Phantasiosiphona Townsend, 1915: 93. Type-species P. tropica Townsend, 1915 (original designation).

Systematics of the Genus Group Taxa of the Siphonini

109

Recognition

Siphona geniculata is the type species of Siphona s.s., and its name literally

means “geniculate siphon”. The name refers to the long prementum and labella of

that species, but it could just as accurately have been applied to any Siphona s.s.

species, as all are characterized by a long proboscis, with the labella at least as long

as the elongate prementum. This state has generally been regarded as unique to

Siphona s.s. species, but is shared with some other siphonines.

Many non-Siphona s.s. species have slight to moderate elongation of the labella,

but only in a few are they as long as the prementum. These can be discerned as

non-Siphona s.s. species by the characteristics they share with other siphonine

lineages. The several Actia species of this type are easily recognized by their row of

katepistemal hairs and the anal vein not extended to the wing margin. The few

species of S. (Aphantorhaphopsis) with the labella almost as long as, to slightly

longer than, the prementum have fully flexible labella (not inflexible on basal half in

life as in Siphona s.s. species) and/or a seta posteriorly on the pregonite (absent from

Siphona s.s. species). S. (Aphantorhaphopsis) Nepal sp. 1 has a very long proboscis

of the typical Siphona s.s. appearance and is the most easily misplaced species of

this taxon, but can be correctly placed by its seta on the pregonite. The proboscis of

S. (Pseudosiphona) sp. 15 is also S. (Siphona)- like, and several other Neotropical

species of that subgenus approach this state, but these species are readily recognized

by characters in the male genitalia (see Recognition section of S. (Pseudosiphona)).

The few species of Neotropical Siphona s.l. species in which the labella approach

the Siphona s.s. condition are recognized by the presence on the pregonite of a seta

posteriorly and/or spinules apically.

Description

Length: 2.5-6.5mm (rarely less than 3.0mm).

Head (Figs. 25-26; ten Palearctic species shown in Mesnil 1965, figs. 150-159; North American species

shown in O’Hara 1983a, figs. 1, 18-59). — Proclinate orbital setae weak to strong, subequal in length or

anterior orbital longer. Eye of male small to very large, 0.62-0.91 head height; eye of female smaller than in

male. Flagellomere 1 of male markedly varied in length, 0.40-0.75 head height; linear to large and almost

triangular. Flagellomere 1 of female smaller than in male. Aristomere 1 short in most species, 1-2X longer

than wide in some species with long flagellomere 1, 3X longer than wide in S. trichaeta. Aristomere 2 varied

from 2.0-10.0X longer than wide, (most species between 2.5-6.0). Aristomere 3 almost bare to

micropubescent, long and evenly tapered in most species, short and thickened to near tip in a few. Clypeus

narrow and enclosed in membrane. Palpus long, clavate. Prementum and labella each slender and elongate;

latter at least as long as prementum, inflexible on basal half or more, with pseudotracheae concentrated

apically and reduced in number. Proboscis length (prementum + labella) varied from 1.3-4.8X head height

(most species between 2.0-3.0).

Thorax (Figs. 28, 32, 34, 36). — Prostemum setulose or bare, varied within some species. Lower

katepistemal seta longer than upper anterior seta in most species (Fig. 32), subequal in length to it in a very

few. Three or four postsutural dorsocentral setae. Upper part of anepistemum with single setula in most

species (Fig. 32), with two in a few. Fore tibia with preapical ad seta much shorter than d seta. Mid tibia

with one ad seta. Tarsomeres normal in size in most species, tarsomere 5 of fore leg slightly to moderately

broadened in females of a very few; claws short in most species, medium to large in a few. Wing vein CuAx

with distal portion 0.26-0.67 length of proximal portion ( i.e . dm-cu near wing margin; mean 0.41). Wing

setulae: in most species Rx dorsally bare or with one or two setulae distally, in a very few entirely setulose;

Quaest. Ent., 1989, 25 (1,2)

110

O’Hara

ventrally bare (most species) or distally setulose (a very few); ^4+5 setulose between base and r-m in most

species, beyond r-m in a few; CuA{ bare in almost all species, with setulae in a very few ( e.g . S. lutea and S.

bevisi).

Abdominal terga 1-5. — Abdomen ovoid in shape in most species, markedly elongate in several

Afrotropical species. Tl+2 with or without median marginal setae (present primarily in members of S.

geniculata group - see O’Hara 1983a: 323-4); one pair (rarely two pair) strong lateral marginal setae in most

species, weak or absent in a few (weak or absent primarily in S. maculata group - see O’Hara 1983a:

321-322). TyT5 average in most species, with extra pair of lateral marginal setae on T3 in a few species.

Male genitalia (Figs. 67, 96, 129, 141, 144). — S5 little varied (Fig. 67; also Andersen 1982, figs. 3-5),

posterior margins of processes approximately U-shaped in most species, obtusely angled in a few; apical

lobe distinctly differentiated, in a very few species apex curved inward (known only in S. cuthbertsoni, S.

mesnili (Andersen 1982, fig. 4) and S. oligomyia (O’Hara 1983a, fig. 11)); median lobe rounded, relatively

unmodified; processes sparsely to moderately setulose. T6 thin to broad and dorsally continuous in most

species, almost absent from a few (apparently not reduced to two lateral sclerites as in some other

siphonines). Ejaculatory apodeme with fan-shaped portion 1.0-2.0X wider than hypandrial apodeme.

Pregonite (Fig. 96) in profile little varied, curved anteriorly and more or less pointed apically; bare.

Epiphallus absent. Distiphallus (Fig. 129) reduced posteriorly, spined anterolaterally in a few species, in

profile apically rounded, truncate or pointed posteriorly; laterally incised in a very few species (e.g.

Philippines sp. 1). Postgonite apically rounded. Surstylus (Fig. 141, 144) average length to slightly elongate,

more or less straight; basally free from epandrium. Cerci (Fig. 141, 144) average length, smoothly curved at

midlength; moderately setose on basal half.

Examined male genitalia of: S. akidnomyia, S. brunnea, S. collini, S. confusa, S. cristata, S. cuthbertsoni, S.

flavifrons, S. futilis, S. geniculata, S. illinoiensis, S. ingerae, S. intrudens, S. kairiensis, S. longissima, S.

lurida, S. lutea, S. macronyx, S. maculata, S. medialis, S. multifaria, S. nigricans, S. oligomyia, S. pacifica,

S. paludosa, S. pauciseta, S. pisinnia, S. rizaba, S. setosa, S. tropica, S. unispina, Ecuador spp. 1-3, Kenya

spp. 1-2, Nepal sp. 1, Philippines sp. 1 and Taiwan sp. 1. Examined published figures of: S. hungarica

(Andersen 1984, figs. 1-3), S. kairiensis (O’Hara 1983b, figs. 3-5), North American Siphona species

(O’Hara 1983a, figs. 60-80), and European Siphona species (aedeagus only, Andersen 1982, figs. 2, 18-28,

and Andersen 1983, fig. 21). Male reproductive system of S. cristata shown in Andersen (1983, fig. 1).

Female genitalia (Figs. 43, 45). — Short. S6 with average length hairs. T6 absent; spiracles of segment 6

in membrane dorsal to anterolateral margins of S6. S7 with long anterior apodeme; not posteromedially

keeled. Tn absent; spiracles of segment 7 in membrane dorsal to anterolateral margins of S7. S8 distinctly

developed, haired. T10 absent or very slightly sclerotized.

Female genitalia of S. melanura (examined externally but not dissected) are uniquely modified, in

marked contrast to the conservative features in other members of the subgenus. S6 is longer than wide,

shovel-like, extremely smooth and shiny black, apically rounded, with hairs confined to single row on

membrane along posterior edge of sternum. S6 in form of flap-like covering over similarly-shaped, apically

more sharply-rounded, blade-like S7.

Examined female genitalia of: S. nr. intrudens, S. lutea, S. maculata, S. nigricans, S. pisinnia and S. nr.

tropica. Examined published figures of: S. hungarica (Andersen 1983, fig. 4), S. nigricans (fig. 12 in O’Hara

1983a, as S. hokkaidensis ) and S. setosa (Andersen 1982, fig. 6, and Andersen 1983, fig. 7). Female

reproductive system of S. ingerae shown in Andersen (1983, fig. 2).

Taxonomic changes

Siphona nigrohalterata Mesnil. — Among the new siphonines described by

Mesnil in his 1959 paper about tachinids of East Africa were Siphona amplicornis

amplicornis and Siphona amplicornis nigrohalterata (pp. 21-22). The types (both

male) were collected from West Kibo, Nigeria - the former at 2800m between

17-22.IV. 1952 and the latter at 3500m between 23-30.IV. 1952. These “subspecies”

do not seem to be geographically or temporally isolated, as a paratype of S. a.

amplicornis bears the same locality data as the type of S. a. nigrohalterata. I infer

from this that these sympatric “subspecies” either freely interbreed (i.e. cannot be

Systematics of the Genus Group Taxa of the Siphonini

111

regarded as separate subspecies) or are reproductively isolated ( i.e . represent two

species). Examination and comparison of the holotypes suggest the latter; the type

of S. a. nigrohalterata has a slightly longer proboscis, less elongate abdomen,

shorter abdominal setae, and markedly different coloration (particularly on the

abdomen). These differences are not major, but in total are not atypical of closely

related Siphona species. Thus I conclude that S. nigrohalterata is probably a valid

species reproductively isolated from S. amplicornis.

Synonymy of Siphona janssensi (Mesnil) with Siphona cuthbertsoni Curran. — S.

cuthbertsoni is more easily recognized than most African Siphona species because

of its very large eye, rather short flagellomere 1, proboscis length less than 2.5X

head height, and generally dark color (including abdomen entirely dark in ground

color). It is probable that Mesnil was unfamiliar with this species of Curran’s,

except perhaps by name, as he did not mention it in his work about the Siphona of

the “Belgian Congo”.

Specimens of S. cuthbertsoni identified by direct comparison with the male

holotype of that species were later compared with the male holotype of S. janssensi.

On the basis of these comparisons, the name S. janssensi is here synonymized with

that of S. cuthbertsoni.

Note about use of the subspecific category . — Subspecies have rarely been

described in the Siphonini, though Mesnil occasionally used this category for taxa

displaying slight morphological differences. Mesnil’ s concept of subspecies was not

based on geographic (and hence I assume not reproductive) isolation. I examined the

types of most of Mesnil’ s subspecies and have reclassified their names as synonyms

or separate species. However, I was unable to examine the types of several

subspecies of Siphona fuliginea Mesnil and Siphona reducta (Mesnil). I leave these

taxa as classified by Mesnil pending study of the types.

Status of Siphona unispina ( Mesnil ) and Siphona unispina infuscata (Mesnil). —

In 1952 Mesnil revised the Siphona species of the “Belgian Congo”, naming many

new species and one pair of subspecies: S. unispina unispina and S. unispina

infuscata (Mesnil 1952a). As was Mesnil’ s practice, these subspecies were

recognized exclusively on morphological criteria without the now standard

requirement of at least partial spatial or temporal separation. Type localities of these

taxa are separated by less than 150km, with paratypes of both collected at one

intermediate locality. This suggests that the subspecific category is inappropriate,

and that the names S. u. infuscata and S. unispina are synonyms or S. u. infuscata is

specifically distinct. Crosskey (1980) chose the former, but upon comparison of the

types of both taxa I have decided to recognize Siphona infuscata as a valid species,

based upon three characteristics that differ between the types: 1) proboscis length of

S. unispina is 3.5X head height as compared with 4.3X head height in S. infuscata,

2) thorax of S. unispina is bluish gray, in S. infuscata mostly brown, and 3) R4+5 is

setulose beyond r-m in S. unispina and not beyond r-m in S. infuscata. Abdominal

characteristics and especially male genitalia cannot be compared between types

Quaest. Ent., 1989, 25 (1,2)

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O’Hara

because the abdomen of the type of S. unispina is missing. Paratypes were not

examined, but a male specimen from the Bwamba river area (SE Zaire) identified by

Mesnil as S. unispina fits the description of that species and possesses the three

characteristics listed above for distinguishing S. unispina from S. infuscata.

Hosts

Most hosts of Siphona s.s. are species of the Noctuidae, and all known

lepidopteran hosts belong to the Macrolepidoptera (Table 2). Some members of the

S. geniculata group are parasites of crane fly larvae (Tipulidae), and are unique in

this habit in the Siphonini and among the few tachinids that parasitize other Diptera.

Phylogenetics

The long, geniculate proboscis of Siphona s.s. species, with labella at least as

long as the elongate prementum, is synapotypic of the subgenus. Elongation of the

proboscis has occurred independently in many siphonine lineages, but only in

Siphona s.s. species are the labella rigid (in life) on basal half or more and at least as

long as the prementum in all species. Hence this condition is considered the

groundplan (and synapotypic) state of Siphona s.s., and a shorter (with a few

exceptions padlike) labella the groundplan state of other siphonine lineages.

Andersen (1983) also considered the reduced number of pseudotracheae in

Siphona s.s. species as a synapotypy of the subgenus, but this state is not treated

separately here because for many species it is (in siphonines) directly correlated

with elongation of the labella. For example, padlike labella may have about 15 pair

of pseudotracheae, elongate labella of S. (Siphonopsis) species (Fig. 18) about eight

pair, and the long labella of Siphona s.s. species about four pair. This correlation is

not constant as certain species of S. (Aphantorhaphopsis), for instance, with

elongate labella have relatively numerous pseudotracheae. The trend is nevertheless

evident, and because of this the number of pseudotracheae on the labella is not

treated as a separate character in this study. No other synapotypies of Siphona s.s.

species are known. The included species have a number of shared characteristics, of

which male genitalic similarities are particularly marked (see figs. 60-80 in O’Hara

1983a), and these are congruent with the hypothesis of monophyly of this group.

The few other siphonines with a S. (Siphona)- like proboscis are recognized as

non-members of that group by synapotypies they share with other taxa, and their

non-S. (Siphona)- like male genitalia.

Andersen’s (1982) revision of European Siphona s.s. species went to press at

about the time I completed a revision of the North American Siphona s.s. species

(O’Hara 1983a). Therefore my comparison of Nearctic and Palearctic species of

Siphona s.s., and discussion of species groups, did not include the four new

European Siphona s.s. species described by him. Here I briefly mention the species

groups to which Andersen’s species belong: S. ingerae is a member of the S.

maculata group, and is closely related to the North American species S. intrudens; S.

Systematics of the Genus Group Taxa of the Siphonini

113

martini is now considered conspecific with S. nigricans (Andersen, pers. comm.),

and is a member of the S. geniculata group; and S. mesnili and S. variata are placed

in the S. geniculata group (formerly called the S. cristata group), because these

species were considered by Andersen {op. cit .) to be close to S. confusa. Andersen’s

new species from Hungary, S. hungarica (Andersen 1984), is closely related to S.

ingerae and S. intrudens , and belongs to the S. maculata group.

Geographic distribution

Siphona s.s. is widely distributed, with 84 described species. Forty-one of these

are Afrotropical in distribution (see Crosskey 1980 for ranges; partial key in Mesnil

1952a), with more species recorded from Madagascar than any other siphonine

taxon (Mesnil 1977a). Three apparently endemic species are on Madagascar and

five species are shared with the mainland (Crosskey 1980), suggesting that Siphona

s.s. includes some markedly vagile members. This suggestion is further evidenced

by the presence of Siphona s.s. species on the Juan Fernandez Islands west of Chile,

the Canary Islands, and four species with Holarctic distributions (excluding S.

geniculata , introduced by man to North America). It is therefore rather surprising

that Siphona s.s., with 41 species in the Afrotropical region and 20 in the Palearctic

region, has only three described species in the Oriental region (Crosskey 1976a) and

one in the Australian region (O’Hara 1983b). From examined material (and Shima,

pers. comm.), the Oriental region and probably islands of the Indoaustralian

archipelago north of Australia contain at least a modest number of undescribed

species. There is as yet no indication of more than one Siphona s.s. species in

Australia.

Palearctic Siphona s.s. species are best known in England (Crosskey 1976b) and

Europe, and are keyed in Mesnil (1964) and Andersen (1982). Additional records

are cited in Mesnil and Pschom-Walcher (1968), Herting (1968b, 1969b, 1973),

Draber-Mohko (1978, 1981), Kugler (1979), Richter (1971, 1975, 1976a,b, 1980,

1986), Richter and Khitsova (1982), Chao and Shi (1982), Karczewski (1983),

Mihalyi and Weinberg (1984) and Rognes (1986), and distributions are summarized

in Herting (1984).

Twenty-one Siphona s.s. species are in North America (including “Middle

America”). These were recently revised and their distributions discussed in O’Hara

(1983a). Only two exclusively South American Siphona s.s. species are described,

but there are specimens of at least ten undescribed species among material I have

examined.

List of described species included in Siphona {Siphona)

A abbreviata (Villeneuve), 1915: 199 {Bucentes). Female syntypes, Madagascar:

Sikora (NMV). Syntypes examined.

N,S akidnomyia O’Hara, 1983a: 311. Holotype male, Mexico: Chiapas, Mt.

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Zontehuitz (CNC). Holotype examined.

A albocincta (Villeneuve), 1942a: 55 ( Bucentes ). Holotype female, Zaire:

Nylragongo (CNC). Holotype examined.

A amoena (Mesnil), 1952a: 12 ( Crocuta ). Holotype male, Zaire: Karisimbi

(MRAC). Holotype examined.

A amplicornis Mesnil, 1959: 21. Holotype male, Tanzania: West Kibo (SMNS).

Holotype examined.

A angusta Mesnil, 1959: 22. Holotype male, Tanzania: West Kibo (SMNS).

Holotype examined.

A antennalis (Mesnil), 1952a: 9 (Crocuta). Holotype male, Zimbabwe: Salisbury

(CNC).

A atricapilla Mesnil, 1959: 20. Holotype male, Tanzania: West Kibo (SMNS).

Holotype examined.

A bevisi Curran, 1941: 7. Holotype male, South Africa: Natal, Durban (AMNH).

Holotype examined.

A bilineata (Mesnil), 1952a: 10 (Crocuta). Holotype male, Rwanda: Niabirehe

(MRAC). Holotype examined.

P boreata Mesnil, 1960: 190. Holotype male. Federal Republic of Germany:

Amsberg (CNC). Holotype examined.

S brunnea O’Hara, 1983a: 308. Holotype male, Mexico: Chiapas, Mt. Zontehuitz

(CNC). Holotype examined.

A capensis Curran, 1941: 7. Holotype female, South Africa: East London (PPRI).

Holotype examined.

P collini Mesnil, 1960: 188. Holotype male. Great Britain: Cambridgeshire,

Burwell (HDE). Holotype examined.

P confusa Mesnil, 1961: 201. Holotype male, Sweden: Granna (CNC). Holotype

examined.

A cothurnata (Mesnil), 1952a: 17 (Crocuta). Holotype male, Rwanda: Burambi

(MRAC). Holotype examined.

A creberrima (Speiser), 1910: 142 (Crocuta). Syntypes, Tanzania: Kilimanjaro

(NRS). Syntypes examined.

P,N cristata (Fabricius), 1805: 281 (Stomoxys). Holotype female, Denmark: Zealand

(ZMUC). Holotype examined.

syn. palpina Zetterstedt, 1859: 6064. Holotype female, Sweden:

Lindholmen (UZI). — Mesnil, 1964: 861.

chetoliga Rondani, 1865: 194. Holotype female, Italy (MZF). —

Herting, 1969a: 192.

chaetolyga. Incorrect subsequent spelling of chetoliga Rondani

(Herting, 1969a: 192).

A cuthbertsoni Curran, 1941: 7. Holotype male, Zimbabwe: Salisbury (AMNH).

Holotype examined.

syn. janssensi (Mesnil), 1952a: 4 (Crocuta). Holotype male (abdomen

Systematics of the Genus Group Taxa of the Siphonini

115

missing), Rwanda: Kibga (MRAC). Holotype examined. New

synonymy.

P ejflatouni Mesnil, 1960: 188. Holotype female, Egypt: Mariout (CNC).

Holotype examined.

P flavifrons Staeger in Zetterstedt, 1849: 3211. Lectotype female (by designation

of Andersen, 1982: 167), Denmark: North East Zealand, Ordrup (ZMUC).

Lectotype examined.

N floridensis O’Hara, 1983a: 288. Holotype male, USA: Florida, Orlando

(AMNH). Holotype examined.

O foliacea (Mesnil), 1953: 113 ( Crocuta ( Siphona )). Holotype male, Burma:

Kambaiti (ZMU). Holotype examined.

A fuliginea Mesnil, 1977a: 77. Holotype male, Madagascar: Ambatolahy

(MNHN).

ssp. cerina Mesnil, 1977a: 76. Holotype male, Madagascar: Amber Mtn.

(MNHN).

rubea Mesnil, 1977a: 77. Holotype male, Madagascar:

Manjakatompo (MNHN).

N,S futilis Wulp, 1890: 125. Lectotype male (by designation of O’Hara, 1983a:

305), Mexico: Guerrero, Omilteme (BMNH). Lectotype examined,

syn. ceres (Curran), 1932: 14 ( Bucentes ). Holotype female, Guatemala:

Antigua (AMNH). — O’Hara, 1983a: 305. Holotype examined.

O gedeana Wulp, 1896: 109. Holotype female, Indonesia: Java, Goenoeng Gedeh

(lost, see Crosskey, 1976a: 214).

syn. nigripalpis (de Meijere), 1924: 223 {Bucentes). Lectotype male (by

designation of Crosskey, 1969: 89), Indonesia: Java, Pangrango

(ZMA). Lectotype examined.

P,N geniculata (De Geer), 1776: 38 ( Musca ).2 Three syntypes, Sweden (NRS;

types erroneously listed as lost in Andersen 1982 and O’Hara 1983a).

syn. urbanis (Harris), 1780: 153 {Musca). Type, England (lost). —

Crosskey, 1976b: 100.

minuta (Fabricius), 1805: 282 {Stomoxys). Holotype female (head

missing), Denmark (ZMUC). — Herting, 1984: 125. Holotype

examined.

cinerea (Latreille), 1809: 338 {Bucentes). Type(s), France: Paris (not

located). — Herting, 1984: 125.

cinerea Meigen, 1824: 156. Holotype female, Europe (MNHN). —

Herting, 1972: 4.

tachinaria Meigen, 1824: 156. Holotype male, Federal Republic of

Germany: Nordrhein, Aachen (MNHN). — Herting, 1972: 13.

Page 20 is cited by various authors for the description of S. geniculata , but this apparently refers to the

German translation of De Geer’s work by Goeze in 1782 (Herting, in lift.).

Quaest. Ent., 1989, 25 (1,2)

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O’Hara

analis Meigen, 1824: 157. Holotype male, Europe (MNHN). —

Herting, 1972: 3.

nigrovittata Meigen, 1824: 157. Holotype male (MNHN). — Herting,

1972: 11.

meigenii (Lepeletier and Serville in Latreille, et al.), 1828: 501

(. Bucentes ). Replacement name for S. cinerea Meigen, 1824

(objective synonym). — Herting, 1984: 125.

A gracilis (Mesnil), 1952a: 13 ( Crocuta ). Holotype male (head missing), Rwanda:

Kibga (MRAC). Holotype examined.

P grandistylum Pandelle, 1894: 108. Holotype male, France: Pyrenees (MNHN).

P griseola Mesnil, 1970: 118 (as subspecies of 5. maculata Staeger). Holotype

male, Israel: Tel Aviv (CNC). Holotype examined.

P hungarica Andersen, 1984: 5. Holotype male, Hungary: Hortobagy N.P.

(HNHM). Paratype examined.

N illinoiensis Townsend, 1891: 368. Lectotype male (by designation of O’Hara,

1983a: 307), USA: Illinois (UKL). Lectotype examined.

A infuscata (Mesnil), 1952a: 14 (as subspecies of Crocuta unispina Mesnil).

Holotype male, Zaire: Tshamugussa (MRAC). Holotype examined. New status.

P ingerae Andersen, 1982: 161. Holotype male, Denmark: North East Zealand,

Bagsvaerd, Sm^rmose (ZMUC). Holotype examined.

N intrudens (Curran), 1932: 14 (Bucentes). Holotype male, USA: Pennsylvania,

Castle Rock (AMNH). Holotype examined.

U kairiensis O’Hara, 1983b: 79. Holotype male, Australia: Queensland, Tinaroo

Lake (CAS). Holotype examined.

S kuscheli (Cortes), 1952: 110 (Phantasiosiphona). Holotype male, Juan

Fernandez Islands: Masatierra (CIE). Holotype examined.

A laticornis Curran, 1941: 9. Holotype male, South Africa: Pretoria (PPRI).

Holotype examined.

A lindneri Mesnil, 1959: 22. Holotype male, Tanzania: Msingi (SMNS). Holotype

examined.

N,S longissima O’Hara, 1983a: 311. Holotype male, Mexico: Chiapas, San

Cristobal (CNC). Holotype examined.

N,P lurida Reinhard, 1943: 20. Holotype male, USA: Oregon, Rainier (CNC).

Holotype examined.

N lutea (Townsend), 1919: 584 (Crocuta). Lectotype female (by designation of

O’Hara, 1983a: 296), USA: New Hampshire, Franconia (USNM). Lectotype

examined.

syn. tenuis Curran, 1933b: 10. Holotype male, Canada: Ontario,

Timagami (AMNH). — O’Hara, 1983a: 296. Holotype examined.

N macronyx O’Hara, 1983a: 313. Holotype male, USA: Washington, Pullman

(WSUP). Holotype examined.

P,N maculata Staeger in Zetterstedt, 1849: 3212. Lectotype male (by designation of

Systematics of the Genus Group Taxa of the Siphonini

117

Andersen, 1982: 162), Denmark: North East Zealand, Charlottenlund (ZMUC).

Lectotype examined.

N medialis O’Hara, 1983a: 303. Holotype male, Canada: Nova Scotia, Cranberry

Island (CNC). Holotype examined.

A melania (Bezzi), 1908: 58 (. Bucentes ). Holotype female, Ethiopia (not located).

A melanura Mesnil, 1959: 23. Holotype female, Tanzania: West Kibo (SMNS).

Holotype examined.

P mesnili Andersen, 1982: 163. Holotype male, Denmark: North East Zealand,

Gribskov, Harager Hegn (ZMUC). Holotype examined.

N multifaria O’Hara, 1983a: 293. Holotype male, Canada: Yukon, Dempster

Hwy., mi. 87 (CNC). Holotype examined.

A munroi Curran, 1941: 6. Holotype female. South Africa: Fort Jackson (PPRI).

Holotype examined.

A murina (Mesnil), 1952a: 15 ( Crocuta ). Holotype male, Zaire: Nyongera

(MRAC). Holotype examined.

