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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

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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

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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

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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

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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

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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

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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

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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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Systematics of the Genus Group Taxa of the Siphonini

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Systematics of the Genus Group Taxa of the Siphonini

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Epiphallus Male distiphallus Postgonite Male surstylus Cerci Female Female

Taxon size posterior posterolateral lateral shape basally shape shape ovipositor sternum 6

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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)

38

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)

40

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

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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

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44

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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

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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.

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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

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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

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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

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60

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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

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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).

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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.

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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.

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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.

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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.

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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).

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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

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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).

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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.

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

88

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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92

O’Hara

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

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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

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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).

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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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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

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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

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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

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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;

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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

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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.).

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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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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.

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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

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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.

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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).

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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

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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.

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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.

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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.

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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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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.

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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.

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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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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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anterior

107. Goniocera io

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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O’Hara

anterior

120. Peribaea apicalis

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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O’Hara

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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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

SIPHONINE & HOST SPECIES

FAMILY

REFERENCE

Actia crassicornis (Meigen)

Depressaria applana Fabr.

Oecophoridae

Lundbeck, 1927:461; Herting, 1960:60;
Mesnil, 1963a:817

Depressaria astrantiae Heinem.

Oecophoridae

Herting, 1960:60; Mesnil, 1963a:817

Depressaria bupleurella Heinem.

Oecophoridae

Herting, 1960:60; Mesnil, 1963a:817

Depressaria costosa Haworth

Oecophoridae

Herting, 1960:60; Mesnil, 1963a:817

Depressaria depressella Hb.

Oecophoridae

Herting, 1960:60; Mesnil, 1963a:817

Depressaria heydenii Zeller

Oecophoridae

Lundbeck, 1927:461; Herting, 1960:60;
Mesnil, 1963a:817

Depressaria hippomarathri Nick.

Oecophoridae

Herting, 1960:60; Mesnil, 1963a:817

Depressaria liturella Schiff.

Oecophoridae

Herting, 1960:60; Mesnil, 1963a:817

Sparganothis pilleriana Sch.

Tortricidae

Herting, 1960:60; Mesnil, 1963a:817

Tortrix viridana L.

Tortricidae

Lundbeck, 1927:461; Herting, 1960:60;
Mesnil, 1963a:817

Yponomeuta mallinellus Z.

Yponomeutidae

Hubenov, 1985:29

Actia diffidens Curran

Acleris variana Femald

Tortricidae

Prebble, 1935; Amaud, 1978:52

Choristoneura conflictana Walker

Tortricidae

Amaud, 1978:52

Spilonota ocellana Denis & Schiff.

Tortricidae

Amaud, 1978:52

Actia eucosmae Bezzi

Crocidosema plebeiana Zeller

Tortricidae

Crosskey, 1973:176; Chadwick &
Nikitin, 1985:588

Actia hargreavesi Curran

Dysdercus sp. (Hemiptera)

Pyrrhocoridae

Mesnil, 1963a:814 (as Actia comitata)
(doubtful record)

Actia infantula (Zett.)

Monopis rusticella Hiibner

Tineidae

Herting, 1960:61; Mesnil, 1963a:819

Actia interrupta Curran

Melanolophia imitata Walker

Geometridae

Amaud, 1978:53

Schizura concinna Smith

Notodontidae

Amaud, 1978:53

Acleris variana Femald

Tortricidae

Amaud, 1978:53

Acleris variegana Denis & Schiff.

Tortricidae

Amaud, 1978:53

Amorbia cuneana Walsingham

Tortricidae

Amaud, 1978:53; Oatman et al.,
1983:53

Archippus packardianus Femald

Tortricidae

Amaud, 1978:53

Ar chips argyrospilus Walker

Tortricidae

Amaud, 1978:53

Archips rosanus L.

Tortricidae

Amaud, 1978:54

Choristoneura conflictana Walker

Tortricidae

Amaud, 1978:54

Choristoneura fumiferana Clemens

Tortricidae

Amaud, 1978:54

Choristoneura pinus Freeman

Tortricidae

Amaud, 1978:54

Choristoneura rosaceana Harris

Tortricidae

Amaud, 1978:54

Croesia albicomana Clemens

Tortricidae

Amaud, 1978:54

Epinotia crenana Hiibner

Tortricidae

Amaud, 1978:53

Epinotia emarginana Walsingham

Tortricidae

Amaud, 1978:53

(Continued on next page)

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O’Hara

Table 2 (continued)

SIPHONINE & HOST SPECIES

FAMILY

REFERENCE

Grapholitha molesta Busck

Tortricidae

Amaud, 1978:53

Phalonia oenotherana Riley

Tortricidae

Amaud, 1978:54

Xenotemna pallorana Robinson

Tortricidae

Amaud, 1978:54

Actia near interrupta Curran

Archips cerasivoranus Fitch

Tortricidae

Amaud, 1978:55

Archips fervidanus Clemens

Tortricidae

Amaud, 1978:55

Actia jocularis Mesnil

Lymantria dispar L.