P,N nigricans (Villeneuve), 1930: 100 {Bucentes). Holotype male, Sweden:

Gallivara (CNC). Holotype examined.

syn. hokkaidensis Mesnil, 1957: 36. Holotype female, Japan: Hokkaido,

Obihiro (CNC). — Herting, 1982: 8. Holotype examined.

silvarum Herting, 1967a: 9. Holotype male, Federal Republic of

Germany: Freiburg (CNC). — Herting, 1982: 8. Holotype examined.

martini Andersen, 1982: 169. Holotype male, Sweden: Scania,

Hyllstofta (UZI). — Andersen (pers. comm.). Paratype examined.

A nigrohalterata Mesnil, 1959: 22 (as ssp. of S. amplicornis Mesnil). Holotype

male, Tanzania: West Kibo (SMNS). Holotype examined. New status.

A nigroseta Curran, 1941: 8. Holotype female, South Africa: Pretoria (PPRI).

Holotype examined.

O nobilis (Mesnil), 1953: 112 ( Crocuta ( Siphona )). Holotype male, Philippines:

Mont Palis (ZMU). Holotype examined.

A obesa (Mesnil), 1952a: 8 (Crocuta). Holotype male, Zaire: Rwindi (MRAC).

Holotype examined.

A obscuripennis Curran, 1941: 8. Holotype female, Zimbabwe: Vumba Mts.

(AMNH). Holotype examined.

N oligomyia O’Hara, 1983a: 297. Holotype male, Canada: British Columbia,

Keremeos (CNC). Holotype examined.

N pacifica O’Hara, 1983a: 291. Holotype male, USA: Washington, Dartford

(WSUP). Holotype examined.

P paludosa Mesnil, 1960: 188. Holotype male, USSR: Tolmatschevo, Luga

(ZIL). Holotype examined.

A patellipalpis (Mesnil), 1952a: 10 (Crocuta). Holotype male, Zaire: Mt. Sesero

(MRAC). Holotype examined.

P pauciseta Rondani, 1865: 193. Eight syntypes, Italy (MZF; mixed series

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O’Hara

according to Herting, 1969a: 198).

syn. oculata Pandelle, 1894: 108. — Lectotype male (by designation of

Herting, 1978: 6), France (MNHN).

delicatula Mesnil, 1960: 190. Holotype male. Great Britain:

Chippenham (HDE). — Herting, 1969a: 198. Holotype examined.

A phantasma (Mesnil), 1952a: 7 ( Crocuta ). Holotype male, Rwanda: Gahinga

(MRAC). Holotype examined.

A pigra Mesnil, 1977a: 78. Holotype female, Madagascar: Moramanga (MNHN).

N,S pisinnia O’Hara, 1983a: 298. Holotype male, USA: New Mexico, 21km. n.

Silver City (CNC). Holotype examined.

S pseudomaculata Blanchard, 1963: 233. Syntypes, Argentina: Santa Fe (MBR).

Syntypes examined.

A reducta (Mesnil), 1952a: 18 {Crocuta). Holotype male, Zaire: Tshamugussa

(MRAC). Holotype examined.

ssp. ludicra Mesnil, 1977a: 78. Holotype male, Madagascar:

Manjakatompo (MNHN).

N,S rizaba O’Hara, 1983a: 310. Holotype male, Mexico: Veracruz, Orizaba (MSU).

Holotype examined.

P rossica Mesnil, 1961: 202. Holotype male, USSR: Leningrad, Lugsk (ZIL).

Holotype examined.

A rubrapex Mesnil, 1977a: 79. Holotype female, Madagascar: Perinet (MNHN).

A rubrica (Mesnil), 1952a: 11 {Crocuta). Holotype male, Zaire: Rutshuru

(MRAC). Holotype examined.

A setinerva (Mesnil), 1952a: 16 {Crocuta). Holotype male, Rwanda: Kibga

(MRAC). Holotype examined.

P setosa Mesnil, 1960: 191. Holotype male, USSR: Tolmatschevo, Luga (ZIL).

Holotype examined.

P seyrigi Mesnil, 1960: 189. Holotype male, Canary Islands: Tenerife (MNHN).

Holotype examined.

A simulans (Mesnil), 1952a: 18 {Crocuta). Holotype male, Rwanda:

Kundhuru-ya-Tshuve (MRAC). Holotype examined.

A sola Mesnil, 1959: 21. Holotype male, Tanzania: Usangi (SMNS). Holotype

examined.

A spinulosa (Mesnil), 1952a: 12 {Crocuta). Holotype male, Zaire: Ngesho

(MRAC). Holotype examined.

A trichaeta (Mesnil), 1952a: 18 {Crocuta). Holotype male, Rwanda: Karisimbi

(MRAC). Holotype examined.

N,S tropica (Townsend), 1915: 93 {Phantasiosiphona). Holotype male, Mexico:

Veracruz, San Rafael (USNM). Holotype examined.

A unispina (Mesnil), 1952a: 14 {Crocuta). Holotype male (abdomen missing),

Zaire: Rutshuru (MRAC). Holotype examined.

P variata Andersen, 1982: 164. Holotype male, Denmark: Frederikshavn

Systematics of the Genus Group Taxa of the Siphonini

119

(ZMUC). Holotype examined.

A vittata Curran, 1941: 8. Holotype male, Zimbabwe: Salisbury (AMNH).

Holotype examined.

A vixen Curran, 1941: 9. Holotype female, Zimbabwe: Salisbury (AMNH).

Holotype examined.

A wittei (Mesnil), 1952a: 5 (Crocuta). Holotype male, Rwanda: Karisimbi

(MRAC). Holotype examined.

Nomina dubia

P analis Robineau-Desvoidy, 1830: 92. Type(s), France (lost).

P clausa Robineau-Desvoidy, 1850: 209. Holotype male, France (lost).

P consimilis Robineau-Desvoidy, 1850: 205. Holotype male, France (lost).

P fuscicornis Robineau-Desvoidy, 1850: 205. Syntypes, France (lost).

P humeralis Robineau-Desvoidy, 1850: 207. Syntypes, France (lost).

P maculipennis Meigen, 1830: 365. Holotype, Portugal: Algarve (lost).

P melanocera Robineau-Desvoidy, 1850: 206. Holotype female, France (lost).

P pusilla Robineau-Desvoidy, 1830: 92. Type(s), France: Paris (lost).

syn. persilla. Incorrect subsequent spelling of pusilla Robineau-Desvoidy

(Coquillett, 1897: 76).

P quadrinotata Robineau-Desvoidy, 1850: 203. Holotype female, France (lost),

syn. quadricincta. Incorrect subsequent spelling of quadrinotata

Robineau-Desvoidy (Rondani, 1859: 10).

P silvatica Robineau-Desvoidy, 1850: 208. Holotype male, France (lost).

P testacea Robineau-Desvoidy, 1850: 207. Holotype male, France (lost).

P tristis Robineau-Desvoidy, 1850: 203. Holotype female, France (lost).

Nomina nuda

S anthomyformis Lynch Arribalzaga in Brauer, 1898: 505 (13).

P dorsalis Brauer and Bergenstamm, 1891: 410.

P pauciseta Mesnil, 1964: 856 (as ssp. of S. geniculata).

O taiwanica (Baranov in Hennig), 1941: 195 {Crocuta).

List of examined, undescribed, species included in Siphona {Siphona)

S. (Siphona) nr. intrudens : Females from Alberta, Canada (JEOH).

S. ( Siphona ) nr. tropica : One female from Coquimbo, Chile (CNC).

S. (Siphona) Ecuador sp. 1: One male from sw. Alausi, Chimborazo (CAS).

S. (Siphona) Ecuador sp. 2: One male from Cerro Pelado, Carchi (CNC).

S. (Siphona) Ecuador sp. 3: One male from sw. Banos (CAS).

S. (Siphona) Kenya sp. 1: One male from Mt. Kenya, Nanyuki (CAS).

S. (Siphona) Kenya sp. 2: Males and females from Kenya and Tanzania (CAS).

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S. (Siphona) Nepal sp. 1: One male from 28°00’N 85°00’E (CNC).

S. (Siphona) Philippines sp. 1: One male from Mt. Apo, Mindanao (BLKU).

S. (Siphona) Taiwan sp. 1: One male from Mt. Alishan (BLKU).

Many undescribed species of S. (Siphona) examined from South America.

Siphona (subgenus Siphonopsis Townsend)

Figs. 18, 24, 30, 97-98, 130.

Siphonopsis Townsend, 1916: 622. Type-species, Siphona plusiae Coquillett, 1895 (original designation).

New subgeneric status in Siphona Meigen.

Recognition

This homogeneous group of three described and approximately 13 undescribed

species is New World in distribution, with greatest diversity in the Neotropics.

The male genitalia of S. ( Siphonopsis ) species possess the only states which are

unique to this taxon, though the following external characteristics distinguish adults

of this taxon from those of most other New World Siphona s.l. species: abdomen

generally dark in ground color on T4 and T5 and medially on T3, with moderate

pruinosity (sparse pruinosity in a few species, and resembling abdomen of most S.

( Pseudosiphona ) species); (head of S. plusiae shown in Fig. 18) parafacial with

several hairs below lowest frontal seta (as in most Siphona s.l. species, but different

from the more haired parafacial of most S. (Pseudosiphona) species); arista almost

bare in most species (also in contrast to the haired arista of most S. ( Pseudosiphona )

species); labella of proboscis elongate, varied from two-thirds prementum length to

slightly less than prementum length; three postsutural dorsocentral setae; lower

katepistemal seta longer than upper anterior one; wing vein R4+5 not setulose beyond

crossvein r-m and CuAx bare.

The most import; nt diagnostic feature of S. ( Siphonopsis ) species externally is

labella length. As mentioned above, it varies from two-thirds prementum length to

slightly less than prementum length. This feature distinguishes members of this

taxon from those of Siphona s.s ., in which the labella are at least as long as the

prementum. Of several minor external differences between members of S.

( Siphonopsis ) and S. (Pseudosiphona) ( cf . Recognition sections), one of the better is

the slightly shorter labella of most (though not all) S. (Pseudosiphona) species.

Since several other Siphona s.l. species have labella as elongate as in S.

(Siphonopsis) species, it is necessary to examine the male genitalia of S.

( Siphonopsis )-\ike specimens for a reliable identification. It is not possible to

reliably identify adult females of all New World Siphona s.l. species.

Adult males with the above external characteristics are recognized as members

of S. (Siphonopsis) by the following genitalic states: sternum 5 with apex of apical

lobe curved inward; pregonite (Figs. 97-98) slender to average in width, lacking

spinules anterolaterally, with (most species) or without (a few species) tiny seta

posteriorly (medium-sized seta in a few species); distiphallus little varied, in profile

Systematics of the Genus Group Taxa of the Siphonini

121

tapered to narrow truncate tip in most species (Fig. 130), more elongate and lateral

apices separated in a few species; cerci not sharply inflexed at midpoint. The unique

shape of the distiphallus of S. ( Siphonopsis ) species readily separates males with the

above external characteristics from males of other Siphona s.l. lineages. Most likely

to be confused externally with S. ( Siphonopsis ) species are several species of S.

( Pseudosiphona ), but features of the pregonite, distiphallus and cerci unequivocally

separate males of these subgenera. Similarly, the few other Siphona s.l. species with

S. (Siphonopsis)- like labella (see Table 1) differ in male genitalic states (most

important of which is distiphallus shape).

Description

Length: 2.5-4.5mm.

Head (Figs. 18, 24, 30). — Proclinate orbital setae subequal in length or anterior one longer. Eye of male

medium to medium-large, 0.73-0.84 head height; eye of female subequal to slightly larger than in male.

Flagellomere 1 of male short to medium-long, 0.37-0.65 head height; shape from average width to large and

triangular. Flagellomere 1 of female smaller than in male in most species, subequal in a few. Aristomere 1

short in most species, as long as wide in at least two species. Aristomere two 1.5-6X longer than wide.

Aristomere 3 short to long and evenly tapered, almost bare to short plumose. Clypeus narrow and enclosed

in membrane. Palpus short, clavate. Proboscis (Figs. 18, 24, 30) with prementum short to medium in length,

between 0.6-0.7 head height in most species (not more than about 0.75 head height), labella elongate, 0.7-0.9

prementum length and inflexible in life on about basal third.

Thorax. — Prostem'um setulose. Lower katepistemal seta longer than upper anterior seta. Three

postsutural dorsocentral setae. Upper part of anepistemum with single setula. Fore tibia with preapical ad

seta much shorter than d seta. Mid tibia with one ad seta. Claws short. Wing vein CuAl with distal portion

0.29-0.55 length of proximal portion (mean 0.43). Wing setulae: R{ dorsally bare, or distally setulose,

ventrally bare; R4+5 setulose between base and r-m\ CuAx bare.

Abdominal terga 1-5. — Abdomen ovoid in shape. T]+2 without median marginal setae, lateral marginal

setae absent or weakly developed. T3-T5 with average setation.

Male genitalia (Figs. 97-98, 130). — S5 with posterior margins of processes approximately U-shaped;

apical lobe large and distinctly differentiated, apex curved inward; median lobe broadly rounded to truncate

and relatively unmodified in most species, with accessory lobe slightly to distinctly differentiated in a very

few species (distinctly differentiated only in S. conata)-, processes moderately setulose. T6 varied from two

lateral sclerites to single dorsally continuous sclerite. Ejaculatory apodeme with fan-shaped portion 1.0-2.0X

width of hypandrial apodeme. Pregonite (Figs. 97-98) in profile curved anteriorly and more or less pointed

apically, in a few species rather slender (Fig. 98); tiny to medium seta posteriorly in most species, seta

absent from at least two species. Epiphallus absent. Distiphallus (Fig. 130) reduced posteriorly, not incised

laterally, without enlarged spines anteriorly or laterally, in profile tapered to a narrow truncate tip, in a few

species distiphallus elongate and very narrow, with apex deeply incised anteriorly and posteriorly (i.e.

distiphallus with two long arms in posterior view). Postgonite reduced in S. conata, short to average length

in other species, apically rounded or pointed. Surstylus about average length in most species, elongate in 5.

conata , straight in most species, slightly curved posteriorly in a few; basally free from epandrium. Cerci

average length in most species, rather short in a few, in profile average width to thick; moderately setose on

basal half.

Examined male genitalia of: S. conata, S. plusiae, S. (Siphonopsis) spp. A-I,K-N.

Female genitalia. — Short. S6 with average length hairs. T6 absent, or present as two tiny to small lateral

sclerites or a single median sclerite; spiracles of segment 6 in membrane dorsal to lateral margins of S6 or in

lateral sclerites of T6. S7 with long anterior apodeme in most species, short in a few; flat and unmodified in

most species, in S. conata elongate and keeled posteromedially and apically incised. T7 absent in most

species, present as two tiny lateral sclerites in S. conata ; spiracles of segment 7 in membrane between

segments 6 and 7. S8 distinctly developed, haired. 7j0 present as median sclerite in most species, as two

sclerites in a very few.

Examined female genitalia of: S. conata, S. plusiae, S. ( Siphonopsis ) spp. A,B,E,M and N.

Quaest. Ent., 1989, 25 (1,2)

122

O’Hara

Hosts

S. conata and S. plusiae attack a serious pest of oak, the California oakworm

(Phryganidia californica). S. plusiae is also recorded from two other insect pests in

California, the cabbage looper ( Trichoplusia ni) and alfalfa looper (Autographa

californica ) (Table 2). Though attacking economically important pest species, S.

conata and S. plusiae do not parasitize their hosts at sufficient densities to be

considered good biological control agents.

S. brasiliensis is listed as a parasitoid of two noctuid species in South America

(Table 2). However, the name might be a misidentification because S. (Siphonopsis)

is very diverse in South America and S. brasiliensis is the only described species.

Phylogenetics

The distiphallus of males of S. ( Siphonopsis ) species is narrowed to a truncate tip

(Fig. 130). The uniqueness of this state within the Siphonini, and its uniformity

thoughout the subgenus, provides the best evidence for monophyly of this taxon. A

number of other similarities externally and in the male genitalia also suggest that the

included species are closely related; however, none can be defended as synapotypic

of S. ( Siphonopsis ). For instance, adults of S. ( Siphonopsis ) species have

characteristically elongate labella (Fig. 18), which are shorter (relative to

prementum length) than in Siphona s.s. species and longer than in most S.

( Pseudosiphona ) species. However, because this state is also present in several other

Siphona s.l. species belonging to several lineages (see Table 2), it is possible that

this state is not independently derived in S. ( Siphonopsis ) but rather synapotypic of

this taxon and one of these lineages. Much convergence has occurred among

Siphona s.l. lineages in the development of elongate labella, so this character can not

be used reliably for the grouping of the supraspecific taxa of this genus.

Geographic distribution

Three described and at least 13 undescribed species are included in S.

( Siphonopsis ). S. plusiae is the most widely distributed of the described species,

ranging in the west from Oregon to southern Mexico. A closely related,

undescribed, species is sympatric with S. plusiae , but restricted in known range to

California and Arizona. S. conata is the only other species of this subgenus known

north of Mexico; it is recorded from the San Francisco Bay area, Santa Cruz Island

and San Bernardino Mtns. of California.

As in S. ( Pseudosiphona ), the greatest diversity of S. ( Siphonopsis ) species is in

the Neotropics, particularly southern Mexico and southeastern Brazil (the latter area

includes the type locality of the only described Neotropical species, S. brasiliensis ).

At least two species range from southern Mexico to southern Brazil. Several species

are recognized from western South America from specimens collected in Colombia,

Ecuador, Peru, and as far south as central Chile. Specimens have also been collected

from Jamaica and Puerto Rico in the Caribbean.

Systematics of the Genus Group Taxa of the Siphonini

123

List of described species included in Siphona ( Siphonopsis )

S brasiliensis (Townsend), 1929: 374 {Siphonopsis). Holotype female, Brazil:

Sao Paulo (USNM). Holotype examined.

N conata (Reinhard), 1959: 162 {Siphonopsis). Holotype male, USA: California,

Marin Co., Mill Valley (CAS). Holotype examined.

N plusiae Coquillett, 1895a: 125. Lectotype male (by designation of Coquillett,

1897: 76), USA: California (USNM). Lectotype examined.

List of examined, undescribed, species included in New World Siphona

{Siphonopsis)

S. ( Siphonopsis ) sp. A: Males and females ranging from Oregon, USA, to northern Mexico (CAS, CNC,

JEOH, MSU, PHA, UCB, UCD, UCR, UKL, USNM, WSUP). [Specimens of this “species” are noticeably

and consistently different from those of S. plusiae , but have been frequently collected along with the latter -

including at aggregation sites - so may represent variants of S. plusiae .]

S. (Siphonopsis) sp. B: Males and females ranging from Durango to Chiapas, Mexico (CNC).

S. ( Siphonopsis ) sp. C: Males from Chiapas, Mexico (CNC).

S. ( Siphonopsis ) sp. D: One male from Chiapas, Mexico (CNC).

S. ( Siphonopsis ) sp. E: Males and females from SE Brazil (BMNH, CAS, CNC, USP). Males from southern

Mexico, Guatemala and Costa Rica are possibly conspecific (CNC, MSU, UCB, USNM).

S. ( Siphonopsis ) sp. F: One male from Veracruz, Mexico (CNC).

S. (Siphonopsis) sp. G: Two males from Ecuador (CNC).

S. (Siphonopsis) sp. H: Two males from Peru (CNC).

S. (Siphonopsis) sp. I: One male from Peru (CNC).

[No S. (Siphonopsis) sp. J.]

S. (Siphonopsis) sp. K: Males from southern Mexico and SE Brazil (BMNH, CNC, UCB, USP).

S. (Siphonopsis) sp. L: Males from SE Brazil (CNC).

S. (Siphonopsis) sp. M: Males and females from Brazil (CAS, CNC, USP) and Argentina (BMNH), and

possibly Ecuador (CNC) and Peru (AMNH, USNM).

S. (Siphonopsis) sp. N: One male from Ecuador, males and two females from Chile (CNC).

Unstudied material includes specimens from the Caribbean: Puerto Rico (USNM) and Jamaica (USNM).

Siphona (subgenus Uruactia Townsend)

Figs. 68, 99, 131.

Uruactia Townsend, 1927: 256. Type-species, U. uruhuasi Townsend, 1927 (original designation). New

subgeneric status in Siphona Meigen.

Recognition

This taxon of one described and one undescribed species is known only from

three male specimens collected in Ecuador and Peru. Adults are relatively large,

4.0-5. Omm in length, with mostly yellow legs, rather dark thoracic dorsum, wings

slightly darkened, abdomen yellow laterally on Tl+2 and T3 and rest reddish brown,

and sparse abdominal pruinosity. The prementum is about half head height in length,

and labella are padlike. These external features help in the recognition of S.

( Uruactia ) species, but examination of the male genitalia is necessary for a positive

Quaest. Ent., 1989, 25 (1,2)

124

O’Hara

identification.

The distiphallus of S. (Uruactia) species is the most distinctive feature of this

taxon. It is smoothly tapered in profile, and with (Fig. 131) or without apical spines.

It is similar in appearance to the distiphallus of S. (Aphantorhapha) species, but

these taxa differ in other genitalic features and externally ( cf ’. Recognition sections).

Other male genitalic states of S. ( Uruactia ) species include the flattened median

lobe of sternum 5 (Fig. 68), bare pregonite (Fig. 99), and inflexed cerci. These states

complement the characteristic shape of the distiphallus and help to distinguish

members of this taxon from other Siphona s.l. species.

The two known species of S. ( Uruactia ) are distinguished from one another by

number of postsutural dorsocentral setae, setulation on wing vein R4+5, relative

development of lateral marginal setae on abdominal Tl+2, and slight differences in

male genitalia (as indicated in description below).

Description

Length: 4.0-5. Omm.

Head. — Proclinate orbital setae subequal in length or anterior one longer. Eye of male medium-large,

0.84 head height (same size in all three examined specimens). Flagellomere 1 of male medium-short to

medium, 0.48-0.50 head height; shape average (sp. 31) or broad ( S . uruhuasi). Aristomere 1 short.

Aristomere two 2X longer than wide. Aristomere 3 long and evenly tapered, micropubescent. Clypeus

narrow and enclosed in membrane or slightly broadened. Palpus short, clavate. Proboscis with prementum

medium in length, slightly longer than half head height, labella padlike.

Thorax. — Prostemum setulose. Lower katepistemal seta longer than upper anterior seta. Three (S.

uruhuasi ) or four (sp. 31) postsutural dorsocentral setae. Upper part of anepistemum with one or two setulae.

Fore tibia with preapical ad seta much shorter than d seta. Mid tibia with one ad seta. Claws short. Wing

vein CwA, with distal portion 0.26-0.46 (mean 0.36). Wing setulae: /?, distally setulose dorsally, ventrally

bare; ^4+5 setulose between base and r-m in sp. 31, beyond r-m in S. uruhuasi ; CuA] bare.

Abdominal terga 1-5. — Abdomen ovoid in shape. Tl+2 without median marginal setae, lateral marginal

setae absent from S. uruhuasi , strong in sp. 31. T3-T5 with average setation.

Male genitalia (Figs. 68, 99, 131). — S5 (Fig. 68) with posterior margins of processes approximately

U-shaped; apical lobe distinctly differentiated; median lobe flattened posteromedially, more prominently so

in sp. 31 (Fig. 68); processes moderately setulose. T6 apparently absent. Ejaculatory apodeme with

fan-shaped portion subequal in width to width of hypandrial apodeme. Pregonite in profile apically pointed,

smoothly curved anteriorly in S. uruhuasi, only slightly curved anteriorly in sp. 31 (Fig. 99); bare.

Epiphallus absent. Distiphallus reduced posteriorly, entire laterally in S. uruhuasi, slightly incised laterally

with short posterolateral arm in sp. 31 (Fig. 131), in profile rather narrow and apically pointed, apical margin

with small (5. uruhuasi) to large (sp. 31) spines. Postgonite average. Surstylus long and straight, basally free

from epandrium. Cerci average length, in profile sharply inflexed at midlength; moderately setose on basal

half.

Examined male genitalia of: 5. uruhuasi and sp. 31.

Female genitalia. — Not examined.

Hosts. Unknown.

Phylogenetics

The two species are interpreted as sister species on the basis of shared states in

the male genitalia (particularly shape of median lobes on sternum 5 and shape of

distiphallus) and concordant external similarities. This taxon is retained as a

Systematics of the Genus Group Taxa of the Siphonini

125

subgenus because its sister group within Siphona s.l. is unknown.

Geographic distribution

The type species of S. (Uruactia) is known only from the male holotype,

collected from Uruhuasi, Peru, on 3.II.1910, at an elevation of 1980m, on flowers of

Baccharis sp. The only other species, sp. 31 (originally numbered in series of

unplaced New World Siphona s.l.), is represented by two male specimens, one with

the same label data as the type of S. uruhuasi (USNM) and the other collected from

Cerro Tinajillas in the Azuay province of southern Ecuador (18-21. III. 1965 at

3100m; CNC).

List of described species included in Siphona ( Uruactia )

S uruhuasi (Townsend), 1927: 364 {Uruactia). Holotype male, Peru: Uruhuasi

(USNM). Holotype examined.

List of examined, undescribed, species included in Siphona {Uruactia)

S. (Uruactia) sp. 31: One male each from Peru (USNM) and Ecuador (CNC).

New and described New World species unplaced to subgenus in Siphona s.l.

New World Siphona species group 1

Figs. 69, 100-101, 132.

This group of approximately nine known species (all undescribed) is Neotropical

in distribution, ranging from southern Mexico to southeastern Brazil. They are

typical members of Siphona s.l., with no external features interpretable as

synapotypic of the group. They include spp. 19,21,24-303, and are diagnosed as

follows (also see Table 1): male eye medium-small to medium-large; male

flagellomere 1 medium-short to medium-long, markedly varied in shape from linear

or broadened to very broad and subquadrangular or almost triangular; aristomere 1

as long as wide in spp. 26 and 27, short in others; proboscis with labella varied from

padlike (a few species) to almost length of prementum (most species), flexible in life

on apical half or more; three or four postsutural dorsocentral setae; lower

katepistemal seta longer than upper anterior seta; postscutellum unusually narrow

and projected in spp. 29 and 30; Rx and R4+5 with setulation varied, CuAx bare;

crossvein dm-cu near wing margin in most species, moderately removed from it in a

few.

Male genitalia of all nine species examined. Similarities in male genitalic

features (particularly of the distiphallus) suggest monophyly of the group. S5 (Fig.

69) with posterior margins of processes approximately U-shaped; apical lobe

3

Numbering of species is explained in Methods chapter.

Quaest. Ent., 1989, 25 (1,2)

126

O’Hara

distinctly differentiated, apex curved inward in a few species; median lobe markedly

varied from narrow to elongate, rounded to truncate, or rather flattened

posteromedially; slightly differentiated accessory lobe in sp. 27, others without.

Pregonite (Figs. 100-101) in profile slender to average width, apically rounded to

pointed; patch of small spines on apical half or less of sclerotized portion in most

species (Fig. 100), spines absent from others (Fig. 101), tiny to moderate-sized seta

posteriorly in a few species (absent from most). Epiphallus absent. Distiphallus (Fig.

132) relatively distinctive, not incised laterally, without enlarged teeth, in profile

nearly parallel-sided on apical half or more and truncate apically. Postgonite

markedly reduced in most species, average in a few. Surstylus varied from short to

long, narrow to broad, straight to curved posteriorly; basally free from epandrium.

Cerci thick on apical half in sp. 21, average in others, smoothly curved at midlength.

Female genitalia (examined in spp. 24,27,30) short, unmodified except sternum 8

broader than average in sp. 30.

List of examined, undescribed, species included in New World Siphona species

group 1

Siphona species group 1, sp. 19: One male from Chiapas, Mexico (CNC).

Siphona species group 1, sp. 21: Males and females from SE Brazil, one male from Colombia (CNC).

Siphona species group 1, sp. 24: Males and females from northern Argentina (CNC), one male from Ecuador

(USNM).

Siphona species group 1, sp. 25: One male from northern Argentina (CNC).

Siphona species group 1, sp. 26: Males from Ecuador (BMNH, CNC).

Siphona species group 1, sp. 27: One male and one female from Ecuador (CNC).

Siphona species group 1, sp. 28: One male from Chiapas, Mexico (CNC), possibly conspecific males from

Guatemala (USNM) and Costa Rica (USNM).

Siphona species group 1 , sp. 29: Males and one female from Ecuador (CNC).

Siphona species group 1, sp. 30: Males and females from Ecuador (CNC).

New World Siphona species group 2

Figs. 19, 70, 102-103, 133-134.

This group comprises about ten species4 distributed between southern Arizona

and southeastern Brazil. They are typical members of Siphona s.l., with the

following characteristics (also see Table 1): (head of sp. 1 shown in Fig. 19) male

eye medium-large to large; male flagellomere 1 medium-short to medium, linear to

subquadrangular in shape; aristomere 1 short; proboscis with labella about

two-thirds length of prementum in sp. 8, padlike in others; three postsutural

dorsocentral setae; lower katepistemal seta longer than upper anterior seta; wing

setulation varied; crossvein dm-cu slightly removed from wing margin in most

species.

These species are numbered from one to ten, separately from those of species groups 1 and 3 and unplaced

New World species, as explained in Methods chapter.

Systematics of the Genus Group Taxa of the Siphonini

127

Male genitalia of all ten included species examined. S5 (Fig. 70) with posterior

margins of processes approximately U-shaped; apical lobe distinctly differentiated,

apex curved inward in a few species; median lobe markedly varied from broadly

rounded to elongate, and truncate or flattened posteromedially, with accessory lobe

distinctly differentiated in a few species. Pregonite (Figs. 102-103) varied in profile,

smoothly to sharply curved anteriorly, average to broad on basal half; with or

without patch of small spines on apical half or less of sclerotized portion and in sp. 2

(Fig. 103) with longitudinal row of spines, bare posteriorly or with tiny to

moderate-sized seta. Epiphallus absent. Most included species are recognized as

members of this species group by features of the distiphallus: most possess

posterolaterally on each side a ventrally or anteroventrally directed projection or

hook (Figs. 133-134); a few species lack this feature of the distiphallus but are

included in this species group (with varied degrees of confidence) because of other

external and genitalic similarities; posterior margin of distiphallus entire in sp. 2,

incised in others. Postgonite markedly reduced in spp. 3 and 10, short in sp. 2,

average in others. Surstylus average length to elongate, straight to curved

posteriorly; basally free from epandrium. Cerci rather short to average length,

straight to smoothly curved.

Female genitalia (examined in spp. 3 and 6) short, unmodified except sternum 8

broader than average.

The conservative external features of members of Siphona species group 2 make

this group difficult to recognize. In addition certain species are less apotypic than

most in their male genitalic features, so their inclusion here is tentative and is based

on general similarities.

List of examined, undescribed, species included in New World Siphona species

group 2

Siphona species group 2, sp. 1: Males and females ranging from southern Arizona, USA, to Chiapas, Mexico

(CAS, CNC, UAT).

Siphona species group 2, sp. 2: One male from Michoacan, Mexico (USNM).

Siphona species group 2, sp. 3: Males and females from Chiapas, Mexico (CNC).

Siphona species group 2, sp. 4: One male from Chiapas, Mexico (CNC).

Siphona species group 2, sp. 5: One male from Panama (USNM), males and one female from Peru (CNC).

Siphona species group 2, sp. 6: Males and females from SE Brazil (CNC, INPA, USP).

Siphona species group 2, sp. 7: Males from Peru (CNC).

Siphona species group 2, sp. 8: Males from Peru (CNC).

Siphona species group 2, sp. 9: One male from Costa Rica (USNM).

Siphona species group 2, sp. 10: One male from Chiapas, Mexico (CNC).

New World Siphona species group 3

Figs. 104-105, 135.

Five undescribed New World species are recognized as belonging to a

monophyletic lineage possibly related to Siphona species group 1 (see below).

Quaest. Ent., 1989, 25 (1,2)

128

O’Hara

Included are spp. 3, 4, 6, 12 and 22 (SE Brazil)5. Externally these flies are quite

varied, particularly in head features and wing setulation, and for this reason

specimens of some of these species appear more externally similar to excluded

species than to their own group members. As a rule, male genitalic characteristics

are more reliable indicators of relationship within Siphona s.l. than are external

features (species of S. (P seudo siphona) and S. (Siphonopsis) being good examples),

so little weight is given here to such homoplastic external similarities.

Siphona species group 3 is diagnosed as follows (also see Table 1): male eye

medium to medium-large; male flagellomere 1 markedly varied, medium-short to

long; shape linear to very broadly triangular; proboscis with labella slightly

lengthened in some specimens of sp. 22, padlike in others; three postsutural

dorsocentral setae; varied wing setulation; position of crossvein dm-cu varied; legs

and abdomen average; (male genitalia examined in all five species) sternum 5 with

posterior margins of processes approximately U-shaped, apical lobe not apically

curved inward, median lobe rounded to elongate and truncate, distinctly

differentiated accessory lobes in sp. 4; pregonite (Figs. 104-105) in profile rather

sharply curved at midlength or smoothly curved anteriorly, with longitudinal ridge

of spines laterally (resembling pregonite of Entomophaga and Proceromyia except

possessing tiny to moderate-sized seta posteriorly in most specimens \i.e. of two

examined specimens of sp. 22, one has tiny seta and the other is bare, Fig. 105] -

this seta only present among Siphona group taxa); pregonite without group of small

spines apically; epiphallus absent; distiphallus (Fig. 135) not incised laterally,

characteristically squared-off basally, with short, parallel-sided apical portion

ending in truncate tip, moderately developed spines anteriorly in sp. 22, only

spinules present in others; postgonite average to reduced; surstylus and cerci

average; female genitalia short and unmodified (examined only in sp. 3).

Two members of Siphona species group 1 (spp. 26 and 27) have similar

distiphallus structure to species of group 3, but differ in features of the pregonite.

Species of group 3 possess a spined longitudinal ridge on the outer surface of the

pregonite, a state not found in other species of Siphona s.l. except in Siphona sp.

grp. 2, sp. 2 (Fig. 103). There are both external similarities and dissimilarities

between members of both groups 1 and 3, but these involve such homoplastic

characters that their phylogenetic interpretation is difficult. I have not studied the

groups in sufficient detail to offer any definite views about the interrelationships of

groups 1 and 3, but I suggest that certain male genitalic similarities may indicate

that the species of group 3 are derived members of group 1 or the groups are sister

groups.

I have presented the information above about Siphona group 3, as I have

elsewhere for groups 1 and 2, to provide a tentative framework upon which a formal

taxonomic investigation of the numerous undescribed and unplaced New World

5

Numbering of these species is explained in Methods chapter.

Systematics of the Genus Group Taxa of the Siphonini

129

Siphona s.l. species can be built. These notes about undescribed New World

siphonines ought to prove useful in the eventual revision of this large and unworked

assemblage of Siphona species.

List of examined, undescribed, species included in New World Siphona species

group 3

Siphona species group 3, sp. 3: Males and females ranging from Washington state to southern California and

Arizona, USA (CAS, CNC, PHA, UCB, UCR, USNM, WSUP).

Siphona species group 3, sp. 4: One male each from Veracruz and Chiapas, Mexico (CNC).

Siphona species group 3, sp. 6: One male from Florida, USA (AMNH), one possibly conspecific male from

Chiapas, Mexico (USNM).

Siphona species group 3, sp. 12: Two males from Iowa, USA (PHA), one male from Ohio, USA (CNC) and

one female from Ontario, Canada (CNC).

Siphona species group 3, sp. 22: Males from SE Brazil (CAS, CNC, USP).

Unplaced New World species of Siphona sensu lato

Figs. 20, 106, 136.

I have left two described species unplaced within New World Siphona s.l.:

Siphona panamensis (Curran) and Siphona pulla (Reinhard). Though typical

members of Siphpna s.l., these species cannot be associated with any of the

subgenera here recognized nor the three additional species groups discussed. Both

are difficult to place because they are only known from female holotypes, and if

males were available then unequivical placement of these species might be possible.

Specimens of many undescribed species were examined during the course of this

revision. Those belonging to relatively easily recognized genera are incorporated

into the descriptions of those genera and not themselves described or specially

treated. This practise is more difficult within Siphona s.l. because monophyletic

lineages are difficult to recognize and many species cannot be associated with the

lineages that are. The complexity is all the more difficult to discuss since most of the

species are undescribed. As this work is primarily a generic level revision it is

impractical at this time to describe these new species, though discussing them in an

informal manner will convey some idea of their diversity. Where possible,

descriptions of certain subgenera (particularly S. (Pseudosiphona) and S.

( Siphonopsis )) have been modified for inclusion of appropriate undescribed species,

but even though this has been done and Siphona groups 1 to 3 informally diagnosed,

at least 15 undescribed New World species remain that I have studied but not been

able to place. I outline below some of the diversity contained within this group of

mostly unrelated species because they may remain undescribed for some time. As

new groups or subgenera are erected for some of these species and other species

identified as related to groups recognized here, the descriptions of the latter will

have to be modified for such inclusions. For the present I have retained a rather

conservative approach toward recognition of New World Siphona subgenera in

order to keep them as monophyletic lineages (and thus, I believe, more meaningful

Quaest. Ent., 1989, 25 (1,2)

130

O’Hara

entities) in the midst of a plethora of undescribed forms.

The species discussed here include S. panamensis, S. pulla , and 15 numbered

species: spp. 1,2,5,7,8,10,13-17,20,23,32 and 346. Their range of variation is given

in Table 1 under the heading of “ Siphona s.l., unplaced”, and the assemblage is

briefly diagnosed here: (head of sp. 23 shown in Fig. 20) male eye medium-small to

very large; male flagellomere 1 short to medium, shape linear, broadened,

subquadrangular or broad and triangular; proboscis with labella padlike in most

species, slightly lengthened in a few, about as long as prementum in spp. 13 and 23

(Fig. 20); three postsutural dorsocentral setae in most species, four in a few; lower

katepistemal seta longer than upper anterior seta in most species, subequal in length

in a few, shorter in spp. 14 and 17; R j and i?4+5 with setulation varied, CuAx bare;

crossvein dm-cu positioned near or removed from wing margin; legs average.

Male genitalia of 14 species examined (spp. 1-2,5,7-8,10,13-17,23,32,34). S5

with posterior margins of processes approximately U-shaped in most species,

obtusely angled in a few, nearly V-shaped in spp. 7 and 15 (though median cleft

more distinct than in Actia species); apical lobe distinctly differentiated, apex curved

inward in some species; median lobe varied, rounded, elongate, truncate, or slightly

flattened posteromedially, accessory lobe present in sp. 5; pregonite (Fig. 106)

varied, in profile slender to broad, apically pointed to rounded, small spines on

sclerotized portion apically in a few species, with tiny to moderate-sized seta

posteriorly in most species, seta absent from a few. Epiphallus absent. Distiphallus

(Fig. 136) markedly varied, laterally incised or complete, with or without enlarged

spines apically, in profile varied apically from pointed or rounded to truncate.

Postgonite average in most species, reduced in spp. 32 and 34. Surstylus and cerci

average in most species, surstylus basally free from epandrium.

Female genitalia examined in seven species (spp. 1-2,5,10,14,20,23), short; S6

sharply keeled posteromedially in sp. 20, flat in others; T6 absent, or present as

lateral sclerites or (in sp. 3) as a dorsally continuous sclerite; S7 with rather short

anterior apodeme in sp. 20, long in others, sharply keeled posteromedially only in

sp. 20; S8 average in most species, very broad in sp. 5, with thick setae in sp. 20.

The highest concentration of species (six) is in southern Mexico, with seven

other species distributed throughout North America from Alaska to northcentral

Mexico and four species distributed between southeastern Brazil and lower Central

America.

Hosts are known only for New World species 7 and 8, and include larvae of

several species of the Geometridae and one of the Noctuidae. Species 7 and 8 are

recorded from southern British Columbia, Canada.

6

Numbering of these species is explained in Methods chapter.

Systematics of the Genus Group Taxa of the Siphonini

131

List of described species unplaced to subgenus in New World Siphona sensu

lato:

S Actia panamensis Curran, 1933a: 3. Holotype female, Panama: Barro Colorado

Is. (AMNH). Holotype examined. (Removed from Actia.)

N,S Aphantorhapha pulla Reinhard, 1974: 1157. Holotype female, Mexico:

Morelos, Cuernavaca (CNC). Holotype examined. (Removed from

Aphantorhapha. )

List of examined, undescribed, species unplaced in New World Siphona sensu

lato

Unplaced Siphona sp. 1: Males and females from western North America (CNC, PHA, UCB, USNM,

WSUP).

Unplaced Siphona sp. 2: Males and females ranging from Alaska to Wisconsin, USA (CNC, USNM).

Unplaced Siphona sp. 5: Males and females from southern Mexico (CNC), one male from El Salvador

(USNM).

Unplaced Siphona sp. 7: Males and females from southern British Columbia, Canada, one male from

California, USA, and one possibly conspecific male from Arizona, USA (CNC).

Unplaced Siphona sp. 8: Males and females from southern British Columbia, Canada (CNC).

Unplaced Siphona sp. 10: One male from Ontario, Canada (CNC) and one female from New Brunswick,

Canada (JEOH).

Unplaced Siphona sp. 13: One male from Florida, USA (CNC).

Unplaced Siphona sp. 14: Males and females from Chiapas, Mexico (CNC).

Unplaced Siphona sp. 15: One male from Oaxaca, Mexico (JEOH).

Unplaced Siphona sp. 16: One male from Durango, Mexico (CNC).

Unplaced Siphona sp. 17: One male from Chiapas, Mexico (CNC).

Unplaced Siphona sp. 20: One female from Chiapas, Mexico (CNC).

Unplaced Siphona sp. 23: Males and females from SE Brazil, Ecuador, Costa Rica and Canal Zone (AMNH,

BMNH, CNC, USNM, USP).

Unplaced Siphona sp. 32: Males from SE Brazil (USP).

Unplaced Siphona sp. 34: Males and females from SE Brazil (CNC, USNM, USP), one male from Ecuador

(CNC) and one female from Costa Rica (USNM).

Actia heterochaeta Bezzi, unplaced species of the Siphonini

I have been unable to locate the female type of this species, described from

Ethiopia by Bezzi (1908: 59). The type apparently does not reside in Milan (MCSN)

with many of the other Bezzi types (Amaud 1982).

Actia heterochaeta was described by Bezzi under a concept of Actia now

equivalent to the entire Siphonini. The original description fails to mention a single

generic characteristic that would help in the placement of this species. Bezzi

compares A. heterochaeta with A. bicolor (now in Ceromya ) in his original

description, and then with A. stiglinae (now in Peribaea) in a later publication

(1928: 204). It is clear that Bezzi’s concept of “allied species” was based on shared

similarities and not commonality of descent, so his species comparisons provide no

clue about the correct placement of A. heterochaeta.

Quaest. Ent., 1989, 25 (1,2)

132

O’Hara

I examined a male specimen in the BMNH collected from Nigeria and identified

by Villeneuve as A. heterochaeta. This specimen is probably the one mentioned by

Crosskey (1980: 855) as a questionable new locality record for the species.

However, Villeneuve’ s identification is probably in error because the specimen (a

Peribaea species) differs from the description of A. heterochaeta in two important

respects: Rx is bare in A. heterochaeta and distally setulose in Villeneuve’s

specimen, and coloration differs markedly (even given that Bezzi’s specimen was a

female and Villeneuve’s specimen a male). Without any concrete indication of the

generic characteristics of A. heterochaeta , I must leave this species unplaced in the

Siphonini, as did Crosskey (1980: 855).

Systematics of the Genus Group Taxa of the Siphonini

133

Figs. 1-4. Head profiles of male siphonines (vestiture of postcranium omitted). Scale bars = 0.5mm: 1,

Goniocera io; 2, Proceromyia macronychicr, 3, Proceromyia pubioculata\ 4, Entomophaga exoleta.

Quaest. Ent., 1989, 25 (1,2)

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O’Hara

Figs. 5-8. Head profiles of male siphonines (vestiture of postcranium omitted). Scale bars = 0.5mm: 5,

Entomophaga nigrohalteratcr, 6, Ceromya flaviceps-, 7, Ceromya New Guinea sp. 15; 8, Actia lamia.

Systematics of the Genus Group Taxa of the Siphonini

135

Figs. 9-12. Head profiles of male siphonines (vestiture of postcranium omitted). Scale bars = 0.5mm: 9,

Actia Nepal sp. 1; 10, Actia parviseta (from holotype, proboscis reoriented); 11, Peribaea cervina (from

holotype, vibrissae missing); 12, Peribaea tibialis.

Quaesl. Ent., 1989, 25 (1,2)

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O’Hara

Figs. 13-16. Head profiles of male siphonines (vestiture of postcranium omitted). Scale bars = 0.5mm: 13,

Siphona (Actinocrocuta) singularis complex; 14, Siphona ( Aphantorhapha ) arizonica; 15, Siphona

( Baeomyia ) xanthogaster (from holotype); 16, Siphona (Ceranthia) flavipes.

Systematics of the Genus Group Taxa of the Siphonini

137

Figs. 17-20. Head profiles of male siphonines (vestiture of postcranium omitted). Scale bars = 0.5mm: 17,

Siphona (Pseudosiphona) brevirostris\ 18, Siphona (Siphonopsis) plusiae\ 19, Siphona s.l., sp. grp. 2, sp. 1;

20, Siphona s.l., unplaced sp. 23.

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138

O’Hara

i i

21. Actia interrupta

22. S. (Baeomyia) xanthogaster

Figs. 21-22. Right wings of siphonines. Scale bars = 0.5mm: 21, Actia interrupta, with wing veins labelled;

22, Siphona (Baeomyia) xanthogaster.

139

Systematics of the Genus Group Taxa of the Siphonini

Figs. 23-26. Comparison of proboscis structure. Scale bars = 200pm: 23, Actia diffidens , with padlike

labella; 24, Siphona (Siphonopsis) plusiae, with elongate labella; 25, Siphona (Siphona) maculata, with long

labella; 26, Siphona ( Siphona ) pisinnia, with very long labella (Ibl, labella; prem, prementum). Figs. 27-28.

Left lateral view of prothorax, for comparison of proepimeral setae. Scale bars = 100pm: 27, Peribaea sp.,

illustrating two strong and opposed proepimeral setae (a spr , anterior spiracle; / prepm s , lower proepimeral

seta; u prepm s, upper proepimeral seta); 28, Siphona (Siphona) maculata, illustrating single strong

proepimeral seta (upper and lower setae indicated by arrows). Figs. 29-30. Comparison of labella of

proboscis. Scale bars = 100pm: 29, labella of Actia diffidens, illustrating padlike condition and numerous

pseudotracheae; 30, labella of Siphona (Siphonopsis) plusiae, illustrating elongate condition and relatively

few pseudotracheae.

Quae st. Ent., 1989, 25 (1,2)

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O’Hara

Figs. 31-32. Left lateral view of thorax, for comparison of katepistemal and anepistemal setae. Scale bars =

250pm: 31, Actia diffidens, illustrating short lower katepistemal seta (/ kepst s) and two anepistemal setulae

(anepst s); 32, Siphona (Siphona) maculata, illustrating long lower katepistemal seta and single anepistemal

setula (indicated by arrows). Figs. 33-34. Ventral view of mesothorax, for comparison of posteroventral

region of katepistemum. Scale bars = 200pm: 33, Actia diffidens, illustrating row of hairs on katepistemum

( kepst h) anterior to mid coxa; 34, Siphona (Siphona) maculata, illustrating presence of several hairs on

katepistemum restricted to midline region (indicated by arrow). Figs. 35-36. Comparison of anterior surface

of mid femur. Scale bars = 200pm: 35, Ceromya varichaeta, illustrating patch of tiny hairs on anterior

surface of mid femur (fem h)\ 36, Siphona (Siphona) maculata, illustrating typical condition of anterior

surface of mid femur. Figs. 37-38. Comparison of anterior surface of mid tibia. Scale bars = 200pm: 37,

Ceromya varichaeta, illustrating presence of strong anterodorsal seta (ad 5) on mid tibia; 38, Siphona

(Baeomyia) xanthogaster, illustrating absence of anterodorsal seta on mid tibia.

Systematics of the Genus Group Taxa of the Siphonini 141

Figs. 3<M-2. Comparison of male pregonite structure. Scale bars = 40|lm: 39, left lateral view of pregonite of

Ceromya americana complex, illustrating spinules on membranous portion ( memb p, membranous portion;

scl p, sclerotized portion; spin, spinules); 40, ventrolateral view of pregonite of Ceromya americana

complex; 41, left lateral view of pregonite of Actia diffidens, illustrating large spines (sp) on sclerotized

portion; 42, ventral view of pregonite of Actia diffidens. Figs. 43^46. Comparison of female genitalia. Scale

bars = 100|im: 43, ventral view of female genitalia of Siphona (Siphona) maculata, illustrating unmodified

sterna (sterna numbered; cer, cercus); 44, ventral view of female genitalia of Siphona (Ceranthia) flavipes,

illustrating keel ( [Id ) posteromedially on sternum 7; 45, lateral view of female genitalia of Siphona ( Siphona )

maculata ; 46, lateral view of female genitalia of Siphona (Ceranthia) flavipes.

Quaest. Ent., 1989, 25 (1,2)

142

O’Hara

posienor

47. Goniocera schistacea

50. Enlomophaga exolela

48. Proceromyia macronychia

49. Proceromyia pubioculala

51 . Enlomophaga nigrohalterala

52. Ceromya silacea

54. Ceromya New Guinea sp. 18

55. Ceromya americana complex

Figs. 47-58. Ventral view of male sternum 5. Scale bars = 0.1mm: 47, Goniocera schistacea (Y68, CNC)

(ap /, apical lobe; bs p, basal plate; descl a, desclerotized area; med c, median cleft; med /, median lobe); 48,

Proceromyia macronychia (Y40, CNC); 49, Proceromyia pubioculata (Y85, JEOH); 50, Entomophaga

exoleta (Y132, ZMUC); 51, Entomophaga nigrohalterata (Y27, CNC); 52, Ceromya silacea (Y179, CNC);

53, Ceromya flaviceps (Y107, CNC); 54, Ceromya New Guinea sp. 18 (Y175, BLKU); 55, Ceromya

americana complex (Y26, CNC); 56, Ceromya amblycera (Y45, UCS); 57, Ceromya lutea (Y121, CNC);

58, Actia lamia (Y17, JEOH).

Systematics of the Genus Group Taxa of the Siphonini

143

60. Peribaea apicalis

61. Peribaea Australia sp. 1

Figs. 59-70. Ventral view of male sternum 5. Scale bars = 0.1mm: 59, Actia autumnalis (Y22, CNC); 60,

Peribaea apicalis (Y82, CNC); 61, Peribaea Australia sp. 1 (Y83, DPI); 62, Siphona (Actinocrocuta)

singulars complex (0120, CNC); 63, Siphona (Aphantorhapha) arizonica (unnumbered, JEOH); 64,

Siphona (Aphantorhaphopsis) Nepal sp. 1 (W75, CNC); 65, Siphona (Ceranthia) flavipes (B8, CNC); 66,

Siphona ( Pseudosiphona ) sp. 4 (R22, CNC); 67, Siphona ( Siphona ) cristata (W38, JEOH); 68, Siphona

( Uruactia ) sp. 31 (R45, CNC); 69, Siphona s.l., sp. grp. 1, sp. 28 (027, CNC); 70, Siphona s.l., sp. grp. 2, sp.

9(0154, USNM).

Quaest. Ent., 1989, 25 (1,2)

71. Goniocera io

72. Goniocera versicolor

73. Proceromyia macronychia

76. Entomophaga nigrohallerala

77. Ceromya flaviceps 78. Ceromya New Guinea sp. 6 79. Ceromya americana complex

Figs. 71-79. Lateral view of left pregonite of male. Scale bars = 0.05mm: 71, Goniocera io (Y41, CNC); 72,

Goniocera versicolor (Y69, CNC); 73, Proceromyia macronychia (Y40, CNC); 74, Proceromyia

pubioculata (Y85, JEOH); 75, Entomophaga exoleta (Y132, ZMUC); 76, Entomophaga nigrohalterata

(Y27, CNC); 77, Ceromya flaviceps (Y107, CNC); 78, Ceromya New Guinea sp. 6 (Y176, BLKU); 79,

Ceromya americana complex (Y25, CNC).

Systematics of the Genus Group Taxa of the Siphonini

145

83. Ceromya lulea

84. Ceromya silacea

85. Actia lamia

86. Actia diffidens

87. Peribaea tibialis

88. Peribaea Australia sp. 3

Figs. 80-88. Lateral view of left pregonite of male. Scale bars = 0.05mm: 80, Ceromya cornuta (Y 1 14,

CNC); 81, Ceromya Mexico sp. 5 (Y1 18, CNC); 82, Ceromya Brazil sp. 3 (Y125, INPA); 83, Ceromya lutea

(Y121, CNC); 84, Ceromya silacea (Y179, CNC); 85, Actia lamia (Y17, JEOH); 86, Actia diffidens (Y21,

CNC); 87, Peribaea tibialis (Y15, JEOH); 88, Peribaea Australia sp. 3 (Y77, DPI).

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O’Hara

89. S. ( Actinocrocuta ) singular is complex 90. S. ( Aphantorhapha ) arizonica

91. S. ( Aphaniorhaphopsis ) crassutala

92. S. (Aphaniorhaphopsis) starkei

93. S. ( Baeomyia) xanthogaster

95. S. ( Pseudosiphona) brevirostris

96. S. ( Siphona) crislaia

97. S. (Siphonopsis) plusiae

Figs. 89-97. Lateral view of left pregonite of male. Scale bars = 0.05mm: 89, Siphona ( Actinocrocuta )

singularis complex (0120, CNC); 90, Siphona ( Aphantorhapha ) arizonica (unnumbered, JEOH); 91,

Siphona (Aphantorhaphopsis) crassulata (B3, CNC); 92, Siphona (Aphantorhaphopsis) starkei (B96,

JEOH); 93, Siphona ( Baeomyia ) xanthogaster (B98, JEOH); 94, Siphona (Ceranthia) flavipes (B8, CNC);

95, Siphona (Pseudosiphona) brevirostris (W89, CNC); 96, Siphona ( Siphona ) cristata (W37, JEOH); 97,

Siphona ( Siphonopsis ) plusiae (R97, JEOH).