Lymantriidae

Schaefer & Shima, 1981:370

Actia lamia (Meigen)

Ourapteryx sambucaria L.

Geometridae

Herting, 1960:60; Mesnil, 1963a:822

Lasiocampa querus L.

Lasiocampidae

Wainwright, 1928:208; Herting,
1960:60; Mesnil, 1963a:822

Epiblema luctuosana Dup.

Tortricidae

Herting, 1960:60; Mesnil, 1963a:822

Epiblema pflugiana Haworth

Tortricidae

Lundbeck, 1927:460; Herting, 1960:60;
Mesnil, 1963a:822

Laspeyresia cosmophorana Treitschke

Tortricidae

Mesnil, 1963a:822

Actia maksymovi Mesnil

Cacoecia murinana Hiibner

Tortricidae

Herting, 1960:61; Mesnil, 1963a:823

Semasia diniana Hiibner

Tortricidae

Herting, 1960:61; Mesnil, 1963a:823

Spilonota lariciana Hein.

Tortricidae

Herting, 1960:61; Mesnil, 1963a:823

Actia near maksymovi Mesnil

Dioryctria abietella Denis & Schiff.

Pyralidae

Crosskey, 1976a:291

Actia nigroscutellata Lundbeck

Elachista megerleella Staint.

Elachistidae

Herting, 1960:61; Mesnil, 1963a:825

Laspeyresia cosmophorana Tr.

Tortricidae

Mesnil, 1963a:825

Laspeyresia servillana Dup.

Tortricidae

Herting, 1960:61 (as questionable);
Mesnil, 1963a:825

Olethreutes roseomaculana Herr.-Sch. or
Olethreutes dalecarliana Guenee

Tortricidae

Lundbeck, 1927:463; Herting, 1960:61;
Mesnil, 1963a:825

Semasia ustomaculana Curtis

Tortricidae

Lundbeck, 1927:463; Herting, 1960:61;
Mesnil, 1963a:825

Tortrix viridana L.

Tortricidae

Hubenov, 1985:29

Actia nudibasis Stein

Heringia dodecella L.

Gelechiidae

Herting, 1960:60; Mesnil, 1963a:826

Dioryctria splendidella Herr.-Sch.

Pyralidae

Herting, 1960:60; Mesnil, 1963a:826;
Shima, 1970c: 187

Evetria buoliana Schiff.

Tortricidae

Lundbeck, 1927:459; Herting, 1960:61;
Mesnil, 1963a:826

(Continued on next page)

Systematics of the Genus Group Taxa of the Siphonini

161

Table 2 (continued)

SIPHONINE & HOST SPECIES FAMILY REFERENCE

Evetria resinella L.

Tortricidae

Lundbeck, 1927:459; Herting, 1960:60;
Mesnil, 1963a:826

Actia painei Crosskey

Agonoxena pyrogramma Meyrick

Agonoxenidae

Crosskey, 1962:175

Actia pamirica Richter

Parapandemis chondrillana H. S.

—

Richter, 1974:1269

Spilonota ocellana F.

Tortricidae

Richter, 1974:1269

Actia parviseta Malloch

Isotenes miser ana Walker

Tortricidae

Crosskey, 1973:176; Chadwick &
Nikitin, 1985:590

Actia pilipennis (Fallen)

Platyptilia rhododactyla Schiff.

Pterophoridae

Herting, 1960:59

Acalla aspersana Hiibner

Tortricidae

Herting, 1960:59; Mesnil,

1963a:828

Acalla hastiana L.

Tortricidae

Mesnil, 1963a:828

Acalla logiana Schiff.

Tortricidae

Herting, 1960:59; Mesnil,

1963a:828

Ancylis mitterbacheriana Schiff.

Tortricidae

Herting, 1960:59; Mesnil,

1963a:828

Ancylis tineana Hiibner

Tortricidae

Herting, 1960:59; Mesnil,

1963a:828

Capua angustiorana Haworth

Tortricidae

Herting, 1960:59; Mesnil,

1963a:828

Eucosma sordidana Hiibner

Tortricidae

Mesnil, 1963a:828

Olethreutes schulziana Fabr.

Tortricidae

Mesnil, 1963a:828

Sparganothis pilleriana Schiff.

Tortricidae

Herting, 1960:59; Mesnil,

1963a:828

Tortrix bergmanniana L.

Tortricidae

Herting, 1960:59; Mesnil,

1963a:828

Tortrix loefflingiana L.

Tortricidae

Herting, 1960:59; Mesnil,
Hubenov, 1985:29

1963a:828;

Tortrix pronubana Hiibner

Tortricidae

Herting, 1960:59; Mesnil,

1963a:828

Tortrix viridana L.

Tortricidae

Herting, 1960:59; Mesnil,
Hubenov, 1985:29

1963a:828;

Actia spp. (North America)

Phyganidia californica Packard

Dioptidae

Amaud, 1978:56

Gnorimoschema spp.

Gelechiidae

Amaud, 1978:56

Catocala relicta Walker

Noctuidae

Amaud, 1978:56

Heterocampa guttivitta Walker

Notodontidae

Amaud, 1978:56

Epinotia similana Hiibner

Tortricidae

Amaud, 1978:56

Episimus argutanus Clemens

Tortricidae

Amaud, 1978:56

Actia spp. (Old World)

Gaesa bisignella Snellen

Gelechiidae

Crosskey, 1976a:291

Herpetogramma licarsisalis Walker

Pyralidae

Cantrell, 1986:259

Epiphyas postvittana Walker

Tortricidae

Cantrell, 1986:259

Ceromya americana (Townsend)

Schizura concinna Smith

Notodontidae

Amaud, 1978:51

(Continued on next page)

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O’Hara

Table 2 (continued)

SIPHONINE & HOST SPECIES FAMILY REFERENCE

Ceromya apicipunctata (Malloch)
Unidentified sp.

Noctuidae

Crosskey, 1976a:291

Ceromya bicolor (Meigen)

Phragmatobia fuliginosa L.
Dendrolimus pini L.

Eriogaster lanestris L.

Gastropacha quercifolia L.
Lasiocampa quercus L.

Dolerus pratensis L. (Hymenoptera)

Arctiidae

Lasiocampidae

Lasiocampidae

Lasiocampidae

Lasiocampidae

Tenthredinidae

Herting, 1960:61; Mesnil, 1963a:835
Herting, 1960:61; Mesnil, 1963a:835
Lundbeck, 1927:466; Herting, 1960:61;
Mesnil, 1963a:835
Herting, 1960:61; Mesnil, 1963a:835
Lundbeck, 1927:466; Herting, 1960:61;
Mesnil, 1963a:835
Mesnil, 1963a:835

Ceromya cibdela (Villeneuve)

Stylochilus sp.

Sphingidae

New record (CNC)

Ceromya ?fergusoni (Bezzi)
Unidentified sp.

Geometridae

Crosskey, 1973:176

Ceromya luleicornis (Curran)
Unidentified sp.

Nudaurelia belina

Satumiidae

Curran, 1933c: 162
Curran, 1933c: 162

Ceromya Ontario (Curran)

Ennomos subsignarius Hiibner

Geometridae

Amaud, 1978:55

Ceromya palloris (Coquillett)

Drepana arcuata Walker
Drepana bilineata Packard

Drepanidae

Drepanidae

Amaud, 1978:55
Amaud, 1978:55

Ceromya near palloris (Coquillett)

Ennomos subsignarius Hiibner
Eugonobapta nivosaria Guenee

Geometridae

Geometridae

Amaud, 1978:55
Amaud, 1978:55

Ceromya patellicornis Mesnil
Callopistria repleta Walker

Noctuidae

Crosskey, 1976a: 291

Ceromya pruinosa Shima

Dendrolimus undans flaveola Motsch.

Lasiocampidae

Shima, 1973:155

Ceromya silacea (Meigen)

Erastria fasciana L.

Noctuidae

Herting, 1960:61; Mesnil, 1963a:839

Ceromya spp. (Australia)

Ectropis excursaria Guenee
Chrysodeixis argentifera Guenee
Euplexia nigerrima Guenee

Geometridae

Noctuidae

Noctuidae

Cantrell, 1986:259
Cantrell, 1986:259
Cantrell, 1986:259

(Continued on next page)

Systematics of the Genus Group Taxa of the Siphonini

163

Table 2 (continued)

SIPHONINE & HOST SPECIES

FAMILY

REFERENCE

Homoeosoma vagella Zeller

Pyralidae

Cantrell, 1986:259

Goniocera io (Aldrich)

Malacosoma americanum Fab.

Lasiocampidae

Amaud, 1978:126

Malacosoma disstria Hiibner

Lasiocampidae

Amaud, 1978:126

Goniocera schistacea (B.B.)