Systematics of the Genus Group Taxa of the Siphonini

147

98. S. (Siphonopsis) sp. N

99. S. (llruactia) sp. 31

100. Siphona s.l., sp. grp. 1, sp. 27

103. Siphona s.l., sp. grp. 2, sp. 2

104. Siphona s.l.. sp. grp. 3, sp. 3

105. Siphona s.l.. sp. grp. 3, sp. 22

106. Siphona s.l., unplaced sp. 1

Figs. 98-106. Lateral view of left pregonite of male. Scale bars = 0.05mm: 98, Siphona (Siphonopsis) sp. N

(024, CNC); 99, Siphona ( Uruactia ) sp. 31 (R45, CNC); 100, Siphona s.l., sp. grp. 1, sp. 27 (R46, CNC);

101, Siphona s.l., sp. grp. 1, sp. 28 (027, CNC); 102, Siphona s.l., sp. grp. 2, sp. 1 (0125, JEOH); 103,

Siphona s.l., sp. grp. 2, sp. 2 (B92, USNM); 104, Siphona s.l., sp. grp. 3, sp. 3 (B81, WSUP); 105, Siphona

s.l., sp. grp. 3, sp. 22 (050, USP); 106, Siphona s.l., unplaced sp. 1 (R62, WSUP).

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110. Entomophaga exolela

108. Proceromyia macronychia

111. Entomophaga nigrohallerata

109. Proceromyia pubioculata

112. Ceromya silacea

Figs. 107-1 12. Lateral view of male distiphallus. Scale bars = 0.05mm: 107, Goniocera io (Y41, CNC); 108,

Proceromyia macronychia (Y40, CNC) (r lat marg, reduced lateral margin); 109, Proceromyia pubioculata

(Y85, JEOH); 110, Entomophaga exoleta (Y132, ZMUC); 111, Entomophaga nigrohalterata (Y27, CNC);

1 12, Ceromya silacea (Y179, CNC) (/ pslat marg, long apical extension of posterolateral margin).

Systematics of the Genus Group Taxa of the Siphonini

149

115. Ceromya cornuta

118. Actia autumnalis

Figs. 113-118. Lateral view of male distiphallus. Scale bars = 0.05mm: 113, Ceromya flaviceps (Y107,

CNC); 1 14, Ceromya americana complex (Y25, CNC); 1 15, Ceromya cornuta (Y 1 14, CNC); 1 16, Ceromya

lutea (Y121, CNC) (pos marg, posterior margin entirely sclerotized; cf. sclerotized projection labelled in

Figs. 125, 127); 117, Actia lamia (Y17, JEOH); 1 18, Actia autumnalis (Y22, CNC).

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121 . Peribaea fissicornis

122. S. ( Actinocrocuta) singulars complex 123. S. ( Actinocrocuia) sp. 1 124. S. ( Aphanlorhapha) arizonica

Figs. 119-124. Lateral view of male distiphallus. Scale bars = 0.05mm: 119, Actia New World sp. 4 (Y48,

USP); 120, Peribaea apicalis (Y82, CNC); 121, Peribaea fissicornis (Y31, CNC); 122, Siphona

(Actinocrocuta) singularis complex (0120, CNC); 123, Siphona (Actinocrocuta) sp. 1 (R4, CNC); 124,

Siphona (Aphantorhapha) arizonica (unnumbered, JEOH) (undif p slat marg, undifferentiated posterolateral

margin).

Systematics of the Genus Group Taxa of the Siphonini

151

■+ anterior

126. S. ( Baeomyia) hurdi

128. S. ( Pseudosiphona) sp. 1 129. S. ( Siphona) crislala

130. S. ( Siphonopsis) plusiae

Figs. 125-130. Lateral view of male distiphallus. Scale bars = 0.05mm: 125, Siphona (Aphantorhaphopsis)

nigronitens (R13, CNC) ( m pslat marg, moderate apical extension of posterolateral margin beyond rest of

lateral margin; scl proj, sclerotized projection posteriorly); 126, Siphona ( Baeomyia ) hurdi (RIO, CNC);

127, Siphona (Ceranthia) flavipes (R81, CNC) (/ pslat marg, long apical extension of posterolateral margin

beyond rest of lateral margin; scl proj, sclerotized projection posteriorly); 128, Siphona ( Pseudosiphona ) sp.

1 (B69, CNC); 129, Siphona (Siphona) cristata (W36, JEOH); 130, Siphona (Siphonopsis) plusiae (B18,

CNC).

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131. S.(Uruactia) sp. 31

132. Siphona s.l., sp. grp. 1, sp. 25 133. Siphona s.l., sp. grp. 2. sp. 1

Figs. 131-136. Lateral view of male distiphallus. Scale bars = 0.05mm: 131, Siphona (Uruactia) sp. 31

(R45, CNC); 132, Siphona s.l., sp. grp. 1, sp. 25 (R2, CNC); 133, Siphona s.l., sp. grp. 2, sp. 1 (0148, CAS);

134, Siphona s.l., sp. grp. 2, sp. 3 (R65, CNC); 135, Siphona s.l., sp. grp. 3, sp. 3 (B81, WSUP); 136,

Siphona s.l., unplaced sp. 1 (R62, WSUP).

Systematics of the Genus Group Taxa of the Siphonini

153

138. Ceromya americana complex

141. S. (Siphona) cristala

142. S. ( Actinocrocuta) 143 5 ( Aphantorhapha) sp. 1 144. S. (Siphona) kairiensis

singularis complex

Figs. 137-141. Lateral view of male surstylus and cerci. Scale bars = 0.1mm: 137, Goniocera io (Y41,

CNC); 138, Ceromya americana complex (Y25, CNC); 139, Peribaea apicalis (Y82, CNC); 140, Siphona

(Pseudosiphona) sp. 1 (B69, CNC); 141, Siphona ( Siphona ) cristata (W36, JEOH). Figs. 142-144. Posterior

view of male genitalia, vestiture omitted. Scale bars = 0.1mm: 142, Siphona (Actinocrocuta) singularis

complex (lectotype of S. chaetosa (Tnsd.), USNM); 143, Siphona (Aphantorhapha) sp. 1 (0182, BMNH);

144, Siphona (Siphona) kairiensis (holotype, CAS).

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O’Hara

Figs. 145-148. Lateral and ventral views of female genitalia. Scale bars = 0.2mm: 145, Goniocera io (Y63,

CNC); 146, Proceromyia macronychia (Y134, CNC); 147, Entomophaga nigrohalterata (Y135, CNC) ( ant

apod, anterior apodeme of sternum 7); 148, Ceromya silacea (Y137, CNC).

Systematics of the Genus Group Taxa of the Siphonini

155

149. Ceromya flaviceps

151. Actia interrupta

Figs. 149-152. Lateral and ventral views of female genitalia. Scale bars = 0.2mm: 149, Ceromya flaviceps

(Y133, CNC); 150, Ceromya Ontario (Y65, USNM); 151, Actia interrupta (Y74, JEOH); 152, Actia Zaire

sp. 1 (Y 105, CNC).

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O’Hara

Figs. 153-156. Lateral and ventral views of female genitalia. Scale bars = 0.2mm: 153, Peribaea fissicornis

(Y60, JEOH); 154, Peribaea Africa sp. 5 (Y96, USNM); 155, Peribaea ugandana (Y93, USNM); 156,

Peribaea Australia sp. 1 (Y84, DPI).

Systematics of the Genus Group Taxa of the Siphonini

157

162. S. ( Pseudosiphona) sp. 14 163. S. ( Siphona) lulea

Fig. 157. Lateral view of first instar of Ceromya americana complex, with ventral view of abdominal

segments 6 to 8 (Y73, UCB). Scale bar = 0.1mm. Figs. 158-163. Lateral view of cephalopharyngeal

skeleton and ventral view of abdominal segments 6 to 8 of first instars. Scale bars = 0.05mm: 158,

Goniocera io (CNC slide) ( d corn, dorsal cornu; hypo scl, hypopharyngeal sclerite; lat scl, lateral sclerite;

lab, labrum; v corn, ventral comu); 159, Ceromya silacea (Y137, CNC); 160, Actia lamia (Y75, CNC); 161,

Peribaea ugandana (Y93, USNM); 162, Siphona (Pseudosiphona) sp. 14 (B93, CNC); 163, Siphona

(Siphona) lutea (W44, CNC).

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O’Hara

HOSTS

Introduction

Hosts are recorded for over 60 siphonine species, or about 20% of the described

fauna, in Table 2. It is difficult to speculate about host preferences from such

meagre information, though a few general patterns are suggested by this compilation

of host records. Most apparent is that siphonines are primarily parasitoids of larval

Lepidoptera, as host species are known from only three families in other orders, the

Pyrrhocoridae (Hemiptera), Tenthredinidae (Hymenoptera) and Tipulidae (Diptera).

Within the Lepidoptera there is marked preference for the Macrolepidoptera,

with only Peribaea and Actia known with certainty from hosts in both the

Microlepidoptera and Macrolepidoptera. Within the Macrolepidoptera the

Geometridae are parasitized by species in the most supraspecific taxa (nine),

followed closely by the Noctuidae (seven). Actia seems the most generalized

siphonine group in terms of host preference, with records from nine families of

Microlepidoptera and six families of Macrolepidoptera, though this may in part be

attributable to the large number of hosts known for this genus.

A brief section about hosts follows each generic or subgeneric description in the

Classification chapter of this work. Because no striking patterns of host preference

are evident between siphonine groups, and hosts are still inadequately known, host

information is not used in the phylogenetic analysis of the Siphonini.

Explanation of parasite-host list

Table 2 is a compilation of the known hosts of siphonine species. Almost all

records have been taken from the literature, though a few are new. New records

were obtained from label data and are accompanied in Table 2 by the museum

where specimens bearing the listed host data are deposited.

Siphonine species names are listed in bold face and their hosts in italics. The

former are listed alphabetically by genus and species and the latter first by family

and second by genus and species (thereby grouping together host species belonging

to the same family). Siphonine names are those used herein, and an attempt has been

made to update host names to correspond with current usage.

The literature includes many host records for tachinids that are unreliable

because of questionable species identifications. The records used here are mostly

from critically compiled works published by tachinid specialists, and as such are

more reliable than the primary literature (though even these authors undoubtedly

repeat some published errors because of the paucity of information about most

siphonine species).

Where records or species identifications were listed as questionable in a

reference cited here, I have preceded the questionable name with a question mark.

Systematics of the Genus Group Taxa of the Siphonini

159

Table 2. Parasite-host list for world Siphonini

(Continued on next page)

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O’Hara

Table 2 (continued)

(Continued on next page)

Systematics of the Genus Group Taxa of the Siphonini

161

Table 2 (continued)

SIPHONINE & HOST SPECIES FAMILY REFERENCE

(Continued on next page)

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O’Hara

Table 2 (continued)

SIPHONINE & HOST SPECIES FAMILY REFERENCE

(Continued on next page)

Systematics of the Genus Group Taxa of the Siphonini

163

Table 2 (continued)

(Continued on next page)

Quae st. Ent., 1989, 25 (1,2)

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O’Hara

Table 2 (continued)

SIPHONINE & HOST SPECIES FAMILY REFERENCE

(Continued on next page)

Systematics of the Genus Group Taxa of the Siphonini

165

Table 2 (continued)

(Continued on next page)

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O’Hara

Table 2 (continued)

(Continued on next page)

Systematics of the Genus Group Taxa of the Siphonini

167

Table 2 (continued)

Quae st. Ent., 1989, 25 (1,2)

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O’Hara

EVOLUTION OF THE SIPHONINI

Introduction

Many methods are presently available for inferring phylogenies through cladistic

analyses, but most are theoretically similar in that they begin with polarization of

character states into plesiotypic (ancestral) and apotypic (derived), generally based

on an outgroup comparison. Taxa are then ordered into a hierarchy of nested sets (a

cladogram) on the basis of synapotypic (shared derived) states. General methods of

phylogenetic analysis are described by Wiley (1981) and are under constant review

and refinement in the pages of Systematic Zoology.

Polarization of character states in the Siphonini is not possible for many of the

characters given in the Structural Features chapter. One of the problems is the

uncertainty regarding the sister group to the Siphonini. This problem required that

the outgroup comprise a selection of species from several tachinid tribes of possible

close relationship to the Siphonini. Names of species selected for this purpose are

listed in Table 3 along with their geographic distribution. Such an extended

outgroup was useful for detecting and dismissing many very homoplastic states

from the phylogenetic analysis, but also required the rejection of possibly useful

states which might have been phylogenetically interpretable if tribal relationships

between the Siphonini and the extended outgroup were better resolved. A

preliminary analysis of the phylogenetics of the Siphonini established Siphona s.l. as

a monophyletic clade within the tribe, so non -Siphona s.l. siphonines were

considered the functional outgroup (Watrous and Wheeler 1981) for polarization of

states among Siphona s.l. subgenera.

Character weighting is a controversial subject because any system of weighting

is subjective in practice, yet equal weighting of all apotypic states is even more

undesirable because a simple tally of apotypic states does not recognize the

fundamental value of evolutionary novelty and complexity in phylogenetic

reconstruction.

A very simple approach to character weighting has been adopted here. Two

levels of synapotypy are recognized - primary and secondary. A primary synapotypy

is one founded on state complexity and a minimum of homoplasy within both

outgroup and the Siphonini. A secondary synapotypy is usually structurally less

complex and/or more homoplastic than a primary synapotypy, or the character has

been insufficiently studied for the apotypic state to be hypothesized as a primary

synapotypy at this time.

Primary and secondary synapotypies of the Siphonini provide good evidence for

the monophyly of most siphonine genera and subgenera but resolve few

intergeneric, or Siphona s.l. intersubgeneric, relationships (Fig. 164).

The synapotypies for siphonine genera and subgenera are discussed under the

Phylogenetics section of each taxon in the Classification chapter. They are

summarized below and designated as primary or secondary, and incidents of

homoplasy are noted. Many characters of the Structural Features chapter are deemed

Systematics of the Genus Group Taxa of the Siphonini

169

unsuitable for phylogenetic analysis so the numbers below do not correspond to

those in that chapter. Similarly, certain states are defined differently below than in

the Structural Features chapter so a direct correlation is not possible between the

states in both sections. Characters and states of the Structural Features chapter that

are the same as those discussed here are indicated below in square brackets. The

three species groups of undescribed Siphona s.l. are considered inadequately studied

to include under the discussion of primary and secondary synapotypies.

Intergeneric relationships of non -Siphona s.l. siphonines and intersubgeneric

relationships of Siphona s.l. are not interpretable by primary or secondary

synapotypies except for the Proceromyia-Entomophaga lineage. However, the

distribution of certain, less reliable, states suggest possible higher relationships

within these groups, and these possibilities are explored at the end of this section

and depicted in Figs. 166-171.

Monophyly of the Siphonini

IP. The best supported synapotypy of the Siphonini is the presence of only two

spermathecae in the reproductive system of adult females. This synapotypy was

proposed by Andersen (1983) in his revision of the Old World Siphonini, and is

supported here by its universality among all female siphonines dissected during this

study (comprising approximately 90 species, or almost 25% of the known described

and undescribed siphonine species). Only species of the unrelated Neotropical genus

Phaenopsis Townsend are known to share this state within the Tachinidae

(Andersen 1983).

2P. The pregonite of adult male siphonines is membranous anteriorly from its apex

to its basal articulation with the hypandrium (Figs. 71-106; universal among

dissected males of over 250 siphonine species). According to Tschorsnig (1985 and

pers. comm.), this state is unique to the Siphonini, with other tachinids lacking

membrane in this position. Based on Tschorsnig’s findings, the state in the

Siphonini is tentatively proposed as a synapotypy of the tribe.

3S. Possibly synapotypic of the Siphonini is absence of accessory glands from the

reproductive system of adult males. This synapotypy was suggested by Andersen

(1983), based on examination of dissected males of a limited number of Old World

siphonine species. Andersen noted that male accessory glands are absent from

several other, apparently unrelated, tachinids. This synapotypy is in need of

corroboration because it is based on few representatives of the Siphonini and was

not investigated during this study.

Andersen (1983) also proposed that convergent subapical setae on the scutellum

are synapotypic of the Siphonini. This state is shared with possible sister groups to

the Siphonini (in particular some species of the Leskiini and Clausicellini) and

therefore cannot be defended as synapotypic of the tribe at this time.

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O’Hara

Book Review

Table 3. Tachinid species examined for outgroup comparison with the Siphonini.

Systematics of the Genus Group Taxa of the Siphonini

171

Monophyly of genera and subgenera

Synapotypies of the genera and subgenera of the Siphonini are summarized

below. Most are discussed in more detail in the Phylogenetics section of each genus

and subgenus.

Goniocera Brauer and Bergenstamm

4P. Goniocera species are unique among siphonines in having a row of ad setae on

the mid tibia. This state is shared with a few species in the outgroup, but most

outgroup species have one seta in this position. This latter state is probably in the

groundplan of both the outgroup and the Siphonini. [Character 24.]

5P. Known hosts are larvae of Malacosoma species (tent caterpillars), a parasitic

habit unique to this genus (host of G. montium unknown).

6S. Median lobe of male sternum 5 prominent and flattened on posteromedial

surface (Fig. 47; G. montium not examined). This state is approached in a few other

siphonines and similar in a few Ceromya s.s. species, but is probably independently

derived in Goniocera. [Character 40, state F.]

7S. Posterolateral margin of distiphallus curved inward toward midline and slightly

incised from rest of lateral surface (not visible in Fig. 107 and not examined in G.

montium). This state is only known to be shared with Ceromya flaviseta and C.

Ontario , but other characters do not support a close relationship between these

species. [Character 49, state S.]

8S. Female sternum 8 reduced in size (Fig. 145). This state is not considered a

primary synapotypy because the female genitalia of two of the four species, G.

montium and G. versicolor, were not examined. [Character 62, state R.]

9S. Male surstylus narrowly fused basally with the epandrium (Fig. 137, not

examined in G. montium). This state is common in many Ceromya species but few

other siphonines, so may be independently derived in Goniocera. [Character 52,

state F.]

10S. Male cerci sharply inflexed at midpoint and densely setose on basal half (Fig.

137, not examined in G. montium). Like the former character, this condition of the

cerci in Goniocera species is shared with many Ceromya species and a few other

siphonines. [In part, character 54, state I.]

Proceromyia Mesnil

IIP. Distiphallus in lateral view with long, broad, anteriorly spined, anterolateral

arm and short lateral projection (Figs. 108-109). Shape of the distiphallus is closely

approximated only in the hypothesized sister group, Entomophaga (Figs. 110-111).

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O’Hara

12P. Male sternum 5 (Figs. 48-49) with apical lobe slightly differentiated

[character 38, state O] and median lobe slightly sinuous and rounded [character 40,

state R].

13P. Head with narrow vertex (Mesnil and Shima 1978: 325).

14P. Large tarsal claws. This state is also present in a few Siphona s.s. species,

though certainly independently evolved in that group. [Character 26, state L.]

15S. A possible synapotypy is the bare prostemum. This state is shared with

Ceromya monstrosicornis, some specimens of Goniocera io and a few other

siphonines. Setulation on the prostemum is a difficult character to polarize because

it is obviously very homoplastic. A bare prostemum predominates in the outgroup,

but some species of the Clausicellini and Neaerini have a setulose prostemum. The

latter state is almost universal among siphonines, but particularly for two reasons -

its presence in Entomophaga (the hypothesized sister group to Proceromyia), and its

requirement of fewer reversals - a setulose prostemum is here considered the more

likely groundplan state of the Siphonini.

Entomophaga Lioy

16P. Aristomere 1 elongate, at least 1.5X longer than wide (Figs. 4-5). This state is

also independently derived in a few Siphona s.l. species. [Character 9, state L.]

17S. The general habitus of the two Entomophaga species, particularly with

respect to head features (Figs. 4-5; small eye, similar-shaped flagellomere 1)

suggests they are sister species.

18S. Female sternum 8 is absent from E. nigrohalterata (Fig. 147). If similarly

absent from the female of E. exoleta , then this condition would represent a primary

synapotypy.

Ceromya s.l. Robineau-Desvoidy

19S. It is not possible to cite a single state which is synapotypic of Ceromya s.l.

species, yet Ceromya s.s. and the C. silacea species group are probably each

monophyletic and have certain features in common which attest to their close

evolutionary grade if not strict monophyly. These include the diagnostic features of

Ceromya s.l. given in the Classification chapter as well as the following features of

the male genitalia shared by some but not all species of both subgroups (and a few

other siphonines, most notably Goniocera species (Fig. 137), the significance of

which is discussed later in this chapter): surstylus basally fused with epandrium

[character 52, state F], cerci sharply inflexed at midpoint [character 54, state I] and

densely setose on basal half (Fig. 138). All the above states are grouped together

here as a secondary synapotypy of Ceromya s.l. because they may be

Systematics of the Genus Group Taxa of the Siphonini

173

phylogenetically important even though they do not clearly establish the monophyly

of the genus.

Andersen (1983) proposed as a synapotypy of Ceromya s.l. : “Praegonite oval

with the sclerotized part more or less bilobed and recurved” (p. 12). Shape of the

sclerotized portion of the pregonite is more varied than suggested by Andersen and

is therefore not considered a synapotypy here. However, the oval shape is

attributable to the enlarged membrane anteriorly, and this is here considered a

synapotypy of Ceromya s.s.

Ceromya s.s. Robineau-Desvoidy

20P. Features of the pregonite provide the only convincing synapotypy of this

group. The membrane anteriorly on the pregonite is characteristically enlarged and

in most species spinulose (Figs. 39-40, 77-83). (The spinules are very tiny in some

species and consequently have been overlooked by previous workers.) Almost all

Ceromya s.s. species examined and lacking spinules on the pregonite are assignable

to species groups which possess them, so this state is considered that of the

groundplan of Ceromya s.s. The phylogenetic significance of similar spinules on the

pregonite of Goniocera io is discussed later in this chapter. [Character 45, state M.]

Ceromya Nepal sp. 1 and C. monstrosicornis are tentatively included in Ceromya

s.s. until their phylogenetic position is better resolved (see Phylogenetics section of

Ceromya s.s.).

Ceromya silacea species group

21P. Distiphallus with internal sclerotized structure, which is continuous with

infolded posterior margins (Fig. 112). This structure is unique to members of the C.

silacea species group.

22S. The first instar of C. silacea has a hook-like labrum (Fig. 159). This is

proposed only tentatively because first instars of other members of this group have

not been examined.

The phylogenetic history of Ceromya s.l. is not clearly understood. The present

division of the genus into Ceromya s.s. and the C. silacea species group is based on

male genitalic states, and may be incomplete until males of all described Ceromya

species are examined. It seems premature to erect a new subgenus for the C. silacea

species group until the phylogenetics of the genus are better understood.

Recognition of two subgenera must await corroboration of Ceromya s.s. and the C.

silacea species group as monophyletic lineages and sister groups.

Actia Robineau-Desvoidy

23P. Male pregonite (Figs. 41-42, 85-86) approximately J-shaped [character 44,

state J] and spinose [character 45, state S].

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O’Hara

24P. Actia species, with few exceptions, possess a row of hairs on the

katepistemum anterior to the mid coxa (Fig. 33). This state is unique to Actia

species and Entomophaga exoleta among siphonines, though is also present in some

species of the outgroup. No evidence suggests that Actia is the primitive lineage of

the Siphonini so presence of a katepistemal row of hairs is hypothesized as

synapotypic of Actia species. The same state in E. exoleta is hypothesized as

independently derived. [Character 20, state S.]

25P. An approximately V-shaped sternum 5 with an indistinct median lobe (Figs.

58-59) is apparently the groundplan state of the genus and approximated by few

other siphonines. [Character 38, state V and character 40, state U.]

26S. Almost all Actia species have two setulae on the upper portion of the

anepistemum (Fig. 31), whereas one setula is found in most other siphonines (Fig.

32) and is almost universal in the outgroup. This state is weighted as a secondary

synapotypy because it is widely distributed throughout different siphonine lineages

even though present in relatively few non -Actia species. [Character 19.]

27S. The lack of a dorsal comu in the cephalopharyngeal skeleton of first instars of

examined Actia species is certainly apotypic (Fig. 160), but the universality of this

state throughout the genus needs to be investigated (see O’Hara in press “a”).

[Character 65, state A.]

Peribaea Robineau-Desvoidy

28P. A strong, downwardly directed, proepimeral seta (Fig. 27) is shared by all

known Peribaea species and absent from other siphonines. Most neaerines and some

other tachinids also share this state, but are not closely related to Peribaea.

[Character 18, state P.]

29P. Distiphallus sclerotized posteriorly (partially unsclerotized [incised] in all but

three other examined and unrelated siphonine species). Andersen (1983) considered

Peribaea to be the most primitive lineage of the Siphonini and a posteriorly

sclerotized distiphallus both plesiotypic and the groundplan state in the tribe. My

outgroup comparison does not support this polarization (i.e. nothing comparable to

the Peribaea condition was found in the outgroup) and I place Peribaea near

Siphona based on characters discussed below. [Character 48, state E.]

30P. Females of most Peribaea species have a distinct sternum 8, though bare as

compared with haired as in other siphonines. The female ovipositor is elongate and

sternum 8 lacking in a few highly derived Peribaea species. [Character 62, state B.]

31S. First instars with narrowed, and in most species hook-like, labrum (Fig. 161).

Outgroup comparison is not helpful in the polarization of this state because both a

Systematics of the Genus Group Taxa of the Siphonini

175

hook-like and hatchet-like labrum are widely distributed throughout the Tachinidae.

Andersen (1983) interpreted Peribaea as primitive and a hook-like labrum as

plesiotypic, but this state is more likely apotypic if the present phylogenetic

placement of Peribaea is correct. [Character 64, state N.]

Siphona Meigen sensu lato

32P. First instars with from two to a row of large spinules or hooks on the

posteroventral margin of segment 6 (Figs. 162-163), with the exception of S.

( Aphantorhaphopsis ) Uganda sp. 1. Except for Ceromya Australia sp. 3,

non -Siphona species have tiny spinules or are bare in this position (Figs. 158-161).

[Character 66.]

33P. First instars, with few exceptions, with single dominant row of spinules on

the posteroventral margin of segment 7. Almost all examined first instars of

non -Siphona species have two or more rows of spinules or spines in this position

[Character 67, state S; refer to figure numbers cited therein.]