Malacosoma castrensis L.

Lasiocampidae

Herting, 1960:119; Mesnil, 1963:801

Goniocera versicolor (Fallen)

Malacosoma neustria L.

Lasiocampidae

Herting, 1960:120; Mesnil, 1963:803

Peribaea argentifrons (Malloch)

Copromorpha prasinochroa Meyrick

Copromorphidae

Crosskey, 1973:176; Chadwick &
Nikitin, 1985:592

Homoeosoma vagella Zeller

Pyralidae

Crosskey, 1973:176

Peribaea fissicornis (Strobl)

Angi rona prunaria L.

Geometridae

Herting, 1960:59; Mesnil, 1963a:807

Boarmia bistortata Goeze

Geometridae

Herting, 1960:59; Mesnil, 1963a:807

Boarmia consortaria Fabr.

Geometridae

Herting, 1960:59; Mesnil, 1963a:807

Bupalus piniarius L.

Geometridae

Herting, 1960:59; Mesnil, 1963a:807

Himera pennaria L.

Geometridae

Herting, 1960:59; Mesnil, 1963a:807

Ourapteryx sambucaria L.

Geometridae

Herting, 1960:59; Mesnil, 1963a:807

Phigalia pedaria Fabr.

Geometridae

Herting, 1960:59; Mesnil, 1963a:807

Poecilocampa populi L.

Lasiocampidae

Herting, 1960:59; Mesnil, 1963a:807

Peribaea hyalinata (Malloch)

Hyblaea puera Cramer

Hyblaeidae

Crosskey, 1976a:291

Pyrausta machoeralis Walker

Pyralidae

Crosskey, 1976a:291

Peribaea orbata (Wiedemann)

Acantholeucania loreyi Dup.

Noctuidae

Crosskey, 1973:176

Aedia leucomelas

Noctuidae

Shima, 1981:450

“Cirphis” sp.

Noctuidae

Crosskey, 1976a:291

Earias sp(p).

Noctuidae

Mesnil, 1963a:806

Heliothis armigera Hiibner

Noctuidae

Shima, 1981:450; Crosskey, 1976a:291

Heliothis sp.

Noctuidae

Crosskey, 1973:176

Laphygma sp(p).

Noctuidae

Mesnil, 1963a:806

Leucania separata

Noctuidae

Shima, 1981:450

Leucania venalba Moore

Noctuidae

Shima, 1981:450; Crosskey, 1976a:291

Mythimna convecta Walker

Noctuidae

Broadley, 1986:61

Platysenta dolorosa Walker

Noctuidae

Cantrell, 1986:259

Prodenia sp(p).

Noctuidae

Mesnil, 1963a:806

Pseudaletia unipuncta Haworth

Noctuidae

Shima, 1981:450; Crosskey, 1973:176
& 1976a:291

Spodoptera exempta Walker

Noctuidae

Crosskey, 1973:176

Spodoptera exigua Hiibner

Noctuidae

Shima, 1981:450; Crosskey, 1976a:291

Spodoptera littoralis Boisd.

Noctuidae

Kugler, 1979:56

(Continued on next page)

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164

O’Hara

Table 2 (continued)

SIPHONINE & HOST SPECIES FAMILY REFERENCE

Spodoptera litura Fabr.

Noctuidae

Shima, 1981:450; Crosskey, 1973:176
& 1976a:291; Jayanth & Nagarkatti,
1984:77

Spodoptera mauritia Boisd.

Noctuidae

Shima, 1981:450; Crosskey, 1976a:291

Spodoptera sp(p).

Noctuidae

Mesnil, 1963a:806

Hedylepta indicata

Pyralidae

Shima, 1981:450

Peribaea palaestina (Villeneuve)

Spodoptera exigua Hiibner

Noctuidae

Kugler, 1979:56

Spodoptera littoralis Boisd.

Noctuidae

Kugler, 1979:56

Peribaea plebeia (Malloch)

Earias huegeli Rogenhofer

Noctuidae

Crosskey, 1973:176

Peribaea sp. ? plebeia (Malloch)

? Anthela sp.

Anthelidae

Crosskey, 1973:176; Chadwick &
Nikitin, 1985:592

Peribaea stiglinae (Bezzi)

Stiglina superior Butl.

Thyrididae

Bezzi, 1928:204

Peribaea suspecta (Malloch)

Earias vittella Fabr.

Noctuidae

Crosskey, 1976a: 291

Earias sp.

Noctuidae

Curran, 1928:237; Crosskey, 1976a:291

Peribaea tibialis (Robineau-Desvoidy)

Ennomos autumnaria Womb.

Geometridae

Hubenov, 1985:29

Ennomos quercinaria Hufn.

Geometridae

Hubenov, 1985:29

Prosopolopha jourdanaria Vill.

Geometridae

Herting, 1960:59; Mesnil, 1963a:813

Lasiocampa grandis Stgr.

Lasiocampidae

Herting, 1960:59; Mesnil, 1963a:813;
Kugler, 1979:56

Orgyia dubia Tausch.

Lyman triidae

Herting, 1960:59; Mesnil, 1963a:813;
Kugler, 1979:55

Anarta myrtilli L.

Noctuidae

Herting, 1960:59; Mesnil, 1963a:813

Asticta pastinum Tr.

Noctuidae

Mesnil, 1963a:813

Spodoptera exigua Hiibner

Noctuidae

Kugler, 1979:56

Yponomeuta malinellus Z.

Yponomeutidae

Hubenov, 1985:29

Peribaea sp. (Australia)

Spodoptera exigua Hiibner

Noctuidae

Cantrell, 1986:259

Crocidosema plebejana
Zeller

S. ( Aphantorhaphopsis ) mallochiana
(Gardner)

Tortricidae

Cantrell, 1986:259

Pelopidas mathias Fabr.

Hesperiidae

Crosskey, 1976a: 291

“Turmeric skipper”

Hesperiidae

Crosskey, 1976a: 291

“Ginger lily leaf-roller”

Hesperiidae

Crosskey, 1976a:291

(Continued on next page)

Systematics of the Genus Group Taxa of the Siphonini

165

Table 2 (continued)

SIPHONINE & HOST SPECIES

FAMILY

REFERENCE

“Rice leaf-roller skipper”

Hesperiidae

Crosskey, 1976a:291

S. (Aphantorhaphopsis) norma (Malloch)

Mythimna convecta Walker

Noctuidae

Crosskey, 1973:176; Chadwick &
Nikitin, 1985:590

S. ( Aphantorhaphopsis ) samarensis (Vill.)

Porthetria dispar L.

Lyman triidae

New record (USNM)

S. (Aphantorhaphopsis) selecta (Pand.)

Paidia murina Hiibner

Arctiidae

Herting, 1978:7

S. ( Aphantorhaphopsis ) siphonoides
(Strobl)

Delilinia pusaria L.

Xanthorhoe biriviata Borkh.

Geometridae

Geometridae

Herting, 1960:62; Mesnil, 1964: 850
Herting, 1960:62; Mesnil, 1964: 850

S. ( Baeomyia ) juniperi (O’Hara)

Semiothisa trivata B. & McD.

Geometridae

O’Hara, 1984:1395

S. (Baeomyia) xanthogaster (O’Hara)

Semiothisa sexmaculata Packard
Semiothisa spp.

Unidentified spp.

Geometridae

Geometridae

Geometridae

O’Hara, 1984:1395
O’Hara, 1984:1395
O’Hara, 1984:1395

S. (Ceranthia) abdominalis (R.-D.)

Cosymbia annulata Schultze
Cosymbia orbicularia Hiibner

Cosymbia pendularia Cl.

Cosymbia porata F.

Cosymbia ruficiliaria H.S.

Geometridae

Geometridae

Geometridae

Geometridae

Geometridae

Herting, 1960:62; Mesnil, 1963a: 841
Herting, 1960:62 & 1966:6; Mesnil,
1963a: 841
Herting, 1966:6

Herting, 1960:62 & 1966:6; Mesnil,
1963a: 841
Herting, 1966:6

S. (Ceranthia) lichtwardtiana (Villeneuve)

Acasis viretata Hiibner

Geometridae

Herting, 1966:6

S. (Ceranthia) pallida (Herting)

Eupithecia denotata Hiibner

Geometridae

Herting, 1960:62; Mesnil, 1963a: 843

S. (Ceranthia) tristella (Herting)
Eupithecia silenata Standf.
Eupithecia undata Frr.

Geometridae

Geometridae

Herting, 1966:6
Herting, 1966:6

S. (Pseudosiphona) brevirostris Coquillett

Oidaematophorus homodactylus Walker

Pterophoridae

Amaud, 1978:452 (doubtful record)

S. (Pseudosiphona) spp.

Chloropteryx sp.

Geometridae

New record (USNM)

(Continued on next page)

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O’Hara

Table 2 (continued)

SIPHONINE & HOST SPECIES

FAMILY

REFERENCE

Unidentified sp.

Lycaenidae

New record (USNM)

S. ( Siphona ) collini Mesnil

Cerapteryx graminis L.