The Old World component of Siphona s.l., comprising the subgenera

Aphantorhaphopsis, Ceranthia and Siphona, was interpreted as monophyletic by

Andersen (1983) based on two hypothesized synapotypies. One was presence of an

anterior apodeme on female sternum 7, which is correctly polarized as apotypic in

the Siphonini based on outgroup comparison. However, this state is widely

distributed throughout the tribe (see Table 1) and therefore too homoplastic to

interpret as a synapotypy of Siphona s.l. [character 58, state W]. The second state

that Andersen considered synapotypic of Old World Siphona s.l. was presence of a

long lower katepistemal seta. I interpret this state differently from Andersen, as

discussed under the section on monophyly of Peribaea and Siphona s.l.

Siphona (subgenus Actinocrocuta Townsend)

34S. Pregonite (Fig. 89) very slender and elongate [character 44, state A], with

outer surface short spinose apically [character 45, state T].

35S. Distiphallus slender and parallel-sided (Figs. 122-123).

36S. Cerci in posterior view broadened to near apex (Fig. 142).

The above states are individually present in a few other siphonines but the

combination of all three is unique to this subgenus and none appear to be

synapotypic between S. ( Actinocrocuta ) and another Siphona subgenus.

Siphona (subgenus Aphantorhapha Townsend)

37S. Distiphallus tapered to a rounded or pointed tip (Fig. 124).

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O’Hara

38S. Cerci broadened at midlength in posterior view (Fig. 143). The cerci of S.

(Aphantorhapha) atoma do not share this apotypy and may be plesiotypic in this

respect, though other states support a sister group relationship between this species

and S. (Aphantorhapha) arizonica.

Siphona (subgenus Aphantorhaphopsis Townsend)

This taxon is non-monophyletic and comprises all Old World Siphona s.l.

species not assignable to S. (Siphona) or S. (Ceranthia). The included species are

too inadequately known to be reclassified into monophyletic lineages at this time.

None of the examined species appears to belong in the otherwise strictly New World

supraspecific taxa of Siphona s.l.

Siphona (subgenus Baeomyia O’Hara)

39P. Mid tibia without an ad seta near midlength (Fig. 38). Species of this

subgenus are not closely related to the few other siphonines with this state.

[Character 24.]

40P. Aristomere 1 very elongate, at least 2X longer than wide (Fig. 15). This state

is independently derived in a few other, unrelated, siphonines. [Character 9, state L.]

41P. Adults are the smallest of siphonines, with a total length of 2.0-3.0mm. Few

other siphonines are typically as small; the average length of adults in other lineages

is 3mm or longer. [Character 1.]

42P. Aristomere 3 very short (Fig. 15). [Character 11, state V.]

43S. Crossvein dm-cu far removed from wing margin (Fig. 22). This state is also

found in a few other siphonine lineages, and may not be synapotypic of S.

( Baeomyia ) species. However, the state is obviously homoplastic so is not used here

to define a higher subgroup within Siphona s.l. [Character 34.]

Siphona (subgenus Ceranthia Robineau-Desvoidy)

44P. Adults with cylindrical palpus (Fig. 16). This feature is considered

independently derived in two species of S. (Aphantorhaphopsis). [Character 14,

state C.]

45P. Pregonite with long seta posteriorly (Fig. 94). The bare state of the pregonite

in S. (Ceranthia) Mauritius sp. 1 is interpreted as autapotypic. [Character 46, state

L.]

46P. Distiphallus with short to long posterolateral arm which is distinctly incised

from rest of lateral surface (Fig. 127). The characteristic shape of the distiphallus in

S. (Ceranthia) species is at least subtlely different from other siphonines which also

Systematics of the Genus Group Taxa of the Siphonini

177

possess distinct posterolateral projections. [Character 49, states M and E.]

Siphona (subgenus Pseudosiphona Townsend)

47P. Pregonite (Fig. 95) broadened in lateral view [character 44, state W], with

spinules anterolaterally [character 45, state T] and two to several tiny setae

posteriorly [character 46, state F]. Few S. ( Pseudosiphona ) species lack one or

another of these states, but other siphonines do not possess more than one.

48P. Distiphallus triangular and anterolaterally spined (Fig. 128). This state is

interpreted as that of the groundplan in this subgenus, and the few species which

depart from it are regarded as apotypic in this respect.

49S. Labrum of first instars elongate and hook-like in most species (Fig. 162).

Species of S. (Pseudosiphona) with a more broadened labrum seem derived from

other species in which the labrum is hook-like. Shape of the labrum is a somewhat

homoplastic character so this state is ranked as a secondary synapotypy. [Character

64, state N.]

50S. The groundplan condition, and possible synapotypy of this subgenus, is

moderately lengthened labella (slightly longer than half prementum length; Fig. 17).

[Character 16, state M.]

Siphona (subgenus Siphona Meigen sensu stricto )

51P. Proboscis with elongate prementum and labella, with latter at least as long as

the former (Figs. 25-26). This state is present in all S. (Siphona) species, and

convergently present in some other siphonines. [Character 16, state L.]

52S. The male genitalia of S. (Siphona) species are remarkably homogeneous for

such a large group of species. There is relatively little variation in shapes of

pregonite (Fig. 96), surstylus and cerci (Figs. 141, 144), while differences in

distiphallus shape (Fig. 129) are more conservative than in any other large siphonine

taxon. Perhaps in combination, these features of the male genitalia can be

considered synapotypic of the subgenus, though individually none is unique.

Siphona (subgenus Siphonopsis Townsend)

53P. Distiphallus tapered to a narrow and truncate tip of varied length (Fig. 130).

This state is evidently unique to S. (Siphonopsis).

54S. Labellar length characteristically slightly less than three-fourths prementum

length (Figs. 18, 24). This length is not unique to, but is probably synapotypic of,

the subgenus. [Character 16, state E.]

Siphona (subgenus Uruactia Townsend)

Quaest. Ent., 1989, 25 (1,2)

178

O’Hara

55P. Distiphallus in profile smoothly tapered and apically pointed (Fig. 131). S.

(Aphantorhapha) species are characterized by a somewhat similar- shaped

distiphallus.

There are only two species included in S. (Uruactia), and these are easily

recognized as sister species because of several similarities, but each shared state is

too widely distributed among other siphonine species to be included here.

Monophyly of the Proceromyia Mesnil and Entomophaga Lioy lineage

56P. Distiphallus of all four described species characterized by long, spined,

anterolateral arm and incised lateral surface (Figs. 108-1 1 1). No examined outgroup

species or other siphonine has very similar distiphallus features.

57P. Pregonite smoothly curved and apically pointed, with longitudinal ridge

which is bare or spined (Figs. 73-76). A spined longitudinal ridge is also present in a

few New World Siphona s.l. species, but the shape of the pregonite is different

(Figs. 103-105) and other character states (for example, presence of a seta

posteriorly on pregonite as in Figs. 103-104) clearly indicate that these species are

not closely related to the Proceromyia-Entomophaga lineage. [Character 45, state

L.)

Monophyly of the Peribaea R.-D. and Siphona Meigen lineage

58P. Anal vein extended to wing margin at least as fold (Fig. 22). The groundplan

state for the Tachinidae and presumably the Siphonini is anal vein not extended to

wing margin. This state in three species of Actia is interpreted as independently

derived. Andersen (1983) considered Peribaea the primitive lineage of the

Siphonini and accordingly interpreted anal vein reaching wing margin as

synapotypic of the Siphonini and anal vein not reaching wing margin as an apotypic

loss in his Actia group ( Goniocera , Ceromya and Actia). [Character 33, state E.]

59P. Lower katepistemal seta subequal in length to, or longer than, upper anterior

seta (Fig. 32). The near universality of this state within this lineage compared with a

shorter lower seta in the outgroup and almost all other siphonines is good evidence

that this state is synapotypic of this lineage. Andersen (1983) also polarized a long

lower katepistemal seta as apotypic, but interpreted the apotypic state in a more

restricted sense. He considered the state in which this seta is longer than the upper

anterior seta as synapotypic of Old World Siphona s.l. (his Asiphona, Ceranthia and

Siphona ). This state is the most common among species of world Siphona s.l., but

the lower and upper anterior setae are subequal in length in species of some lineages

(Table 1). The subequal condition of these setae is found in the majority of Peribaea

species and is here interpreted as the groundplan state of the Peribaea-Siphona s.l.

lineage because a short lower seta characterizes all other siphonine lineages.

[Character 21, states E and L.]

Systematics of the Genus Group Taxa of the Siphonini

179

Phylogenetics of non-Siphona s.l. siphonine lineages

There are no convincing synapotypies among non-Siphona s.l. genera of the

Siphonini except those already discussed for the Proceromyia-Entomophaga

lineage. As equivocal as the generic relationships may be, two characters warrant

discussion because the interpretation of their states suggest different phylogenetic

scenarios concerning the evolution of Goniocera. (No phylogenetic statements about

the interrelationships of Actia, Ceromya s.l. and the Proceromyia-Entomophaga

lineage are made at this time because no shared states which are phylogenetically

interpretable were found among these taxa.)

Goniocera is well established as a monophyletic taxon of the Siphonini (Fig.

164), but its phylogenetic position within the tribe can be interpreted in several ways

depending upon the polarization of the states of two characters: (1) length of

preapical ad seta on fore tibia [character 23], and (2) presence or absence of spinules

on the membranous portion of male pregonite [character 45]. The distribution of

states for these characters among the four Goniocera species and major lineages of

the Siphonini is shown in Fig. 165, without polarization of states into plesiotypic

and apotypic. The states of three other characters important to the discussion of the

phylogenetics of Goniocera are also included in Fig. 165.

Length of the preapical ad seta on the fore tibia is a difficult character to polarize

into plesiotypic and apotypic states. Several species of the outgroup (both species of

the Emestiini, Triarthria setipennis and Neaera robertsonii; Table 3) have an ad

seta subequal in length to the preapical d seta, but in the others it is short. The

character is also varied among siphonines, as documented in Table 1 and Fig. 165,

so reversals have presumably taken place in both the outgroup and the Siphonini.

Yet a possibility is raised that needs to be investigated further: perhaps the presence

of a long preapical ad seta in some Goniocera species and the four species of

Proceromyia and Entomophaga represents a synapotypy of these three genera, with

a reversal within Goniocera and independent acquisitions of the state in two species

of Ceromya s.s.1, several Actia species and one species of S. (Aphantorhaphopsis).

This scenario is illustrated in Fig. 166. The evidence for this interpretation is very

weak, though it is worth noting that the members of these three genera are also

similar in appearance, with mostly dark coloration and non-vittate, uniformly

pruinose abdomens. Are the long preapical ad seta and similar appearance of these

taxa indicative of their common ancestry, their primitiveness among siphonines, or

the result of frequent homoplasy and hence of little value in phylogenetic

reconstruction? The evolutionary history of Goniocera is far from clear because the

states of the second character to be discussed are non-congruent with any but the last

interpretation.

7the presence of this state in two Ceromya s.s. species is discussed in the Phylogenetics section of Ceromya

s.s.

Quaest. Ent., 1989, 25 (1,2)

180

O’Hara

Considerable weight is placed here upon the presence of spinules on the

membranous portion of the male pregonite being an apotypic state [character 45,

state M]. This state is considered synapotypic of Ceromya s.s. for the reasons

discussed in the Phylogenetics section of that taxon. As shown in Fig. 165, this state

is also found in Goniocera io and apparently nowhere else in the Siphonini. The

distribution of this apotypic state can be interpreted in many ways, of which three

seem more reasonable than the rest: (1) the state arose independently (and

convergently) in G. io and Ceromya s.s. (Fig. 167), (2) the state arose once, and

Goniocera and Ceromya s.s. are sister groups (with subsequent reversals within both

taxa; Fig. 168), or (3) the state arose once, and Goniocera is a derived subgroup of

Ceromya s.s. (Fig. 169).

Under the first of the above interpretations (illustrated in Figs. 164 and 167), no

sister group relationship is indicated between Goniocera and another siphonine

taxon. Under the next two interpretations (illustrated in Figs. 168 and 169,

respectively), the apotypic state is considered synapotypic of Goniocera and

Ceromya s.s. and differ only with respect to the monophyly of Ceromya s.s.

The hypothesis of Goniocera and Ceromya s.s. sharing a common ancestry at

first seems highly improbable, for it requires loss of spinules from the pregonite of

all Goniocera species except G. io as well as loss in many Ceromya s.s. species (or

non-expression of the apotypic state if it is interpreted as an underlying

synapotypy8). There is reason to believe that the apotypic state is the groundplan

state of Ceromya s.s. (as discussed in the Phylogenetics section of that taxon), but

not convincingly (or even probably) for Goniocera. Nevertheless, the cladograms

pairing Goniocera and Ceromya s.s. (Figs. 168-169) are presented as alternative

(and testable) hypotheses because monophyly of Goniocera+Ceromya s.s. would

help explain the distribution of three other male genitalic character states. The three

characters are shape of median lobe on sternum 5, articulation between surstylus and

epandrium, and shape of cerci in profile; the distribution of their states is shown in

Fig. 165 (along with the states for the two characters discussed above). If the solid

dots are interpreted as apotypic states, then these are present in all examined

Goniocera species and at least some species of Ceromya s.s. and the C. silacea

species group. Decreasing the probability of a close relationship between Goniocera

and Ceromya is the possibility of the three states being primitive, the possibility of

homoplasy (note the presence of these states in a few other siphonines), and the fact

that the three states are not all present in any Ceromya s.s. species.

In summary, the • phylogenetic position of Goniocera among non-Siphona

siphonines remains enigmatic, with a possible close relationship being indicated

between it and the Proceromyia-Entomophaga lineage (Fig. 166) or Ceromya s.s.

(Figs. 168-169). Since both these interpretations are highly speculative, the

polychotomus (and unresolved) depiction of the origin of Goniocera in Fig. 164

underlying synapotypies are discussed in the next section.

Systematics of the Genus Group Taxa of the Siphonini

181

(and Fig. 167) is that best reflecting the current state of knowledge of this problem.

Phylogenetics of Siphona subgenera

Nine subgenera of Siphona s.l. are recognized, with all but S.

(Aphantorhaphopsis) hypothesized as monophyletic (Fig. 164). No clear

synapotypies were found between subgenera, though three derived character states

which are widely distributed among the subgenera provide some information about

possible higher groupings within the genus and weak additional support for its

monophyly. These character states, all belonging to characters of the male genitalia,

are reviewed below and their phylogenetic significance discussed.

The least informative of the three states concerns shape of the apex of the apical

lobe on abdominal sternum 5 [character 39]. As documented in Table 1, the tip of

the apical lobe is curved inward in at least some members of most Siphona lineages

(state C), yet this state is virtually absent from other siphonines (observed in only

one Ceromya s.s. species). The distribution of state C in the Siphonini suggests that

this state is apotypic within Siphona, but note that nine of the ten Siphona taxa with

state C in Table 1 also have state A (apical lobe not curved inward at apex;

plesiotypic). This character state distribution may be the result of one or more of the

following: (1) homoplasy, possibly resulting from parallel selection (as defined

below), (2) reversals to state A from groundplan (and apotypic) state C (also a type

of homoplasy, though groundplan state different from that of previous explanation),

and/or (3) an underlying synapotypy (as defined below). It is unlikely that the first

type of homoplasy alone could account for such a wide distribution of state C

among Siphona lineages. Equally unlikely is the interpretation of state C as the

groundplan state of most Siphona lineages with subsequent reversals in almost all

lineages. (The groundplan state is clearly A in at least S. (Siphona), as only three

unrelated species of the 40-odd species examined for this character possess state C.)

Saether (1979: 305) recognizes two types of parallelisms: (1) “parallelism as a

result of parallel selection influencing homologous structures” and (2) “parallelism

as the result of common inherited genetic factors including parallel mutations, i.e.

underlying synapomorphies'\ An underlying synapomorphy (or more generally,

synapotypy) has also been described by Saether as “the capacity or tendency to

develop parallel similarity” (1977: 31). I suspect that the inwardly curved apical

lobe on male sternum 5 results from one of these forms of parallelism, for neither

homoplasy nor state reversals provide as satisfactory an explanation for the presence

of both states A and C in nine of 13 Siphona taxa listed in Table 1. It is not so clear

which form of parallelism the shape of the apical lobe is governed by, yet this is the

key to its interpretation since, as Saether comments, “parallel selection is of no

greater value than convergence, while underlying synapomorphy is comparable to

true synapomorphy” (1979: 306). Perhaps there is an inherent tendency

^underlying synapotypy) among most Siphona lineages to develop an inwardly

curved apical lobe. This would explain both the near ubiquity of state C throughout

Quaest. Ent., 1989, 25 (1,2)

182

O’Hara

Siphona lineages and its virtual absence from other siphonines. Yet the character in

question is also one in which the difference between one state (A) and another (C)

involves little change, and the taxa grouped by the presence of the apotypic state

seem subjectively no more phylogenetically significant than the polychotomy

depicted in Fig. 164. Several factors, including the following, may be involved in

expression of state C, all of which contribute to my interpretation of this state as a

non-informative apotypic state: (1) state C is not complex so homoplasy may be

involved to some degree, (2) there may be a selective advantage to state C which

would therefore reduce its phylogenetic value, and (3) Siphona lineages from which

state C is unrecorded may nevertheless possess the underlying synapotypy.

Two states of the male pregonite are interpreted as apotypic within Siphona s.l.

because they are absent from all other siphonines. One is presence of spinules on the

distal portion of the pregonite [character 45, state T], which is restricted to New

World Siphona species of subgenera Actinocrocuta (Fig. 89) and Pseudosiphona

(Fig. 95), some members of New World species groups 1 (Fig. 100) and 2, and

several unplaced New World Siphona s.l. species (Table 1). The other state,

presence of a seta posteriorly on the pregonite [character 46, states T, M, L, F;

figures numbers for each state listed in Structural Features chapter], is more widely

distributed throughout Siphona s.l., and only unrecorded from subgenera Siphona

and Uruactia (Table 1).

The apotypic states of both pregonite characters are regarded as underlying

synapotypies (as defined above) because each involves a novel and complex feature

that almost certainly would not arise convergently, nor doubtfully by parallel

selection, among so many different lineages. As with the sternum 5 character, some

lineages with an apotypic state of the pregonite also have species with a plesiotypic

state, so the term underlying synapotypy is preferable to synapotypy, and arguably

more correct than postulating a synapotypic groundplan state followed by numerous

reversals. The difference in interpretation between the apotypic states of the

pregonite and state C of the apical lobe of sternum 5 is primarily due to the greater

complexity of the former, though as will become apparent other difficulties in

interpretation remain the same. Each pregonite character will be discussed in turn,

beginning with the setal character.

A seta posteriorly on the pregonite is a widely distributed, apotypic state among

Siphona s.l. lineages, though is absent from one or more species of most of these

lineages (Table 1). It is particularly evident that lineages in which most species have

a tiny seta also have species which lack it, so to some extent there seems to be a

graded potential for development of a pregonite seta among Siphona lineages (with

a large seta almost unique to, and hypothesized as a synapotypy of, S. (Ceranthia)).

One of the difficulties in interpretation of an underlying synapotypy is how to infer

its presence or absence in a taxon from which the apotypic state is unrecorded.

Certainly there is no reason to suppose that a pregonite seta is an underlying

synapotypy of any non-Siphona siphonines, but how does one interpret the absence

Systematics of the Genus Group Taxa of the Siphonini

183

of the apotypic state in subgenera Siphona and Uruactial Only two species of S.

(Uruactia) are known, and male genitalia of only two specimens were examined, so

these findings provide only weak (and inconclusive) evidence for the lack of the

underlying synapotypy in this subgenus.

The situation is different for S. (Siphona), as male genitalia of nearly 40 species

of this diverse subgenus were examined, and none possess a seta posteriorly on the

pregonite. These findings are far from conclusive, but they are suggestive of S.

(Siphona) occupying a rather primitive position among Siphona s.l. lineages. This

interpretation is neither supported nor refuted by other characters at this time. It

might be argued that a primitive position within Siphona s.l. is consistent with the

speciose nature and wide geographic distribution of the subgenus, but the

conservative features of its many members do not preclude a more recent and rapid

radiation of the taxon.

Even if one considers S. (Siphona) as a rather primitive lineage of Siphona s.l., it

is doubtfully the most primitive. That position probably belongs to certain members

of the Old World (probably polyphyletic) taxon S. ( Aphantorhaphopsis ). This

subgenus includes species with or without a pregonite seta, and it is possible that

some of the latter represent the most primitive lineages of Siphona s.l. This

interpretation is illustrated in Fig. 170, with S. (Siphona) shown in a primitive

position relative to most other subgenera, and S. (Uruactia) questionably placed

near S. (Siphona). Fig. 170 represents one of the more parsimonious interpretations

of the character under discussion, though it must be noted that it is presented as a

hypothetical model to be tested, and not as a well corroborated cladogram.

I return now to a consideration of the phylogenetic significance of the field of

spinules distally on the sclerotized portion of the pregonite of S. (Actinocrocuta) and

S. (Pseudosiphona) species, some species of New World species groups 1 and 2, and

some unplaced New World Siphona s.l. species (Table 1 - character 45, state T).

Much of the discussion above concerning the pregonite seta applies here as well, as

a field of spinules is interpreted as an underlying synapotypy of those taxa in which

the state is both present and absent. A cladogram based strictly upon the criterion of

presence of the apotypic state in at least one species of a Siphona lineage is shown

in Fig. 171, but again it must be cautioned that non-expression of this underlying

synapotypy in taxa grouped on the left side of the cladogram would lead to

misinterpretation of the phylogenetic affinities among these taxa. Of significance is

the fact that all taxa grouped by presence of the spinulose state are New World in

distribution, and there almost entirely restricted to the tropics and subtropics.

Whether or not one or more of the other New World taxa belong in this group, there

is at least evidence here for evolution of a higher clade of Siphona s.l. within the

New World tropics.

One final conclusion can be drawn from the discussion of the three genitalic

characters above: the distribution of the apotypic states is consistent with the

interpretation of Siphona s.l. as a monophyletic lineage, which is independently

Quaest. Ent., 1989, 25 (1,2)

184

O'Hara

supported by two synapotypies of first instars (states P32 and P33). Unfortunately

the distributions of the apotypic states of these three characters are too

phylogenetically equivocal to include as primary or secondary synapotypies of

higher clades of Siphona s.l.

Systematics of the Genus Group Taxa of the Siphonini

185

Fig. 164. Cladistic relationships and geographic distribution of the genera and subgenera of the Siphonini.

Primary and secondary synapotypies are denoted by the suffixes P and S respectively. The numbered

synapotypies are explained in the Evolution chapter and do not correspond to the numbers given to

characters in the Structural Features chapter.

Quaest. Ent., 1989, 25 (1,2)

Goniocera schistacea

186

O’Hara

8 8

O O

o g

6 6

•- JS

6 4 — G

a — n

o — o — 6 — Q 6 — 4

o — 0 — 4 — o — 0 — 4 — 4

< > — n — o — 6 — 6—4 — 6 — Q — 6 — 4

6 — 6 — 6 — 6 — 6 — 6 — 4

Characters and unpolarized states

Length of preapical ad seta on fore tibia [character 23]

O short [state S]

• long [slate E]

long seta in C. Australia sp. 3 and C. Nepal sp. 1

©' long seta in several Actia species

Q ! long seta in one S. ( Aphanlorhaphopsis ) species

Spinules on membranous portion of pregonite [character 45]

O absent

• present [state M]

Q polytypic

Shape of median lobe on male sternum 5 [character 40]

O not flattened

9 rather flattened [state I] in some species, most with state O

• prominently flattened [state F]

Articulation between surstylus and epandrium [character 52]

O membranous [state M]

• sclerotized (fused) [state F]

©’ sclerotized in one specie ‘

Shape of cerci in profile [character 54]

O not sharply inflexed

• sharply inflexed [state I]

Figure 165

Fig. 165. Distribution of states for five characters relevant to the interpretation of the cladistic relationships

of Goniocera B. & B. The male genitalia of Goniocera montium were not examined.

Systematics of the Genus Group Taxa of the Siphonini 187

Fig. 166. Long preapical ad seta on

fore tibia interpreted as synapotypic of

Goniocera and the Proceromyia-

Entomophaga lineage.

Fig. 167. Independent evolution of

spinules on pregonite of Goniocera io

and Ceromya s.s. (same as Fig. 210).

cx

00

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Fig. 168. Spinules on pregonite

interpreted as synapotypic of

Goniocera and Ceromya s.s.

Fig. 169. Spinules on pregonite

interpreted as synapotypic of

Goniocera and Ceromya s.s., with the

former a derived subgroup of the latter.

Figs. 166-169. Varied interpretations of the cladistic relationships of Goniocera B. & B. See Fig. 165 for

explanation of character states and text for discussion. Open circles indicate plesiotypic states, solid dots

indicate apotypic states, and half solid dots indicate the presence of both states within a taxon: 166, cladistic

relationship of Goniocera based on long preapical ad seta on fore tibia as an apotypic state; 167, cladistic

relationship of Goniocera based on independent evolution of spinules on the male pregonite of Goniocera io

and Ceromya s.s. species; 168, Goniocera and Ceromya s.s. as sister groups based on spinules on male

pregonite as synapotypy; 169, Goniocera as derived subgroup of Ceromya s.s., based on one interpretation

of spinules on male pregonite as synapotypy.

Quaest. Ent., 1989, 25 (1,2)

188

O’Hara

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170, One interpretation of the cladistic relationships within Siphona s.l., based on presence of a seta

posteriorly on male pregonite as synapotypy. 171, One interpretation of the cladistic relationships within

Siphona s.l., based on presence of spinules on the sclerotized portion of male pregonite as synapotypy.

Systematics of the Genus Group Taxa of the Siphonini

189

Historical zoogeography

The Siphonini are not ideal for zoogeographic analysis because they are

unknown from fossils, their sister group has not been determined and their

phylogenetic history is not well resolved. It is therefore necessary to take a general

approach to interpreting their zoogeographic history. Accordingly, I draw very

heavily upon the studies of others concerning both physical and biotic aspects of

earth history to help infer a little about how siphonines came to be distributed as

they are. In this section emphasis is placed on interpreting the geographic history of

New World siphonines.

The geographic distribution of the major lineages of the Siphonini are detailed in

the species lists and Geographic Distribution sections of the Classification chapter,

and summarized by region in Fig. 164. Eight patterns of distribution are evident

among these lineages:

(1) cosmopolitan — Ceromya s.s., Actia and S. (Siphona).

(2) widespread Old World-Nearctic — S. (Ceranthia).

(3) widespread Old World — Ceromya silacea species group, Peribaea and S.

( Aphantorhaphopsis ) (the latter is polyphyletic or paraphyletic with respect to other

Siphona s.l. species, but is apparently without close affinities with New World

Siphona s.l. species).

(4) Holarctic — Goniocera.

(5) Palearctic — Proceromyia and Entomophaga.

(6) Nearctic — S. (Baeomyia).