Noctuidae

Andersen, 1982:157

Euxoa temera Hiibner

Noctuidae

Andersen, 1982:157

Euxoa tritici L.

Noctuidae

Andersen, 1982:157

S. ( Siphona ) confusa Mesnil

Mythimna littoralis Curtis

Noctuidae

Andersen, 1982:157

Phlogophora meticulosa L.

Noctuidae

Andersen, 1982:157

5. ( Siphona ) cristata (Fabricius)

Anticollix sparsata Treitschke

Geometridae

Andersen, 1982:157

Erannis defoliaria Clerck

Geometridae

Andersen, 1982:157

Antitype chi L.

Noctuidae

Andersen, 1982:157

Caradrina morpheus Hufnagel

Noctuidae

Andersen, 1982:157

Hadena bicruris Hufnagel

Noctuidae

Andersen, 1982:157

Lacanobia suasa Denis & Schiff.

Noctuidae

Andersen, 1982:157

Lacanobia oleracea L.

Noctuidae

Andersen, 1982:157

Mamestra brassicae L.

Noctuidae

Andersen, 1982:157

? Mythimna f err ago Fabr.

Noctuidae

Andersen, 1982:157

?Mythimna littoralis Curtis

Noctuidae

Andersen, 1982:157

? Mythimna obsoleta Hiibner

Noctuidae

Andersen, 1982:157

?Phlogophora meticulosa L.

Noctuidae

Andersen, 1982:157

Smerinthus ocellata L.

Sphingidae

Andersen, 1982:157

S. ( Siphona ) geniculata (De Geer)

Ceramica pisi L.

Noctuidae

Andersen, 1982:158 (doubtful record)

Tipula fulvipennis De Geer

Tipulidae

Andersen, 1982:158

Tipula lateralis Meigen

Tipulidae

Andersen, 1982:157

Tipula maxima Poda

Tipulidae

Andersen, 1982:158

Tipula montium Egger

Tipulidae

Andersen, 1982:157

Tipula oleracea L.

Tipulidae

Andersen, 1982:157

Tipula paludosa Meigen

Tipulidae

Carter^ ai, 1981:257; Andersen,
1982:157

Tipula subnodicornis Zett.

Tipulidae

Andersen, 1982:157

Tipula vittata Meigen

Tipulidae

Andersen, 1982:158

S. ( Siphona ) maculata Staeger

Euxoa obelisca Denis & Schiff.

Noctuidae

Andersen, 1982:157

Ochropleura candelisequa Denis & Schiff.

Noctuidae

Andersen, 1982:157

S. ( Siphona ) nigricans (Villeneuve)

Tipula irrorata Macquart

Tipulidae

Herting, 1967a:9; Andersen, 1982:158

S. ( Siphona ) pseudomaculata Blanchard

Faronta albilinea Hiibner

Noctuidae

Blanchard, 1963:252; Guimaraes,
1977:74

(Continued on next page)

Systematics of the Genus Group Taxa of the Siphonini

167

Table 2 (continued)

SIPHONINE & HOST SPECIES

FAMILY

REFERENCE

S. ( Siphona ) setosa Mesnil

Eupithecia succenturiata L.
Allophyes oxyacantae L.

Geometridae

Noctuidae

Andersen, 1982:157
Andersen, 1982:157

S. ( Siphonopsis ) brasiliensis (Townsend)
Rachiplusia nu Guenee
Syngrapha gammoides Blanchard

Noctuidae

Noctuidae

Guimaraes, 1977:74
Guimaraes, 1977:74

S. ( Siphonopsis ) conata (Reinhard)

Phryganidia californica Packard

Dioptidae

Amaud, 1978:458

S. ( Siphonopsis ) plusiae Coquillett

Phryganidia californica Packard
Autographa californica Speyer
Trichoplusia ni Hiibner
Unidentified spp.

Dioptidae

Noctuidae

Noctuidae

Noctuidae

Amaud, 1978:458
Amaud, 1978:459
Amaud, 1978:459
Amaud, 1978:459

Siphona s. 1. (New World) sp. 7

Cladara limitaria Walker
Stenoporpia sp.

Panthea portlandia Grt.

Geometridae

Geometridae

Noctuidae

New record (CNC)
New record (CNC)
New record (CNC)

Siphona s. 1. (New World) sp. 8

Ectropis crepuscularia Denis & Schiff.
Epirrita autumnata Bkh.

Melanolophia imitata Walker

Geometridae

Geometridae

Geometridae

New record (CNC)
New record (CNC)
New record (CNC)

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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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170

O’Hara

Book Review

Table 3. Tachinid species examined for outgroup comparison with the Siphonini.

TRIBE AND SPECIES
Ernestiini

DISTRIBUTION

Eloceria delecta (Meigen)
Synactia parvula (Rondani)

Europe

Europe

Triarthriini

Triarthria setipennis (Fallen)

Europe

Aphriini

Aphria ocypterata Townsend
Solieria pacifica (Meigen)
Solieria sp.

Nearctic

western Palearctic
Arizona (USA)

Leskiini

Atylostoma towadensis (Matsumura)
Drepanoglossa lucens Townsend
Genea brevirostris (James)

Genea texensis (Townsend)

Genea robertsonii (Townsend)
Leskia aurea (Fallen)

Leskia depilis (Coquillett)

Leskia sp.

Trochiloleskia loriola (Reinhard)

eastern Palearctic & SE Asia
Nearctic
eastern Nearctic
Nearctic

southern Nearctic
Palearctic
eastern Nearctic
New Mexico (USA)
southern Nearctic

Clausicellini

Clausicella sp.

Crocinosoma cornuale Reinhard
Dolichopalpellus mirabilis Townsend
Epicoronimyia mundelli (Blanchard)
Ginglymia acirostris Townsend
Ginglymia johnsoni (Coquillett)
Phantasiomyia sp.

Spathipalpus philippi Rondani

Arizona (USA)
southern Nearctic
SE Brazil
Argentina & Chile
Nearctic
Nearctic

California (USA)
Argentina & Chile

Neaerini

Camposodes evanescens Cortes
Neaera laticornis (Meigen)
Neaera leucoptera (Johnson)
Neaera robertsonii (Townsend)
Phytomyptera setigera (Thomson)
Phytomyptera sp.

Chile

Palearctic

central and eastern Nearctic
central and eastern Nearctic
western Nearctic
Arizona (USA)

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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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).

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

176

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]

© polytypic

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]

© most species with state O, few with ©or#

Articulation between surstylus and epandrium [character 52]

O membranous [state M]

• sclerotized (fused) [state F]

© polytypic

©’ sclerotized in one specie ‘

Shape of cerci in profile [character 54]

O not sharply inflexed

• sharply inflexed [state I]

© polytypic

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

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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).

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Fig. 169. Spinules on pregonite
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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).

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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

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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)

200

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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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

Publication of Quaestiones Entomologicae was started in 1965 as part of a
memorial project for Professor E.H. Strickland, the founder of the Department
of Entomology at the University of Alberta in 1922.

It is intended to provide prompt relatively low cost publication for compre-
hensive accounts of entomological research of greater than average length.
However, shorter papers about insects in the Prairie Provinces of Canada are
acceptable. Page charges are normally levied, the rate determined by printer’s
charges. For information about current page charges, consult the Editor.

Copy for all types of papers should conform to the Style Manual for
Biological Journals, published by the American Institute of Biological Sci-
ences, Second Edition, 1964, except that title of periodicals should be given in
full. For style of taxonomic papers, the Editor should be consulted. Two copies
of a manuscript are requested. All manuscripts will be reviewed by referees.

Abstracts are required, one in English and one in another language, prefer-
ably French.

Copy for illustrations must accompany the manuscript, and be of such char-
acter as to give satisfactory reproduction at page size (less 1/2 inch, or 1.2 cm
on plates of full page size [7X4 1/4 inches or 17.8 X 10.7 cm]). Reprints must
be ordered when proofs are returned, and will be supplied at cost.

Subscription rates are the same for institutions, libraries and individuals,
$20.00* per volume of four issues, normally appearing at quarterly intervals;
single issues $5.00**. Back volumes and issues are available at the same cost.
These prices supersede those previously indicated, and are subject to change as
required by inflationary pressure on the value of money.

Communications regarding subscriptions should be addressed to the
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* Same rate in US$ for non-Canadian subscriptions.

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Published quarterly by;

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Return Undeliverable mail to the address above. Return Postage Guaranteed

Issued September 1989

TROSIEME CONFERENCE INTERNATIONALE DES
ENTOMOLOGISTES D’EXPRESSION FRANCAISE
Gembloux, 9 - 14 juillet 1990

Cette conference - ouverte a tous les entomologistes, arachnologues,
acarologues, professionnels ou amateurs de toutes disciplines fondamentales ou
appliquees - se tiendra a Gembloux (Belgique) a la Faculte des Sciences
agronomiques de 1’Etat. Elle donnera lieu a des conferences plenieres,
communications, tableaux de demonstrations (’’posters”), ateliers. Elle comprendra
aussi des sessions speciales dans d’autres villes de Belgique et sera suivie
d’excursions.