(7) Nearctic-Neotropical — S. (Aphantorhapha), S. ( P seudo siphona ), S.

(Siphonopsis) and New World Siphona species groups 2 and 3.

(8) Neotropical — S. (Actinocrocuta), S. (Uruactia) and New World Siphona

species group 1 .

Absent from the above patterns is any taxon of widely, and strictly, Southern

Hemisphere distribution. Similarly, no such pattern is evident among species groups

of the three cosmopolitan taxa. This negative evidence suggests that the Siphonini

did not diversify in the Southern Hemisphere until after the breakup of

Gondwanaland, either because they were initially of Laurasian distribution or they

evolved somewhere in the Southern Hemisphere after South America, Africa and

Australia drifted apart.

Breakup of Gondwanaland began with the separation and northward movements

of Africa and India from the other southern continents, followed by separation of

South America and then Australia from Antarctica. Timing of each of these events is

inexactly known, though most workers agree that South America and Africa were in

close proximity until the Late Cretaceous, and South America and Australia were

narrowly jointed via Antarctica (or only narrowly separated) into the Early Tertiary

(Adams 1981, Smith et al. 1981). Not only is the geologic timing of these events

important to the zoogeographer, but also are the varied dispersal abilities of different

organisms and the changing environmental conditions of the Cenozoic. All of these

Quaest. Ent., 1989, 25 (1,2)

190

O’Hara

factors affected the probability of particular organisms reaching new habitations.

Siphonines are moderately good dispersers, as judged from their limited species

numbers on Madagascar and islands of the Caribbean and southeast Asia, and their

virtual absence from mid-oceanic islands. This ability to cross moderate water gaps

implies that siphonines cross barriers more readily than most terrestrial vertebrates,

and can doubtfully cross such large distances as those now between South America,

Africa and Australia.

I infer from the lack of a Gondwanaland distribution pattern among siphonines,

the timing of the breakup of Gondwanaland and the dispersal powers of siphonines,

that the history of the Siphonini in the Southern Hemisphere is entirely Cenozoic in

age. This age could be extended to the Late Cretaceous if southern continents were

farther apart at the beginning of the Cenozoic (or siphonines poorer dispersers) than

here suggested.

I noted above that siphonines might have originated in Laurasia prior to the

breakup of Gondwanaland (i.e. during the Cretaceous). This possibility cannot be

corroborated or rejected by an analysis of siphonine relationships and present day

distributions (Fig. 164). Instead, such an early age for the Siphonini is considered

highly unlikely because fossil Calyptratae are virtually unknown from pre-Tertiary

deposits, and extant calyptrates show few and suspect Gondwanian distributions

(O’Hara 1983a: 329).

In summary, no evidence supports a pre-Tertiary origin of the Siphonini.

Therefore, in the ensuing discussion, all aspects of the zoogeographic history of the

Siphonini are couched in terms of Cenozoic events.

Large and small scale Cenozoic changes in earth climate, sea level, biotic

associations and position and topography of continents must have produced a

shifting array of physical and ecological barriers to the dispersal of siphonines. The

interpretation of siphonine distribution patterns is inferred from a comparison of

siphonine distribution (and hypothesized relationships) with major changes in earth

history. As an introduction to the analyses of siphonine distributions, some of the

major physical and ecological changes that are thought to have occurred during the

Cenozoic are outlined. Most of this account is summarized from Adams (1981),

with additions as noted.

Europe and Asia became separated by the Turgai Strait during Early Tertiary

time, while the Tethys Sea slowly narrowed as Africa and India drifted northward.

South America, severed from Africa in the Cretaceous and from Antarctica in the

Late Paleogene, remained isolated from other land masses until the Pliocene. North

America was in land contact with both Europe and Asia in the Early Tertiary. Two

or three land bridges spanned the North Atlantic during the Paleogene, providing

good avenues for biotic exchange between North America and Europe during most

of this period. Though the histories of these land bridges are geologically complex

and difficult to interpret, remnants of a North Atlantic corridor might have persisted

even into the Miocene (Matthews 1979, Eldholm and Thiede 1980, McKenna 1983).

Systematics of the Genus Group Taxa of the Siphonini

191

However, scanty fossil evidence of a functional Miocene corridor implies that it

was, at best, a weak filter bridge at that time (also, Beringian interchange at that

time could have produced similar fossil distributions).

Beringia probably provided a land corridor between Asia and North America

throughout most of the Cenozoic until periodically submerged during the

Quaternary (McKenna 1983). However, Beringia occupied a more northerly

position with respect to the earth’s rotational pole during the Early Tertiary, so

probably experienced a more extreme climate at that time than its North Atlantic

counterpart. Hence, Beringia might have functioned more as a filter bridge than a

land bridge in the Early Tertiary before assuming a relatively more southern position

(McKenna 1983).

The climate during the Paleogene was considerably warmer than now even at

high latitudes, peaking during the Early Eocene after several warm-cool fluctuations

(Wolfe 1978, 1980, Axelrod 1983, Romero 1986). By Middle Eocene the earth’s

climate had begun to deteriorate, causing retreat of paratropical forests ( sensu Wolfe

1978) from high latitudes and their replacement with more temperate adapted

elements. By the end of the Paleogene a seaway opened between Antarctica and

Australia, and the Turgai Straits closed between Europe and Asia.

Climatic cooling continued, with periodic fluctuations, in the Neogene. Faunal

exchange increased between Africa and Eurasia as the Tethys Sea narrowed, though

changes in sea level alternately facilitated and hindered such exchange, as it did also

as the Australian plate converged upon the island archipelago of southeast Asia

(Martin 1982). Large scale climatic and biotic changes took place as the Tethys Sea

closed and major mountain building occurred, particularly between Asia and the

Indian subcontinent and along the western edge of the Americas. Antarctica grew

progressively colder, with development of a polar ice cap in the Late Miocene

(Mercer and Sutter 1982). Marked increase in exchange between North and South

America began in the Pliocene with the emergence of a land corridor between these

continents, which has persisted under changing ecological regimes to the present

day. Most dramatic of all were the wide scale changes to the earth’s climate and

biota during the ice ages of the Pleistocene.

The following discussion about the zoogeographic history of the Siphonini is

partly based on three important assumptions: (1) siphonine evolution and

diversification took place under the general Cenozoic conditions just described, (2)

siphonines were more likely to shift geographically than ecologically in the face of

changing climatic conditions (in the same way as most species are thought to have

responded to Pleistocene climatic changes, for example), and (3) general patterns of

siphonine distribution have not been profoundly limited by host availability (one

might expect that siphonines, as parasitoids, have been constrained in their ability to

diversify in newly invaded regions by lack of suitable hosts, yet no such constraints

appear to have affected siphonine distributions).

Quaest. Ent., 1989, 25 (1,2)

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O’Hara

Two patterns of distribution are recognized among strictly Old World siphonines

- one Palearctic and the other widespread (see list of patterns at beginning of

section). The Palearctic pattern is shared by sister genera Proceromyia (with two

eastern species) and Entomophaga (with two western species). A past vicariance of

a widespread ancestor into an eastern Proceromyia and western Entomophaga

lineage is the most obvious explanation for this pattern. However, these lineages

may be relatively basal to siphonine phylogeny, so might once have been more

diverse (and possibly more widespread) than at present. This might also account for

the marked external differences among the extant species. On the other hand, the

restriction of extant species to the Palearctic region, and a temperate environment, is

suggestive of a more recent origin (perhaps during the Miocene when temperate

forests reached their maximum extent; Graham 1972, Axelrod 1983).

Belonging to the second Old World distribution pattern are Peribaea, the

Ceromya silacea species group and S. (Aphantorhaphopsis). Peribaea is widespread

throughout all Old World regions, with more described species in the Australian

region than any other siphonine lineage (partly, but not entirely, a reflection of

recent taxonomic work by Shima 1970a). Peribaea species are also recorded from

such islands as Fiji, Seychelles, Mauritius and Madagascar, so it is paradoxical that

this highly vagile and speciose group has not reached the New World.

Little can be written about the distributions of the Ceromya silacea species group

and S. (Aphantorhaphopsis) except that they, like other widespread taxa except

Peribaea , have depauperate faunas in the Australian region. This pattern

undoubtedly reflects the isolation of Australia, New Guinea and associated islands

from Asia during most of the Cenozoic.

Two distribution patterns, those of Goniocera and S. ( Ceranthia ), have

Nearctic-Old World elements. The former is known from four temperate species,

one in eastern North America and three in Europe. The phylogenetic relationships

among these species are unknown, though the presence of possible sister groups to

Goniocera in the Old World suggests that the genus arose there. Later dispersal to,

and differentiation in, the New World probably occurred when temperate forests

were widespread at high latitudes and continuous between North America and

Eurasia. Such conditions developed during the Early Neogene, presumably after

severance of a functional North Atlantic land bridge (though Axelrod (1983)

hypothesizes exchange of temperate forest elements across both Beringia and a

North Atlantic corridor in the Miocene). Therefore, barring long range dispersal

from Europe, entry to the New World was probably through Beringia. The restricted

ranges of the extant species, all far removed from Beringia yet cool adapted, are best

accounted for by either an Early Neogene vicariance of a Holarctic species (with

changes in range due to Miocene mountain building and global climatic changes), or

a more recent vicariance followed by Pleistocene displacements (Fig. 172).

The subgenus S. (Ceranthia) also has a Nearctic-Old World distribution pattern.

Old World species of this subgenus are almost equally split in number between the

Systematics of the Genus Group Taxa of the Siphonini

193

Palearctic and Afrotropical regions, while the Nearctic element is strictly temperate.

As with Goniocera, the temperate element of S. (Ceranthia) in the Nearctic region

probably resulted from dispersal from the Old World during the Neogene (Fig. 172),

possibly during the Miocene judging from the moderate number of Nearctic species

(mostly undescribed).

Among the remaining distribution patterns are four Siphona lineages only

recorded from either the Nearctic ( S . (Baeomyia)) or Neotropical ( S .

(Actinocrocuta), S. (Uruactia) and New World Siphona species group 1) region. In

patterns such as these where all members of a lineage are within one region, it is

most parsimonious to postulate that each lineage evolved from an ancestral species

in that region. This is almost certainly true of the three Neotropical lineages, since S.

( Uruactia ) is only known from two species in Peru and Ecuador, and S.

( Actinocrocuta ) and N.W. Siphona sp. grp. 1 are members of a monophyletic group

(Fig. 171) of almost entirely Neotropical distribution. The history of Neotropical

siphonines is discussed in more detail further on.

Three siphonine lineages have cosmopolitan distributions: Ceromya s.s., Actia

and S. (Siphona). The zoogeographic history of S. (Siphona), with special emphasis

on the North American fauna, was treated elsewhere (O’Hara 1983a). I

hypothesized that periodic exchange between the Old and New World took place

among S. (Siphona) species groups from the Eocene to the Pleistocene. In general,

Holarctic elements were hypothesized as Middle Tertiary to Quaternary in age (Fig.

172) and more tropically distributed taxa with Old World-New World affinities of

older ancestry (Fig. 173). I predict that similar hypotheses can explain the

distributions of Ceromya s.s. and Actia species groups when these are

phylogenetically analyzed.

It would appear from lists of described species that cosmopolitan lineages of the

Siphonini are low in species diversity in both the Neotropical and Australian

regions. Though this is so for the Australian region, it is not for the Neotropical

region. The pattern of low diversity in the Australian region is the same as for most

Old World siphonine lineages with Australian members, reflecting the relative

isolation of that region from Asia for most of the Cenozoic (see above). South

America was also isolated from other continents for most of the Cenozoic, so the

high diversity of siphonines in that region poses an especially interesting

zoogeographic problem.

The Neotropical region has three endemic siphonine taxa (S. (Actinocrocuta), S.

(Uruactia) and New World Siphona species group 1), five lineages shared only with

the Nearctic region (S. (Aphantorhapha), S. (Pseudosiphona), S. (Siphonopsis) and

New World Siphona species groups 2 and 3), and elements of three cosmopolitan

groups ( Ceromya s.s., Actia and S. (Siphona)). How can this high diversity of

Neotropical siphonines be explained? Whether the ancestors of most Neotropical

lineages originated in that region or in the Old World is of minor importance, as the

major pathways of movement ( i.e . between the Americas and between North

Quaest. Ent., 1989, 25 (1,2)

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O’Hara

America and Eurasia) would be the same even if the direction were reversed. The

phylogenetic relationships are presently too inadequately known in Ceromya s.s.,

Actia or S. (Siphona) to strongly argue for or against an Old World origin of each

(cf. O’Hara 1983a, in which an Old World origin was favored partly because the

high diversity of these taxa in the Neotropical region was unknown). However, the

possibly basal lineages of the Siphonini are Old World in distribution, so for the

purpose of this discussion an Old World origin will be assumed for these three taxa.

Certain lineages of Siphona s.l. are certainly New World in origin, as the

distributions in Fig. 164 and possible relationships in Fig. 171 illustrate.

It follows from the review above about Cenozoic events and general assumptions

about siphonine evolution, that the only route to (or from) South America during the

Cenozoic would have been through North America. It is well known on the basis of

both geologic and fossil evidence that South America was physically isolated from

other continents during most of the Cenozoic. However, there were two periods of

faunal and floral exchange with North America. The first is hypothesized as

Paleocene (and Late Cretaceous), as evidenced and/or postulated from geologic

evidence (Coney 1982) and distributions of angiosperms (Gentry 1982),

herpetofauna (Savage 1982, Estes and Baez 1985), atherinid fishes (White 1985),

colubrid snakes (Cadle 1985) and mammals (Gingerich 1985, Webb 1985a). The

exact nature of the Paleocene connection is uncertain, though there is general

agreement that it was probably an island chain and functioned as a filter bridge.

Certain organisms evidently crossed this bridge more readily than others. The

suggestion that this filter bridge persisted into the Eocene is not well documented.

The second period of biotic exchange between the Americas is very well

corroborated and the subject of a recent multi-authored book, “The great American

biotic interchange” (Stehli and Webb 1985). This interchange began in the Pliocene

(about 3mybp [million years before present] according to most sources, but 5mybp

according to others) with the emergence of a complete land corridor between the

Americas. This corridor has alternately functioned, from that time to the present, as

a route of dispersal for certain organisms and a barrier to others, as global climate

fluctuated throughout the ice ages of the Pleistocene.

It is extremely doubtful that the diversity of Neotropical Siphonini can be

entirely accounted for by immigrations since the Pliocene, though it is clear that

present day siphonine sister species and sister groups distributed between Middle

and South America are evidence of siphonine participation in the “great biotic

interchange”. The important question to be addressed here is whether earlier

siphonine arrivals to South America were over a water gap or across a filter or land

bridge.

Assuming (as above) that siphonines originated in the Old World during the

Cenozoic and have generally maintained their ecological affinities during periods of

major climatic change, it follows that ancestors of extant Neotropical siphonines

entered North America from Eurasia at a time during the Tertiary when land bridges

Systematics of the Genus Group Taxa of the Siphonini

195

between the continents were subtropical to tropical in nature. As the only such land

bridges were at high latitudes (Beringia and across the North Atlantic), and the

climatic optimum of the Early Eocene was the only time during the Tertiary that

these corridors supported such conditions, it is hypothesized that an ancestral stock

of Neotropical Siphonini entered the New World at that time. (Webb, comparing

mammal faunas of Europe and North America, called the warm period of the Early

Eocene “the most intensive intercontinental interchange between any two

continental mammal faunas before the Great American Interchange.” (1985a: 207))

Then, as global climate cooled in the latter part of the Paleogene and temperate

forests began to replace the retreating tropical forests, siphonines became

increasingly more southerly distributed in North America (Fig. 173). This sequence

of events was proposed to explain Mexican elements of S. (Siphona) (O’Hara

1983a), and to explain some elements of the Central American herpetofauna

(Savage 1982).

The zoogeographic scenario has thus far tended toward a vicariant explanation of

siphonine history. However, only dispersal events over a wide water gap or across a

weak filter bridge can account for the earliest siphonine entries into South America

(Fig 173). Even if siphonines colonized North America in the Eocene and reached

the southern part of the continent shortly thereafter, they would still have arrived too

late to cross the Paleocene filter bridge into South America. There is no evidence

from the Eocene to the Miocene of a land corridor between the Americas, and little

evidence for an even weak filter bridge until the Late Miocene. (An intermittent

filter bridge is mostly hypothesized to explain the first appearance of certain

organisms in the fossil record of either North or South America - e.g. Stehli and

Webb 1985, Chapter 1.) Whether or not a filter bridge existed from time to time

during this interval, there is nevertheless a suggestion of faunal and floral exchange,

as inferred from cricetine rodents (Hershkovitz 1966, 1972), angiosperms (Raven

and Axelrod 1975), carabid beetles (Ball 1978, Allen and Ball 1980, Ball and

Shpeley 1983, Noonan 1985), mammals (Webb 1985b), ectoparasites of mammals

(Wenzel and Tipton 1966), primates and caviomorph rodents (Wood 1985 [though

the zoogeographic interpretations of these groups are highly controversial]) and

herpetofauna (Vanzolini and Heyer 1985). Siphonines, like these taxa, seem to have

preceded, as well as participated in, the “great biotic interchange” in their

colonization of South America. When, and how often, is uncertain. The large clade

of Neotropical Siphona s.l. species (Fig. 171) suggests at least a moderate history in

South America (Oligocene or Miocene?). The high species diversity of Ceromya

s.s. , Actia and S. (Siphona) in the Neotropics also suggests that members of these

lineages reached South America before formation of the Pliocene land bridge (Fig.

173).

A minor element of the Nearctic siphonine fauna is possibly derived from

tropical adapted siphonines. Two sources are hypothesized: Firstly, differentiation

of temperate adapted taxa from tropical adapted taxa as subtropical to tropical

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O’Hara

conditions over most of the region were replaced by warm temperate to boreal

conditions after the climatic deterioration of the Eocene. (Similar to the hypothesis

that the vegetation of the Sonoran Desert is mostly derived from contiguous, less

arid adapted, taxa; Axelrod 1979.) It has been assumed throughout this section that

the evolution of new ecological affinities such as this have played a minor role in

the history of New World Siphonini - an analysis of relationships among North

American S. (Siphona) suggests this is a valid assumption (O’Hara 1983a).

Secondly, a few Nearctic taxa are likely derived from reinvasion of the region from

Middle or South America (Fig. 172). Most of these taxa are southerly distributed in

the Nearctic region.

Perhaps as more organisms are studied, particularly vagile organisms like many

insects, vicariance biogeographers will recognize biotic exchange between North

and South America as a zoogeographic problem demanding of both dispersal and

vicariant explanations. Cracraft (1975), for instance, considered the patterns of

distribution between the Americas as inadequately interpretable by vicariant events

alone. It must be realized that in certain areas dispersal cannot be ignored in favor of

the “rigorous” method of vicariance biogeography. Erwin, discussing how

vicariance biogeographers interpret distributional data, wrote:

“The major flaw of the entire Croizat school [vicariance biogeography] is

that their method is no more “general” [meaning “vicariance ... is to be

assumed and dispersal only explains special cases”] than that of the

dispersal or centrist’s school, because biological organisms are not

generalists! All groups and subsets of groups have their own unique

powers of movement or lack of movement and each group must be dealt

with on its own merits with full knowledge of powers of dispersal,

passive or active, or lack thereof.” (1979: 357)

Summary of the zoogeographic history of the Siphonini

(1) The Siphonini are hypothesized as originating during the Cenozoic, because of

the virtual absence of Mesozoic fossil Calyptratae and lack of recognizable

Gondwanian distributions among extant siphonines.

(2) The present distributions of sister genera Proceromyia and Entomophaga in

eastern and western Palearctic, respectively, suggest a past vicariance of a

widespread Palearctic ancestor. However, the possibly rather basal position of

this clade in the Siphonini might reflect an older ancestry than otherwise

suspected.

(3) A general pattern of low diversity is noted in the Australian region for most

siphonine lineages. This pattern is explained by the relative isolation of

Australia and associated islands from southeast Asia during most of the

Tertiary, which prevented rapid dispersal of siphonines into the region, and

Systematics of the Genus Group Taxa of the Siphonini

197

hence limited the opportunity for speciation therein.

(4) It is assumed from largely inferential evidence that the Siphonini originated in

the Old World, so the high diversity of New World siphonines must be

explained by northern routes of entry (across Beringia and the North Atlantic)

from Eurasia (Figs. 172-173).

(5) Most Nearctic siphonines are descendants of Old World taxa that became

widespread in the Holarctic region since the Middle Tertiary, as temperate and

boreal conditions replaced the more tropical conditions of the Eocene (Fig.

172). A minor component of the Nearctic fauna is derived from tropical adapted

siphonines. These either differentiated in situ from tropical adapted siphonines

as global climate cooled and temperate forests expanded, or reinvaded the

Nearctic region from Middle or South America (Fig. 172).

(6) It is assumed that siphonines have generally maintained their ecological

affinities through periods of climatic change. Therefore, Neotropical siphonines

are hypothesized as descendants of Old World, tropical adapted, siphonines that

became widespread in the Holarctic region during the climatic optimum of the

Eocene. Subsequent deterioration in climate and periodic crossing of a water

barrier (weak filter bridge?) between the Americas during the Middle to Late

Tertiary, and speciation in South America, accounts for the high diversity of

some siphonine lineages in the Neotropical region (particularly the large clade

of Neotropical Siphona s.l. species) (Fig. 173).

(7) Sister species and groups distributed on either side of the Isthmus of Panama

(mostly Middle America-South America) are evidence of siphonine

participation in the “great American interchange” of the Pliocene and

Quaternary.

Predictions and tests of zoogeographic hypotheses

(1) No Gondwanian distribution patterns will be recognized among siphonines.

(2) More resolved phylogenies of both higher taxa and species groups of the

Siphonini will more firmly establish the Old World origin of most lineages and

corroborate the suspected affinities between Nearctic and Palearctic Siphonini

(, i.e . similar relationships are expected to those hypothesized for S. (Siphona)',

O’Hara 1983a).

(3) The largely endemic nature of Neotropical Siphonini will become apparent as

relationships to faunas of other regions are resolved, reflecting the old origin of

the fauna from Early Eocene, North American, ancestors.

(4) Indirect corroboration will be forthcoming from similar interpretations of

Neotropical elements of similarly distributed taxa, and corroboration or

refutation of aspects of earth history herein reviewed.

Quae st. Ent., 1989, 25 (1,2)

198

O’Hara

Fig. 172. General zoogeographic history of Nearctic elements of the Siphonini. Three ancestral sources are

hypothesized as contributing to the siphonine fauna of the Nearctic region:

(1) most taxa are derived from Old World ancestors crossing Beringia from the Middle Tertiary to the

Pleistocene. These taxa evolved in association with warm temperate to boreal biomes.

(2) a minor component is derived from in situ differentiation, as tropical adapted siphonines shifted

southward, and temperate forests expanded, following the climatic deterioration of the Eocene.

(3) another minor component, mostly southern Nearctic in distribution, are taxa that reached the Nearctic

region from Middle or South America. Taxa reaching North America from South America did so either

before (very few taxa) or after a land corridor was established between these continents in the Pliocene.

Systematics of the Genus Group Taxa of the Siphonini

199

Middle Tertiary

southward

shift

Middle and Late

Tertiary over-water

dispersal

Pre-Pliocene history of

Neotropical elements of the Siphonini

Actia

Ceromya s.s.

S. ( Actinocrocuta )

S. ( Aphantorhapfa )

S. ( Pseudosiphona )

S. ( Siphona )

S. (Siphonopsis)

S. ( U ruactia )

New World Siphona sp. grp. 1

New World Siphona sp. grp. 2

New World Siphona sp. grp. 3

Fig. 173. Pre-Pliocene zoogeographic history of Neotropical Siphonini. An ancestral stock of tropical

adapted siphonines are hypothesized to have reached North America across Beringia and/or North Atlantic

land bridges during the climatic optimum of the Early Eocene. These taxa became more southerly distributed

as global climate deteriorated. In the absence of a land bridge between the Americas for most of the

Cenozoic, ancestral siphonines of several major lineages crossed a water gap (weak filter bridge?) to reach

South America. Exchange was facilitated between the Americas after establishment of a land corridor in the

Pliocene (as siphonines participated in the “great American interchange”).

Quaest. Ent., 1989, 25 (1,2)

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O’Hara

CONCLUDING REMARKS

I have set forth in this paper a revised classification of the supraspecific taxa of

the Siphonini. For the first time, the markedly different classifications of the Old

and New World Siphonini have been meshed. Relatively few taxonomic changes are

proposed as a result of this action, except for a number of changes in rank to update

and balance the classification (particularly with respect to genus group names

among New World Siphona group taxa) and several new higher level combinations

required on phylogenetic grounds.

Considerable effort was made to classify - and provide a key for identification of

- Neotropical siphonines, even though most of the known species are undescribed.

However, this effort is but a preliminary attempt, and apt to be inadequate for some

taxa. Present categories will need to be modified, and new subgenera of Siphona s.l.

will almost certainly need to be erected, when the diverse siphonine fauna of the

Neotropical region is more completely studied and the species described.

ACKNOWLEDGEMENTS

It is a pleasure to thank my supervisor, G.E. Ball, for his constant

encouragement, superior judgement and unfailing support throughout the course of

this project. His personal and professional integrity and dedication to science has

been inspirational.

I am grateful to all curators and individuals - listed in the Materials and Methods

chapter - who loaned me specimens or made collections available to me during this

study. My knowledge of the Tachinidae was much improved by frequent visits with

D.M. Wood, who also provided unlimited access to the Canadian National

Collection, gave freely of his time and his extensive knowledge of the Tachinidae,

and made many valuable suggestions for the improvement of this thesis. I am also

indebted to the following individuals for discussions about siphonine and tachinid

systematics during visits to their institutions: S. Andersen, R.W. Crosskey, B.

Herting, C.W. Sabrosky and H.P. Tschorsnig. Their hospitality during my visits is

much appreciated. The following tachinidologists were also particularly helpful

regarding loan of siphonines or other information related to my study: P.H. Amaud,

Jr., B.K. Cantrell, R. Cortes, H. Shima and N.E. Woodley.

I am grateful to the members of my Advisory Committee and the external

examiner for their critical review of my Ph.D. thesis: G.E. Ball, B.S. Heming, C.W.

Sabrosky, D.H. Vitt, M.V.H. Wilson and D.M. Wood. I also thank B.V. Brown and

D.C. Currie for their review of the zoogeography chapter. For review of this

manuscript based on that work for publication, I thank R.W. Crosskey and an

anonymous reviewer.

My ideas about systematic theory have developed over the years, influenced in

part by discussions with G.E. Ball, G.C.D. Griffiths and fellow graduate students:

Systematics of the Genus Group Taxa of the Siphonini

201

R.S. Anderson, J.S. Ashe, J.M. Gumming, D.C. Currie, G.A.P. Gibson, J.-F. Landry,

D.R. Maddison and R.E. Roughley.

G. Braybrook provided technical assistance with the Scanning Electron

Microscope, and J. Scott and K.P. Fennie assisted with the production of the

photographic plates.

Financial support for this study was provided by Natural Sciences and

Engineering Research Council of Canada Grant A- 1399, held by G.E. Ball.

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Zetterstedt, J.W. 1849. Diptera Scandinaviae. Disposita et descripta 8: 2935-3366.

Lund.

Zetterstedt, J.W. 1859. Diptera Scandinaviae. Disposita et descripta 13: Sup. 4, pp.

v-xvi, 4943-6190. Lund.

POSTSCRIPT

Several papers dealing with siphonines have recently been published while this

paper was in press. These are listed below.

O’Hara, J.E. 1988. Survey of first instars of the Siphonini (Diptera: Tachinidae).

Entomologica scandinavica 18: 367-382. This paper is cited as “In press ‘a’” in the

text and references above.

O’Hara, J.E. 1988. Correlation between wing size and position of a hind crossvein

in the Siphonini (Diptera: Tachinidae). Journal of Natural History 22: 1141-1146.

This paper is cited as “In press ‘b’” in the text and references above.

Herting, B. 1987. Beitrage zur Kenntnis der palaarktischen Raupenfliegen (Dipt.

Tachinidae), XVII. Stuttgarter Beitrage zur Naturkunde (A) 408: 1-14. A new

siphonine species, Ceranthia impropria Herting, is described in this work. Dr.

Herting kindly allowed me to examine the only known specimen of that species

when I visited him in 1985. I provisionally follow Herting ’s placement of the

species in Ceranthia ( i.e . Siphona ( Ceranthia ) in my classification); however, I need

to examine a male specimen (particularly the male genitalia) before placing the

species with confidence.

Cantrell, B.K. 1988. The comparative morphology of the male and female

postabdomen of the Australian Tachinidae (Diptera), with descriptions of some

first-instar larvae and pupae. Invertebrate Taxonomy 2: 81-221. First instars and the

male and female genitalia of some Australian siphonines are described and

illustrated.

I discuss in the text a nomenclatural problem involving the name Peribaea orbata

Systematics of the Genus Group Taxa of the Siphonini

219

(see Peribaea Section). The matter is thoroughly reviewed by R.W. Crosskey and H.

Shima (1988) in a case before the International Commission on Zoological

Nomenclature: Case 2632. Tachina orbata Wiedemann, 1830 (currently Peribaea

orbata; Insecta, Diptera) : proposed confirmation of neotype designation. Bulletin of

Zoological Nomenclature 45: 199-201.

Quaest. Ent., 1989, 25 (1,2)

220

O’Hara

INDEX TO NAMES OF TAX A

(Synonyms in italics)

FAMILY GROUP TAX A

Actiini, 34

Ceromydes, 34

Calyptratae, 5, 14

Clausicellini, 172

Crocutini, 34

Diptera, 158

Emestiini, 179

Eutachininae, 34

Geometridae, 158

Goniinae, 5

Hemiptera, 158

Hymenoptera, 158

Lasiocampidae, 43

Lepidoptera, 5

Muscidae, 34

Neaerini, 5, 14, 77, 79, 172

Noctuidae, 158

Pyrrhocoridae, 158

Siphonae, 34

Siphonina, 5, 14, 34

Siphoninae, 34

Tachinidae, 5, 178

Tachininae, 5, 34

Tenthredinidae, 158

Thryptocerides, 34

Thryptoceratae, 34

Thryptoceratidae, 34

Tipulidae, 158

GENERA AND SUBGENERA

Actia Robineau-Desvoidy, 14,

18-21,27, 36-39,41,43,49,

51-52, 58, 60-61,67-71,

80-81,84, 158, 173-174, 179,

189, 193-195

Actinactia Townsend, 53, 61

Actinocrocuta Townsend, 10-11,

85,87, 89, 175, 182-183, 189,

193

Actiopsis Townsend, 67, 72

Aphantorhapha Townsend, 10, 87,

89-91, 175, 189, 193

Aphantorhaphopsis Townsend, 10,

18,36-37,41,55,85,92,

94-95,99, 109, 175-176, 181,

189, 192

Asiphona Mesnil, 13, 36-37, 92,

95, 178

Baeomyia O'Hara, 10, 19, 37, 84,

97-98, 176, 189, 193

Borgmeiermyia Townsend, 16

Bucentes Latreille, 108, 1 13-1 15

Cartocometes Aldrich, 41, 43

Ceranthia Robineau-Desvoidy, 10,

13, 18, 23,37, 85, 93,95,

99-101, 175-176, 178, 182,

189, 192-193,218

Ceromy a Robineau-Desvoidy, 11,

14, 36-37, 46, 49, 52, 57, 59,

95, 175, 178

Ceromya Robineau-Desvoidy

sensu lato, 1 1, 38, 45, 50-53,

57, 172-173, 179

Ceromya Robineau-Desvoidy

sensu stricto, 11, 39, 46,

50-53,55-56, 58, 173, 180,

189, 193-195

Chaetostigmoptera Townsend, 77,

79

Crocuta Meigen, 108, 114

Entomophaga Lioy, 14, 36, 38, 44,

46^17, 50, 52, 58-59, 71-72,

128, 171-172, 178-179, 189,

192, 196

Eogymnophthalma Townsend, 77

Euchaetactia Villeneuve, 41, 43

Systematics of the Genus Group Taxa of the Siphonini

221

Euthryptocera Townsend, 41

Goniocera Brauer and

Bergenstamm, 14, 19, 22, 35,

38, 41 — 42, 52, 54, 58-59, 171,

178-180, 189, 192-193

Gymnopareia Brauer and

Bergenstamm, 67

Gymnophthalma Lioy, 67

Herbstia Edwards, 77

Herbstia Robineau-Desvoidy, 77

Lasioneura Coquillett, 62

Nipponoceromyia Mesnil and

Shima, 36-37, 44, 46

Peribaea Robineau-Desvoidy, 13,

18, 26, 37, 69, 77, 79, 158,

174-175, 178, 189, 192

Phaenopsis Townsend, 34, 169

Phantasiosiphona Townsend, 108

Phytomyptera Rondani, 16

Polychaetoneura Walton, 52, 60

Proceromyia Mesnil, 36-37, 39,

44-47, 49-50, 52, 59, 128,

171, 178-179, 189, 192, 196

Pseudactia Malloch, 53, 61

Pseudosiphona Townsend, 10-11,

18, 86, 103, 106-107,

120-122, 129, 177, 182-183,

189, 193

Schizactiana Curran, 53

Schizoceromyia Townsend, 53, 61

Setasiphona Townsend, 67

Shizotachina Walker, 61

Siphona Meigen, 84, 97, 108,

174-175, 178, 182

Siphona Meigen sensu lato , 10-11,

22-24, 36, 39, 77, 84, 95, 121,

125-126, 128-129, 175, 178,

181-183, 194, 197

Siphona Meigen sensu stricto,

10-11, 13, 18, 35,37, 85, 104,

108, 112, 115, 177, 183, 189,

193-196

Siphonopsis Townsend, 10-11, 18,

68, 86, 104, 112, 120-122,

129, 177, 189, 193

Stenoparia Stein, 37, 46, 58

Stomoxys Geoffroy, 1 14

Strobliomyia Townsend, 13, 37,

77.81

Tachina Meigen, 44, 50, 60-61,

66, 72-73, 75,81,89, 205

Talaractia Malloch, 77, 81

Thryptocera Macquart, 34, 60, 67,

72.81

Uruactia Townsend, 10, 12, 86,

123, 125, 177-178, 182-183,

189, 193

Uschizactia Townsend, 77

Xanthoactia Townsend, 52

SPECIES AND SUBSPECIES

abbreviata (Villeneuve), Siphona,

113

abdominalis (Robineau-Desvoidy),

Siphona, 101-102, 162

aberrans Malloch, Actia, 2

aegyptia (Villeneuve), Peribaea,

78.81

akidnomyia O'Hara, Siphona, 1 13

albocincta (Villeneuve), Siphona,

114

alipes (Villeneuve), Peribaea, 82

altemata Shima, Peribaea, 80

alticola (Mesnil), Siphona, 93, 96

amblycera (Aldrich), Ceromya, 54,

60

americana (Townsend), Ceromya,

54, 60, 159

americana Townsend,

Thryptocera, 52

amicula Mesnil, Ceromya, 60

amoena (Mesnil), Siphona, 1 14

amplicomis amplicomis Mesnil,

Quaest. Ent., 1989, 25 (1,2)

222

O’Hara

Siphona, 110

amplicomis Mesnil, Siphona, 110,

114, 117

amplicomis nigrohalterata Mesnil,

Siphona, 1 10

analis Meigen, Siphona , 1 16

analis Robineau-Desvoidy,

Siphona, 119

angusta Mesnil, Siphona, 114

angustifrons (Malloch), Siphona,

95-96

annulata (Mesnil), Peribaea, 80

anomala Zetterstedt, Siphona , 102

antennalis (Mesnil), Siphona, 114

antennalis (Rondani), Actia, 73

antennata (O'Hara), Siphona,

98-99

anthomyformis Lynch Arribalzaga,

Siphona, 1 19

anthracina Mesnil, Peribaea, 80

antiqua (Mesnil), Actia, 72

apicalis Robineau-Desvoidy,

Peribaea, 77-78, 80

apicipunctata (Malloch), Ceromya,

60, 162

argentifrons (Malloch), Peribaea,

80, 162

aristalis (Rondani), Actia , 73

arizonica (Townsend), Siphona,

90-92, 176

arizonica Townsend,

Aphantorhapha, 89

articulata (Stein), Entomophaga,

51

atoma (Reinhard), Siphona, 90-92,

176

atricapilla Mesnil, Siphona, 1 14

autumnalis (Townsend), Actia, 72

autumnalis Townsend, Actiopsis,

67

baldwini (Malloch), Peribaea, 77,

81

baldwini Malloch, Actia, 77

bellina Mesnil, Ceromya, 60

bevisi Curran, Siphona, 110, 114

bicolor (Macquart), Actia , 72, 131

bicolor (Meigen), Ceromya, 51,

54, 56-57, 59-60, 94, 131, 162

bicolor Macquart, Thryptocera, 67

bigoti (Milliere), Actia, 75

bilineata (Mesnil), Siphona, 1 14

boreata Mesnil, Siphona, 1 14

brasiliensis (Townsend), Siphona,

122-123, 162

brevirostris Coquillett, Siphona,

40, 103, 105-108, 162

brevis Malloch, Actia, 69, 72

broteas (Walker), Actia, 75

brunnea Malloch, Actia, 72

brunnea O'Hara, Siphona, 1 14

brunneipalpis (Villeneuve),

Siphona, 96

brunnescens (Villeneuve),

Siphona, 96

buccalis (Curran), Ceromya, 61

capensis Curran, Siphona, 114

capitata Mesnil, Ceromya, 61

cephalotes Mesnil, Ceromya, 39,

52, 54, 57, 61

ceres (Curran), Siphona, 115

cervina (Mesnil), Peribaea, 77, 81

chaetosa Townsend,

Actinocrocuta, 40, 87

chaetosa Townsend, Siphona,

88-89

chetoliga Rondani, Siphona, 1 14

chrysocera Bezzi, Actia, 39,

67-68, 72

cibdela (Villeneuve), Ceromya, 54,

61, 162

ciligera (Mesnil), Actia, 20, 68

cinerea (Latreille), Siphona, 1 15

cinerea Meigen, Siphona, 115

cinereus Latreille, Bucentes, 108

Systematics of the Genus Group Taxa of the Siphonini

223

clara (Mesnil), Peribaea, 81

claripennis (Robineau-Desvoidy),

Actia, 72

clausa Robineau-Desvoidy,

Siphona, 119

collini Mesnil, Siphona, 114, 162

comitata Villeneuve, Actia, 73

compacta (Curran), Peribaea, 81

completa Malloch, Actia, 39,

67-70, 72

conata (Reinhard), Siphona,

121-123, 162

confusa Mesnil, Siphona,

113-114, 162

consimilis Robineau-Desvoidy,

Siphona, 1 19

comuta (Aldrich), Ceromya, 54, 61

cothumata (Mesnil), Siphona, 1 14

crassicomis (Meigen), Actia, 72,

159

crassicomis Meigen, Tachina, 67

crassulata (Mesnil), Siphona, 93,

96

creberrima (Speiser), Siphona, 1 14

cristata (Fabricius), Siphona,

113-114, 162

cuthbertsoni Curran, Actia, 72

cuthbertsoni Curran, Siphona,

110-111, 114

darwini Malloch, Actia, 69, 72

deferens Malloch), Actia, 72

delicatula Mesnil, Siphona , 1 1 8

diffidens Curran, Actia, 72, 159

dilecta Herting, Ceromya, 61

discicomis (Pandelle), Peribaea,

78,81

dorsalis Brauer and Bergenstamm,

Siphona, 1 19

dorsigera Herting, Ceromya, 66

dubia (Malloch), Ceromya, 61

dubitata Herting, Actia, 27, 71-72

efflatouni Mesnil, Siphona, 1 15

elyii (Walton), Ceromya, 60

elyii Walton, Polychaetoneura, 52

enigmatica Villeneuve, Goniocera,

44

erythrocera Robineau-Desvoidy,

Ceromya, 62

eucosmae Bezzi, Actia, 68, 73-7 4,

159

exoleta (Meigen), Entomophaga,

18, 37,47-50, 58, 67,71, 172,

174

exoleta Meigen, Tachina, 47

exscensa (Walker), Actia, 75

exsecta Villeneuve, Actia, 68, 73

fallax (Mesnil), Actia, 20, 68, 73

fasciata (Stein), Ceromya, 60

femorata Mesnil, Ceromya, 61

fera Mesnil, Siphona, 93, 96

fergusoni (Bezzi), Ceromya, 53,

55-56,61, 162

fergusoni Bezzi, Schizotachina, 53

ferina (Mesnil), Peribaea, 8 1

fissicomis (Strobl), Peribaea, 8 1 ,

162

fissicomis Strobl, Thryptocera, 77

flaviceps (Ratzeburg), Ceromya,

54, 57, 61

flaviceps (Stein), Ceromya, 61

flavicornis Robineau-Desvoidy,

Peribaea, 83

flavifrons Staeger, Siphona, 1 15

flavipalpis (Macquart), Actia, 72

flavipes (Coquillett), Siphona, 40,

102

flavipes (Robineau-Desvoidy),

Siphona, 102

flaviseta (Villeneuve), Ceromya,

43,53,61, 171

flavisquamis

(Robineau-Desvoidy), Actia,

75

floridensis O'Hara, Siphona, 1 15

Quaest. Ent., 1989, 25 (1,2)

224

foliacea (Mesnil), Siphona, 115

frontalis (Macquart), Actia, 73

fuliginea cerina Mesnil, Siphona,

115, 180, 189

fuliginea Mesnil, Siphona, 111,

115

fuliginea rubea Mesnil, Siphona,

115

fulvicauda Malloch, Actia, 39,

67-68, 73

fulvipes (Robineau-Desvoidy),

Siphona , 102

fulvipes Robineau-Desvoidy,

Ceranthia, 99

fuscicomis Robineau-Desvoidy,

Siphona, 1 19

futilis Wulp, Siphona, 115

gedeana Wulp, Siphona, 1 15

geniculata (De Geer), Siphona,

109-110, 112-113, 115, 162

geniculata De Geer, Musca, 108

gibbicomis (Mesnil), Peribaea, 8 1

gracilis (Mesnil), Siphona, 116

grandistylum Pandelle, Siphona,

116

gratiosa (Mesnil), Actia, 68, 73

grisea (Robineau-Desvoidy),

Siphona , 102

griseola Mesnil, Siphona, 116

hargreavesi Curran, Actia, 73, 159

hartigii (Ratzeburg), Goniocera,

44

heterochaeta Bezzi, Actia,

131-132

hirsuta (Shima), Peribaea, 8 1

hirticeps (Malloch), Ceromya, 61

hirticeps Malloch, Actia, 53

hokkaidensis Mesnil, Siphona,

110, 117

humeralis (Robineau-Desvoidy),

Actia, 75

humeralis Robineau-Desvoidy,

O’Hara

Siphona, 119

hungarica Andersen, Siphona, 113,

116

hurdi (Reinhard), Siphona, 99

hurdi Reinhard, Aphantorhapha,

97

hyalinata (Malloch), Peribaea, 81,

162

illinoiensis Townsend, Siphona,

116

illugiana (Shima), Peribaea, 8 1

infantula (Zetterstedt), Actia, 68,

73, 159

infuscata (Mesnil), Siphona, 111,

116

ingerae Andersen, Siphona,

112-113, 116

insularia (Shima), Peribaea, 81

interrupta Curran, Actia, 73, 159

intrudens (Curran), Siphona,

112-113, 116

invalida (Malloch), Ceromya,

53-56,61

io (Aldrich), Goniocera, 24, 37,

42 — 43, 52-53, 57, 162, 172,

180

io Aldrich, Cartocometes, 41

janssensi (Mesnil), Siphona, 111,

114

japonica (Mesnil), Siphona,

101-102

jepsoni (Villeneuve), Peribaea, 77,

81

jocosa (Villeneuve), Siphona,

101-102

jocularis Mesnil, Actia, 68, 73, 159

juniperi (O'Hara), Siphona, 98-99,

162

kairiensis O'Hara, Siphona, 1 16

kuscheli (Cortes), Siphona, 116

labellata Kamran, Actia, 76

laboriosa Mesnil, Siphona, 96

Systematics of the Genus Group Taxa of the Siphonini

225

lacrymans (Mesnil), Siphona, 102

lamia (Meigen), Actia, 68, 73, 159

lamia Meigen, Roeselia, 67

languidula (Villeneuve), Ceromya,

54, 57,61,64

languidulina Mesnil, Ceromya, 61

lata Malloch, Actia, 74

laticomis (Malloch), Siphona, 93,

96

laticomis Curran, Siphona, 116

latifrons Meigen, Goniocera, 44

latifrons Meigen, Tachina, 41

latipalpis (Malloch), Ceromya, 61

lavinia (Curran), Ceromya, 54, 61

leucopheae (Mesnil), Peribaea, 81

lichtwardtiana (Villeneuve),

Siphona, 101-102, 162

lindneri Mesnil, Siphona, 1 16

linguata Mesnil, Actia, 74

livoricolor (Mesnil), Siphona, 102

lobata Mesnil, Peribaea, 77, 81

longilingua (Mesnil), Actia, 68, 74

longimana Mesnil, Ceromya, 61

longiseta (Villeneuve), Peribaea,

81

longissima O'Hara, Siphona, 1 16

ludibunda (Robineau-Desvoidy),

Goniocera , 44

lurida Reinhard, Siphona, 116

lutea (Townsend), Ceromya, 23,

54, 57,61

lutea (Townsend), Siphona, 110,

116

lutea Toensend, Actia, 53

luteicomis (Curran), Ceromya, 53,

62, 162

macronychia Mesnil, Ceromya, 44,

58

macronychia Mesnil, Proceromyia,

36, 44-47, 58

macronyx O'Hara, Siphona, 116

maculata Staeger, Siphona, 1 1 0,

112-113, 116, 162

maculipennis (Malloch), Ceromya,

62

maculipennis Meigen, Siphona,

119

magnicomis Malloch, Actia,

67-69, 74

maksymovi Mesnil, Actia, 74, 159

malaisei (Mesnil), Actia, 68, 74

malayana (Malloch), Peribaea, 81

mallochiana (Gardner), Siphona,

96, 162

martini Andersen, Siphona , 112,

117

medialis O'Hara, Siphona, 117

meigenii Lepeletier and Serville,

Siphona , 116

melania (Bezzi), Siphona, 1 17

melanocera Robineau-Desvoidy,

Siphona, 119

melanura Mesnil, Siphona, 85,

110, 117

mellina (Mesnil), Ceromya, 64-66

mesnili Andersen, Siphona, 110,

113, 117

microcera (Robineau-Desvoidy),

Siphona , 102

mimetica Malloch, Actia, 74

minuta (Fabricius), Siphona , 115

minuta Fabricius, Stomoxys, 108

minuta Robineau-Desvoidy,

Peribaea , 83

mitis (Curran), Peribaea, 81

modesta (Mesnil), Peribaea, 20,

77,81

mongolica Richter, Actia, 74

monstrosicomis (Stein), Ceromya,

37, 39, 46, 50, 52, 54, 57-59,

62, 172-173

monstrosicomis Stein, Stenoparia,

52

monticola (Malloch), Peribaea ,

Quaest. Ent., 1989, 25 (1,2)

226

O’Hara

79, 82

montium (Villeneuve), Goniocera,

41 — 43, 171

montium Villeneuve, Actia, 41

multifaria O'Hara, Siphona, 117

munroi Curran, Actia, 68, 74

munroi Curran, Siphona, 117

murina (Mesnil), Siphona, 117

nana (Curran), Peribaea , 83

natalensis (Curran), Ceromya, 39,

52, 54, 57, 62

nigra Shima, Actia, 74

nigrapex Mesnil, Actia, 74

nigricans (Villeneuve), Siphona,

110, 113, 117, 162

nigrifrons (Robineau-Desvoidy),

Actia, 75

nigripalpis (de Meijere), Siphona,

115

nigripalpis (Robineau-Desvoidy),

Actia, 72

nigripes (Curran), Peribaea, 82

nigritula (Malloch), Peribaea, 82

nigriventris Malloch, Actia, 68, 74

nigrohalterata (Villeneuve),

Entomophaga, 37, 46-50, 58,

172

nigrohalterata Mesnil, Siphona,

111, 117

nigrohalterata Villeneuve,

Ceromya, 46

nigronitens Mesnil, Siphona, 23,

93-94, 96

nigroscutellata Lundbeck, Actia,

74, 159

nigroseta Curran, Siphona, 117

nigrovittata Meigen, Siphona, 116

nitidella Villeneuve, Actia, 69, 74

nitidiventris Curran, Actia, 74

nobilis (Mesnil), Siphona, 117

norma (Malloch), Siphona, 96, 162

normula (Curran), Ceromya,

64-65

nudibasis Stein, Actia, 68, 74-75,

159

obesa (Mesnil), Siphona, 1 17

oblimata Mesnil, Actia, 74

obscurella Robineau-Desvoidy,

Actia, 73

obscuripennis Curran, Siphona,

117

oculata Pandelle, Siphona, 1 18

oligomyia O'Hara, Siphona, 1 10,

117

Ontario (Curran), Ceromya, 43, 53,

62, 162, 171

orbata (Wiedemann), Peribaea, 79,

81, 162,218-219

orbata Wiedemann, Tachina, 78

orientalis (Townsend), Peribaea,

82

orientalis (Townsend), Siphona,

95-96.

orientalis Townsend,

Aphantorhaphopsis, 92

orientalis Townsend,

Eogymnophthalma, 77

pacifica O'Hara, Siphona, 117

painei Crosskey, Actia, 75, 159

palaestina (Villeneuve), Peribaea,

82, 162

pallens Curran, Actia, 75

pallida (Herting), Siphona,

100-102, 162

palloris (Coquillett), Ceromya, 40,

43, 54, 62, 162

palloris Coquillett, Lasioneura, 52

palpalis (Rondani), Actia, 72

palpina Zetterstedt, Siphona, 1 14

paludosa Mesnil, Siphona, 1 17

pamirica Richter, Actia, 75, 159

panamensis Curran, Siphona,

129-131

parviseta Malloch, Actia, 67-69,

Systematics of the Genus Group Taxa of the Siphonini

227

75, 159

patellicomis Mesnil, Ceromya, 66,

162

patellipalpis (Mesnil), Siphona,

117

pauciseta Kamran, Actia, 76

pauciseta Mesnil, Siphona, 119

pauciseta Rondani, Siphona, 1 17

pectinata (Shima), Peribaea, 77, 82

pellex (Mesnil), Actia, 75

pendleburyi (Malloch), Ceromya,

66

perdita Malloch, Actia, 68, 75

perispoliata (Mesnil), Siphona , 96

phantasma (Mesnil), Siphona, 1 18

philippinensis Malloch, Actia, 75

picipalpis (Mesnil), Actia, 75

picturata (Mesnil), Siphona, 96

pigra Mesnil, Siphona, 118

pilipennis (Fallen), Actia, 75, 159

pilipennis Robineau-Desvoidy,

Actia , 73

pisinnia O'Hara, Siphona, 118

plebeia (Malloch), Peribaea, 82,

162

plorans (Mesnil), Siphona, 102

plusiae Coquillett, Siphona, 40,

120, 122-123, 162

pokharana Shima, Actia, 75

portentosa Mesnil, Ceromya, 62

pruinosa Shima, Ceromya, 60, 62,

162

pseudomaculata Blanchard,

Siphona, 118, 162

pubioculata (Mesnil & Shima),

Proceromyia, 15, 34, 36,

44-46, 58-59

pubioculata Mesnil and Shima,

Nipponoceromyia, 44, 46-47

pudica Mesnil, Siphona, 96

pulex Baranov, Actia, 68, 75

pulla (Reinhard), Siphona, 91,

129-131

pulla Mesnil, Peribaea, 82

pulla Reinhard, Aphantorhapha,

131

punctipennis (Malloch), Ceromya,

54, 62

punctum (Mesnil), Ceromya, 62

pusilla Robineau-Desvoidy,

Siphona, 119

quadrinotata Robineau-Desvoidy,

Siphona, 119

quadriseta Malloch, Actia, 75

reducta (Mesnil), Siphona, 111,

118

reducta ludicra Mesnil, Siphona,

118

reducta Villeneuve, Actia , 75

rejecta Bezzi, Actia, 75

repanda (Mesnil), Peribaea, 82

resinellae (Schrank), Actia , 74

rizaba O'Hara, Siphona, 118

robertsonii (Townsend), Neaera,

179

rossica Mesnil, Siphona, 118

rotundicomis (Malloch), Ceromya,

62

rotundipennis (Malloch),

Peribaea , 79, 82

rubea Mesnil, Peribaea, 82

rubiginosa (Mesnil), Actia, 75

rubrapex Mesnil, Siphona, 118

rubrica (Mesnil), Siphona, 118

rubrifrons (Robineau-Desvoidy),

Actia, 76

rufescens (Greene), Actia, 68, 76

rufina (Zetterstedt), Ceromya , 60

russula Mesnil, Actia, 76

samarensis (Villeneuve), Siphona,

94, 96, 162

schistacea Brauer and

Bergenstamm, Goniocera,

41-43, 162

Quaest. Ent., 1989, 25 (1,2)

228

scutellaris (Rondani), Actia, 72

scutellata (Mesnil), Siphona, 102

sedlaceki (Shima), Peribaea, 82

selangor (Malloch), Siphona,

95-96

selecta (Pandelle), Siphona, 96,

162

selecta Pandelle, Thryptocera, 92

setinerva (Mesnil), Siphona, 118

setinervis (Thomson), Peribaea, 82

setipennis (Fallen), Triarthria, 179

setosa Mesnil, Siphona, 118, 162

seyrigi Mesnil, Siphona, 1 1 8

siebeckii (Sintenis), Ceromya, 66

silacea (Meigen), Ceromya, 11,

51-53,56, 59, 63-66, 80, 162,

173, 192

silvarum Herting, Siphona, 1 17

silvatica Robineau-Desvoidy,

Siphona, 1 19

similata (Malloch), Peribaea, 82

similata Mesnil, Ceromya, 64, 66

simulans (Mesnil), Siphona, 118

singularis (Wiedemann), Siphona,

20, 88-89

singularis Wiedemann, Tachina,

87

siphonoides (Strobl), Siphona, 96,

162

siphonosoma Malloch, Actia, 67,

76

sola Mesnil, Siphona, 118

sonorensis (O'Hara), Siphona, 99

sororcula (Mesnil), Peribaea, 82

speciosa Mesnil, Siphona, 97

spinulosa (Mesnil), Siphona, 118

spoliata (Bezzi), Peribaea, 82

starkei (Mesnil), Siphona, 97

stiglinae (Bezzi), Peribaea, 83,

131, 162

stiglinae Bezzi, Actia, 131

subaequalis (Malloch), Peribaea,

O’Hara

79, 82

subopaca (Aldrich), Ceromya, 62

sufferta (Villeneuve),

Entomophaga, 5 1

sulfurea (Mesnil), Siphona,

101-102

suspecta (Malloch), Peribaea, 83,

162

tachinaria Meigen, Siphona, 1 15

taiwanica (Baranov), Siphona, 1 19

takanoi Baranov, Actia, 68, 76

tarsata Richter, Actia, 68, 76

tenuipalpis (Villeneuve), Siphona,

101, 103

tenuis Curran, Siphona, 1 16

terrosa (Mesnil), Siphona, 100,

103

testacea Robineau-Desvoidy,

Ceromya, 52, 60

testacea Robineau-Desvoidy,

Siphona, 1 19

tibialis (Robineau-Desvoidy),

Peribaea, 80, 83, 162

tibialis Robineau-Desvoidy,

Herbstia, 77

timida (Mesnil), Peribaea, 83

trichaeta (Mesnil), Siphona, 109,

118

trifurcata (Shima), Peribaea,

77-78, 83

triseta (Mesnil), Actia, 76

tristella (Herting), Siphona, 101,

103, 162

tristis Robineau-Desvoidy,

Siphona, 119

tropica (Townsend), Siphona, 118

tropica Townsend,

Phantasiosiphona, 108

ugandana (Curran), Peribaea, 77,

83

unicolor (Aldrich), Ceromya, 62

uniseta (Malloch), Peribaea, 83

Systematics of the Genus Group Taxa of the Siphonini

229

uniseta Malloch, Actia, 77

unispina (Mesnil), Siphona, 111,

118

unispina infuscata (Mesnil),

Siphona, 1 1 1

unispina unispina (Mesnil),

Siphona, 111

urbanis (Harris), Siphona, 115

uruhuasi (Townsend), Siphona, 12,

124-125

uruhuasi Townsend, Uruactia, 123

ussuriensis (Mesnil), Peribaea, 83

valida (Curran), Ceromya, 53, 55,

62

valida Curran, Actia, 53

variata Andersen, Siphona, 1 13,

118

varichaeta (Curran), Ceromya,

64-66

verralli (Wainwright), Siphona, 97

versicolor (Fallen), Goniocera,

41-44, 162, 171

versicolor Fallen, Tachina, 41

vidua (Mesnil), Peribaea, 83

villeneuvii (Strobl), Actia, 73

vitripennis Rondani, Actia, 73

vittata Curran, Siphona, 119

vivida (Robineau-Desvoidy),

Siphona, 102

vixen Curran, Siphona, 1 19

vulpina (Mesnil), Actia, 76

wittei (Mesnil), Siphona, 119

xanthogaster (O'Hara), Siphona,

99, 162

xanthosoma Mesnil, Siphona, 97

yasumatsui Shima, Actia, 68, 76

Quaest. Ent., 1989, 25 (1,2)

.

■

Commentary

231

COMMENTARY

[Commentary is a section of Quaest. Ent. that appears from time to time, and will

contain expressions of opinions about general items, controversial or otherwise, that

ought to be of interest to many of our readers. These contributions will not be

refereed because they are intended to be free expressions of opinion. Changes by the

Editor might be made to the form of presentation, but not to its substance. Remarks

that are deliberately abusive or insulting will not be published. Rebuttals to

previously expressed views will be considered, but the journal is under no obligation

to publish them.

The Editor]

Following is an extended book review. Because of the potentially controversial

nature of some of the comments, it seems more appropriate to place the review in a

section of the journal that invites dialogue— hence its location in “Commentary”.

LIEBHERR, J. K. (Editor). 1988. Zoogeography of Caribbean Insects. Comstock

Publishing Associates, Cornell University Press, Ithaca and London, xi + 285 pp.

Price, $39.95 (U.$.)

Printed on acid-free paper, this volume is attractively hardbound, with dark green

covers. On the front cover is a figure illustrating the geographical distribution and

reconstructed phylogeny of the drosophilid genus group Pseudiastata and an adult

of the West Indian Mayagueza argentifera, a member of this taxon. It is an excellent

illustration and provides a focus on the subject matter of the book. The lighter green

end-papers inside the covers contain a useful map of the Caribbean Basin, including

the islands of the West Indies and adjacent portions of the North, Middle, and South

American mainland, complete with scale and indications of longitude and latitude.

The volume includes a preface, list of contributors, and 1 1 chapters. Subject and

taxonomic indices end the volume. Chapter 1 is a general discussion of Caribbean

zoogeography. Chapter 2 treats geological aspects; and Chapter 1 1 is a critique of

biogeographical methods in general, and of work of the other authors of this

volume, in particular.

Chapters 3 to 10 contain analyses of taxa, as follows: Lygaeidae, by J.A. Slater;

auchenorrhynchous Homoptera of the Greater Antilles, by J.A. Ramos; scaritine

Carabidae, by S.W. Nichols; platynine Carabidae, by J.K. Liebherr; polycentropodid

caddisflies, by S.A. Hamilton; relict Drosophilidae, by D.A. Grimaldi; ants, by E.O.

Wilson; and halictid bees, by G.C. Eickwort. Each taxon-based chapter is a rich

source of clearly presented information, containing extensive lists of included taxa

and their distributions. Most of the chapters contain very well executed illustrations

of the insect group treated, or of their work, and maps and diagrams are also

provided that amplify the text. The chapter by Ramos is not illustrated, and overall

seems rather perfunctory.

Quaest. Ent., 1989, 25 (1,2)

232

Ball

As a carabid specialist, I was impressed especially with the execution of the

distribution maps in Nichols’ chapter: each with a photograph illustrating habitus of

one or more species whose range is indicated by dots, etc. Much care went into the

preparation of these figures, and collectively they are a valuable source of data.

Recurrent themes in the text are: occurrence of old relicts on Puerto Rico (noted

by Slater, Nichols, Liebherr, and Grimaldi): and incomplete knowledge of the biota,

because of insufficient collecting by appropriate specialists.

The analyses of various taxonomic groups are inconsistent in delimitation of the

study area. In his concept of the West Indian Biogeographic Region, Nichols

includes the Greater and Lesser Antilles, the Bahamas, Barbados (as a separate

entity). South Florida, and the Yucatan Peninsula of Mexico. Wilson does not

include the mainland in his treatment of the ants, but does include Trinidad and

Tobago. This inclusion increases the size of the ant fauna of the West Indies by

about a third. The other authors confine their study areas to the Bahamas, and

Greater and Lesser Antilles, with Barbados included in the last-named island group.

Such differences in definition of study area must be taken into account by those who

might wish to compare taxonomic diversity of the various taxa in the West Indies.

The Editor claims in the title of the Introduction (Chapter 1) that the Caribbean

area is a “fertile ground for zoogeography”. He provides a clear, even-handed

discussion of the history of ideas about assembly of the West Indian biota; first,

postulation of land bridges to make possible movement of the ancestral terrestrial

biota from mainland to islands; second, carefully reasoned dispersal theory, without

invocation of land bridges; and third, plate tectonic theory, with its pieces of land,

originally close to mainland Central and South America, that drifted eastward, and

brought an essentially mainland biota to the development of the present Greater

Antilles.

Liebherr indicates that this basic question of the mechanism of faunal assembly

has not been settled, and maintains (p. 10) that “geologic data often cannot provide

unequivocal answers about the history of areas, making biological data of utmost

importance in the interpretation of faunal histories”. He advocates use of vicariance

biogeography in this endeavour. However, I doubt that conclusions drawn from

biological data can be more compelling than geological data, by whatever means the

former are interpreted.

The Editor makes a convincing case that the Caribbean area is fertile ground for

zoogeographers. He concludes his opening chapter by indicating the great potential

of data derived from insects, as follows:

1 . Insects are apt to help clarify old patterns of faunal relationships, because

the West Indies have a fair number of taxa that exhibit relations with

Africa rather than with New World taxa, and this implies Gondwanian

connections. Liebherr implies that such relationships are direct, i.e.,

without extinct New World mainland intermediates, or even

Commentary

233

intermediates that have yet to be discovered.

2. Many taxa in the Antilles are conspecific with or very closely related to

mainland Neotropical species. These taxa indicate overwater dispersal

between mainland and islands.

3. There is a rich endemic fauna in the islands, which provides an

“extensive potential data base for intra-island analyses”.

He notes, however, that the insect fauna of the Antilles is not well known, and

that much field work is required to elucidate “species distributions and habitat

requirements”. Mindful of the destruction of habitats that is taking place in the

islands and the effect that such will have on distribution patterns before there is the

chance to study them, he advocates activity in protection of the biota.

Thomas W. Donnelly, a geologist and dragon fly specialist with extensive

experience in the Caribbean area, and thus sympathetic to the requirements of

biogeographers in interpreting geological history, provides important background

information in his chapter entitled “Geologic Constraints on Caribbean

Biogeography”. Reviewing the evidence on the basis of plate tectonic theory,

Donnelly argues that an island arc formed between Central and South America,

providing a tenuops connection between these land masses. In latest Cretaceous and

early Cenozoic times, this arc was broken into fragments as a “flood basalt moved

eastward” (p.33). These fragments, or terranes, formed the present Greater Antilles,

but some (such as proto- Jamaica) were totally submerged for extended periods.

Cuba was formed in the late Cretaceous by diverse terranes that were “swept

northward” with the opening of the Yucatan Basin. In the Middle Cenozoic

(Oligocene to early Miocene), the continued eastward movement of the Caribbean

Plate closed the gap that separated Central and South America and another island

arc system served as a limited filter bridge for terrestrial organisms, between the two

continents. The lesser Antilles, during mid-Cenozoic, was a series of separate

fragments more distant from South American than from the Greater Antilles. The

volcanic arc that formed during the late Cenozoic provided a filter bridge for

dispersal from South America, but geological evidence minimizes the probability of

an earlier Cenozoic connection of the mainland and the Lesser Antillean arc.

For most of late Mesozoic and Cenozoic time, faunal movements into the

proto-Antilles and Greater Antilles would have required overwater dispersals in the

order of tens of kilometers. For brief periods, the water gaps might have been

“relatively narrow”, and there could have been terrestrial connections with northern

Central America, and between the islands. Overwater dispersal must be emphasized,

though geologists are beginning to find evidence for “limited vicariant interchange”.

Donnelly emphasizes that changing climatic conditions during the Cenozoic

must have had profound influence on distribution of the biota. Using evidence of

lateritic soils in the Greater Antilles, he postulates less moderate climates during the

middle Cenozoic than at present, with markedly alternating wet and dry periods. The

Quaest. Ent., 1989, 25 (1,2)

234

Ball

development of the Central American isthmus during the Pliocene must have led to

profound climatic change, and during the Pleistocene there is strong evidence for

aridity during the glacial maxima.

The main point of all this is that biogeographers must be very cautious about

invoking interruptions of continuous land connections to explain present-day

vicariant distributions of related taxa.

One might think that Donnelly’s paper would have caused all of the other

symposiasts to emphasize dispersal theory in explaining extant distribution patterns.

In fact, this did not happen, with the resulting biogeographic analyses forming two

groups: those postulating dispersal theory as the principal means of explaining

extant distribution patterns; and those postulating elimination of former land

connections and subsequent establishment of new connections as the basis for

vicariant patterns.

Of course, vicariant distribution patterns result, whatever mechanism gives rise

to geographically isolated descendants of an originally continuously distributed

ancestral stock. Consequently, it is incorrect to use “vicarism” as a term for a

process. However, for want of a better term, I will use vicarism as is accepted by

some biogeographers to designate postulation of interruptions of continuous parental

ranges as the normal cause of subsequent vicariant distribution patterns of

descendants.

Authors adopting dispersal as the principal cause of vicariant distributions in the

Caribbean area are Slater, Nichols, Wilson, and Eickwort. Vicarists are Liebherr,

Hamilton, and Grimaldi. Wilson hardly acknowledges the existence of the vicarist

school, and interprets the Antillean ant fauna mainly in terms of island

biogeographic theory.

An interesting age correlation emerges. Of the dispersalists, three (Slater,

Wilson, and Eickwort) are appreciably older than the vicarists. Nichols is the

exception. As the youngest and least experienced of the symposiasts, perhaps he was

the most inclined to pay attention to Donnelly. The older individuals had their ideas

formed before the heyday of the vicarists, whereas the others have been developing

their careers during the vicarism period, and thus perhaps they were influenced by

recent events, not to mention forceful protagonists.

Slater argues that congruence of distribution patterns of different groups may not

be applicable in establishing vicariance explanations to islands located relatively

close to different source areas. Taking exception to the late D.E. Rosen’s expressed

antipathy to using dispersal to explain biotic complexity, Slater (p.39) notes that “if

wind patterns, ocean currents, similarity of habitats, and relative proximity of areas

persist over a reasonable period of time, congruent patterns could be developed by

dispersal as well as by vicariance”. He proceeds to establish the high probability of

several mainland-Greater Antillean faunal connections being the result of

over-water dispersal. He concludes by noting the need for improved analyses using

cladistic methods and having more complete collections with which to work.

Commentary

235

Wilson’s analysis is a generally satisfying outline of the geographical history of

the West Indian ant fauna. Based on fossil as well as living taxa, dynamic principles

involving dispersal and possible radiation in situ of certain dispersants, the only

discordant element in Wilson’s account seems to be the presence of the poor -

dispersing Ecitoninae in the fossil record of Hispaniola.

Wilson points out as defects in knowledge of West Indian ants the probable

under-collecting of some islands, particularly Cuba and Jamaica, as well as lack of

knowledge of certain West Indian taxa that are markedly speciose. To this list of

imperfections must be added the absence of phylogenetic analysis of the West

Indian ants. Without this information, the details of geographical history of the

relatively luxuriant Antillean ant fauna will remain obscure.

Nichols supports the argument that the Greater Antilles function as oceanic

islands, using three arguments: first, values derived from Preston’s Similarity Index

indicate that the islandic scaritine fauna is “in a state of flux”: second, the genus

Pasimachus (adults are large, flightless scaritines) is confined to Middle and North

America, with a few species in South Florida and the Yucatan Peninsula: and third,

many of the endemic genera of Coleoptera in the West Indies are borers or live

under bark. Had there been land connections between the islands and mainland,

presumably Pasimachus would be represented in the Greater Antilles. Similarly,

because a preponderance of endemic and thus older genera of the West Indies live in

situations suitable for transport by rafting (i.e., logs) their disproportionate

representation in the islands is argument for overseas dispersal. The “state of flux”

argument based on the Preston Similarity Index seems to indicate that faunal

composition is being determined by forces working on an ecological time scale, and

if so, strength is added to Nichols’ more general argument. However, the pattern

might be an artifact of the method of analysis. As Connor points out (p. 258) this

index emphasizes the effects of forces working on an ecological time scale, and

these may outweigh historical relationships. Thus, the disturbance of the general

order in the fauna implied by its being “in a state of flux” or kaleidoscopic, may not

reflect an older underlying reality. Be that as it may, the present pattern of scaritine

distribution does seem to me to be rather unordered.

Evidence presented by the vicarists for their hypotheses is interesting but less

than convincing. In fact, Hamilton shows, in his phylogenetic reconstruction of the

Polycentropus nigriceps group (Figs. 7-9, notes G and H, p. 159), that only the

terminal clades fit the pattern required by the Rosen model. He concludes that “this

cladistic analysis.... gives no clear evidence of disjunct inter-island patterns of

relationship”, and calls for cladistic analyses of other Greater Antillean groups to

search for a common pattern. Taken at face value, however, the data presented do

not fit the Rosen model, and the author is left in the uncomfortable position of

having to question the value of the data presented, thus: “A cladistic analysis of the

nigriceps group based on the semaphoront [read holomorph] (not just adult

male). ..would undoubtedly test and enhance the results I have presented here”.

Quae st. Ent., 1989, 25 (1,2)

236

Ball

Accepting the analysis presented as preliminary, therefore, I would be more

encouraged to test the Rosen model further if it were supported by the preliminary

data. It seems to me that the pattern of Polycentropus is suggestive of inter-island

dispersal, and accordingly, it is a dispersalist hypothesis that ought to be tested with

a more complete set of data about the species of this genus.

Grimaldi’s reconstruction of the geological history of the Greater Antilles calls

for a close connection of the proto- Antillean land mass with Africa, following near

separation of the former from the American mainland. This seems to be required

because of the relationships of some old lineages of Drosophilidae with Old World,

rather than with New World, extant lineages. On two counts, I find the argument

unconvincing: first, Donnelly’s account of Caribbean geological history does not

support Grimaldi’s hypothesis (in fact, there is no reference to it), and second, just

because relicts with African affinities occur in the West Indies, it is not necessary to

postulate a direct former connection between the two areas. Puerto Rico could be the

last area in the New World where a former widespread lineage has been able to

survive. Of the larger Antillean islands, that one is farthest from the mainland, and

thus might be expected to accumulate relicts, under the strictures of a dispersalist

hypothesis as developed by P. J. Darlington, Jr.

In Liebherr’s treatment of Platynus, the fauna of each island is discussed, with

emphasis on cladistic relationships, and problems therewith. The major problem

with the phylogenetic analysis is lack of characters in which one can have

confidence. Liebherr recognizes one assemblage (the wingless group) that is based

on wing loss (character 40) and displacement of the setae of the posterior angles of

the pronotum (character 14). Neither of these features is very reliable as an indicator

of relationships, and for the setal feature, this instability is highlighted by the

necessity to hypothesize a reversal within the wingless clade, at the base of the P.

jaegeri group (Figs. 6-7 and 6-8). In that same clade, character 17 (width of pronotal

margins) is used to relate the P. cinchonae and P. jaegeri groups. I have no doubt

about the value of this character for determining relationships of similar adjacent

allopatric taxa that differentiated comparatively recently, but I am skeptical that

such a feature is useful to establish relationships of geographically widely separated

clades including a total of 18 species.

Nonetheless, Liebherr uses the hypothesized relationships of the flightless clade

in his geographical analysis, suggesting that Rosen’s vicariance model “adequately

explains taxon relationships among species on Cuba and south and central

Hispaniola, and implying that island vicariance and hybridization have been at work

along the northern edge of the Caribbean plate’’. If, in fact, the extreme mobilist

hypothesis were established, or if relationships of the Platynus species involved in

testing the model were more convincingly demonstrated, one would have cause to

accept the underlying theory of vicariance biogeography as applied to the West

Indies. Under the circumstances, I find little basis for use of Rosen’s model in

interpreting the history of the West Indian biota.

Commentary

237

By accepting at face value the result of the numerical analysis that relates several

montane lineages with brachypterous adults, Liebherr is not in a position to consider

the possibility that each of these lineages came from lowland winged ancestors

which invaded montane habitats on their respective islands, with subsequent

extinction of the lowland ancestors and loss of wings among the upland survivors.

Such an interpretation may be contrary to the principle of parsimony as practiced by

numerical cladists, but the resulting picture might make at least as much sense

biogeographically as the interpretation based on the Rosen vicariance model.

Although I am not persuaded of the Rosen model of vicariance nor of the details

of relationships and historical interpretation of the distribution pattern of the

Platynus taxa as presented by Liebherr, I am impressed by the clarity of the

presentation and appreciate the value of this contribution in highlighting

phylogenetic and zoogeographic areas for further investigation. Discussion of causal

relations between brachyptery and diversity, and between climatic change and origin

of the bromeliad-inhabiting fauna of Jamaica are insightful, valuable contributions

to the more general aspects of West Indian biogeography.

The concluding chapter (No. 1 1), by Edward F. Connor, outlines mathematically

acceptable procedures for inferring historical biogeographic relationships. His

principal conclusions are two: first, none of the other authors of this volume used

methods acceptable to mathematicians for inferring historical biogeographic

relationships: two, “it is best to examine the biogeographical evolution of the

Caribbean biota independently of geologic hypotheses concerning area

relationships”. I suppose that systematists should pay attention to the

pronouncements of their more mathematically inclined brethren, and in a logical

sense, I can appreciate why one might want to analyze biogeographical data

independently of geological hypotheses. However, I believe that, in the absence of

the required mathematical precision that plagues most types of biogeographical data,

Hennig’s principle of reciprocal illumination can be applied to the available

geological and phylogenetic biogeographic data, with reasonable approximations to

the truth thus being obtained. I suspect it will be more useful for biogeographers to

improve the quality of their taxonomic and phylogenetic information than to invest

too much effort in elaborate statistical treatment of what is now available.

Connor’s negative assessments aside, based on study of this volume I conclude

that in view of clear evidence of past crossing of sea barriers by insects, flying or

otherwise, and in the absence of clear evidence for the geologic basis of a vicariance

hypothesis, little is to be gained by developing a research program to test further the

tenets of that hypothesis. Gains are to be made, first by improving the data available

for analysis (i.e., more complete collections of the islandic faunas; better

information about way of life and local distribution of the species), and second by

undertaking phylogenetic analyses of the taxa, to be interpreted using a dispersalist

theory, which takes into account the tenets of the theory of island biogeography and

the Darlington- Wilson-Erwin theory of taxon cycles or pulses. If such analyses fail

Quaest. Ent., 1989, 25 (1,2)

238

Ball

to provide a satisfactory account of the biota and its history, then it will be time to

seek other explanatory means.

In conclusion, I found this volume to be enlightening and interesting, and a

valuable contribution both to entomology and to the growing knowledge of the West

Indian biota. My principal negative comment about the presentation, as such, is the

lack of a concluding chapter that could have discussed and perhaps attempted to

resolve the different interpretations of biotic history by the various authors. It would

have been desirable for each author to have used the same definition of the study

area, so that the resulting data could be compared more easily.

This volume ought to be owned and studied by anyone interested in West Indian

biogeography. Otherwise, various chapters can be read with profit by taxonomists

interested in the taxa treated therein, though not interested in the West Indian fauna,

as such.

George E. Ball

Department of Entomology

January, 1989

Book Review

239

BOOK REVIEW

IVES, W. G. H. and H. R. WONG. 1988. Tree and shrub insects of the prairie

provinces. Information Report NOR-X-292. Northern Forestry Centre, Canadian

Forestry Service. Edmonton, Alberta. T6H 3S5. 327 pages, 117 full page colored

plates. Available at no charge to persons in the forest industry and educational

institutes. ISBNO-662- 15770-2.

Even though I am acknowledged in this book as one of the scientific reviewers,

my contribution consisted primarily of proof-reading several drafts of

computer-processed text that did not at all resemble the very attractive-looking

finished product. I feel justified, therefore, in publicly reviewing it, particularly now

that I have had a chance to browse through it and, more importantly, to use it. After

the title page, the contents are succinctly described in an abstract , which is worth

quoting directly,:

“More than 600 species of insects and mites that feed on trees and shrubs in the

prairie provinces are discussed in terms of their distribution, hosts, and

importance. There are approximately 1 100 color photographs; the life cycle and

damage of major pest species are illustrated, and for common species, primarily

the larval stage is shown. Pests attacking coniferous hosts are separated from

those attacking hardwoods, and each group is subdivided according to the type

of feeding damage and type of insect involved. An extensive bibliography is

provided. In addition to a taxonomic index listing the order and family of each

insect and mite, there is a diagnostic index listing the host species, feeding site

of the pest, and species of insect or mite involved and an insect index”.

Obviously, this is a major scientific work that should appeal to a wide

cross-section of users such as students, professional entomologists and ecologists,

anyone associated with forestry activities, and even the general public; hopefully,

favorable responses from this large audience will generate a lot of good will for the

Canadian Forestry Service.

Restrictions of support for insect identification services for provincial agencies

prompted the need for this insect guide. However, it took the combined talents of an

insect ecologist (W. G. I. ) and an insect taxonomist (R. H. W.) to successfully

surmount the nomenclatural problems that are bound to arise when dealing with so

many species of insects and plants, and to bring together a formidable knowledge of

tree and shrub insects of the prairie regions of Canada. No attempt was made to

emphasize economically important insects so what we have here is an ecological

check list of insects and mites, with descriptions and photographs, associated with

native and introduced trees and shrubs of parklands, shelter belts, urban areas of the

prairies and parts of the Boreal forest and sub-montane prairie. No claim is made

that all insects are included but I believe that a very small proportion have been

Quaest. Ent., 1989, 25 (1,2)

240

Book Review

missed; after all, both authors have had considerable experience in forest

entomology and have had the time to build up large collections, make countless

rearings, and record insect occurrences, over many years.

Even a cursory glance at the contents makes it apparent that a tremendous effort

went into the production of this book. The two-column layout on standard

letter-sized glossy paper is pleasing to the eye and individual species names stand

out in bold type. The color plates are on the left hand page and refer only to the

species covered on the right hand page, making this one of the easiest to use

technical insect identification books I have encountered. All photographs are, in

general, faithful renditions of the actual subjects. Text references consist of numbers

that refer to items in a bibliography of 657 citations preceding the indices at the end

of the book, and are given for species covered in each plate and discussed in the

text. I would quibble about the correctness of the word “pest” as it is sometimes

used, and I think that the inclusion of a photograph of an adult of a representative

looper would have been useful, but these are minor points. The soft covers, though

beautifully illustrated on the front with a full color painting of a Malacosoma

disstria larva, will probably not hold up for very long; in any case, a valuable

reference like this should be bound in hard covers.

Unbelievably, there is no charge for this publication. Yet most of my colleagues

and myself would have gladly bought it for the $75.00 or so per copy that it took to

produce. It is a book to be looked at for its illustrations, read for its interesting

biological information, and used as an identification guide. It should be on the

bookshelf of every entomologist interested in ecological diversity, the beauty of

insects and natural history, recommend it highly.

W. G. Evans

Department of Entomology

University of Alberta

I

'Quaestiones.

Entomolosicae

A periodical record of entomological investigations,

published at the Department of Entomology,

University of Alberta, Edmonton, Canada.

VOLUME 25

NUMBER 3

SUMMER 1989