Les conferences, les differentes communications et ateliers devront s’inspirer du
theme general de la conference:

Methodes et responsabilites des entomologistes d?aujourd’hui

Les communications s’inscriront dans le cadre de sections qui ont ete definies
par le Comite organisateur. Elies ne sont pas encore definitives. En voici la liste
provisoire:

1. Entomologie culturelle; 2. Entomologie medicate, veterinaire et medico-legale; 3.
Insectes, amateurs et grand public; 4. Entomologie agricole; 5. Entomologie agricole
dans les pays en voie de developpement; 6. Role des arthropodes dans la qualite et la
fertilite des sols; 7. Insectes sociaux; 8. Mediateurs chimiques; 9. Entomologie des
milieux forestiers; 10. Tendances actuelles de la systematique; 11. Zoogeographie;
12. Systemes tegumentaires (cuticule, mue et metamorphoses,...).

La langue officielle de la conference (exposes thematiques, communications et
tableaux de demonstrations) sera le fran^ais. Cependant les participants auront
Lentiere liberte de s ’exprimer dans la langue de leur choix au cours des discussions
qui suivront les exposes ex-cathedra et lors des presentations et discussions des
affiches.

Si vous souhaitez participer ou assister aux travaux de Tune ou 1’autre des
sessions de cette conference, veuillez rapidement prendre contact ave le Secretaire
general:

Monsieur Charles VERSTRAETEN
Zoologie generate et appliquee
Faculte des Sciences agronomiques de l’Etat
B-5800 Gembloux (P ique).

Le Comite organisateur

P.S.: Veuillez diffuser cette circulate autour de vous.

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 3 1989

CONTENTS

Freitag and Bames-Classification of Brazilian Species of Cicindela and
Phylogeny and Biogeography of Subgenera Brasiella, Gaymara new
subgenus, Plectographa and South American Species of Cylindera

(Coleoptera: Cicindelidae) 241

Book Review-Schwalm, F.E. 1988. Insect Morphogenesis 387

CLASSIFICATION OF BRAZILIAN SPECIES OF CIC1NDELA AND
PHYLOGENY AND BIOGEOGRAPHY OF SUBGENERA BRASIELLA ,
GAYMARA NEW SUBGENUS, PLECTOGRAPHA AND SOUTH AMERICAN
SPECIES OF CYLINDERA (COLEOPTERA: CICINDELIDAE)

Richard Freitag
Barbara L. Barnes
Department of Biology
Lake head University
Thunder Bay, Ontario
Canada P7B 5 El

Quaestiones Entomologicae
25:241-386 1989

ABSTRACT

Brazilian tiger beetles of the tribes Cicindelini, Ctenostomatini, and
Megacephalini, subtribes Cicindelina, Odontocheilina and Prothymina, and taxa of
the genus Cicindela are distinguished in identification keys. Diagnoses based on
adult characteristics, including genitalia of males and females, new for most
species, are provided for four subgenera, 12 species groups, and 29 species of
Brazilian Cicindela. Species sections consist of nomenclature, recognition,
synonyms and types, description, geographical variation, relationships, habitat and
period of activity, geographical distribution, lists of localities of examined
specimens, distribution map, and figures illustrating taxonomically important
external and internal structures.

A classification of 61 Brazilian and related Neotropical species of Cicindela is
established based on a reconstructed phylogeny employing the methods of Hennig.
Brazilian taxa are arranged in sequence according to the reconstructed phylogeny.
The four subgenera, species groups, and species recognized are: subgenus Brasiella
Rivalier, argentata group- C. argentata Fabricius, C. obscurella Klug, C. pretiosa
Dokhtouroff; aureola group- C. amaenula Chaudoir, C. aureola Klug; misella
group- C. dolosula Rivalier, C. misella Chaudoir; minarum group- C. hamulipenis
Horn, C. brevipalpis Horn, C. banghaasi Horn, C. minarum Putzeys; new subgenus
Gaymara (type species: C. chlorosticta Kollar); chlorosticta group- C. chlorosticta
Kollar, C. staudingeria Horn, C. nigroreticulata Horn, new species C.
paranigroreticulata ( type locality.— Tramandai, Rio Grande do Sul); anulipes group-
C. anulipes Horn; subgenus Plectographa Rivalier; suturalis group- C. suturalis
Fabricius, C. nivea Kirby; melaleuca group- C. melaleuca Dejean, C. patagonica
Brulle; nigrovittata group- C. nigrovittata Horn; apiata group- C. apiata Dejean;
subgenus Cylindera Westwood; morio group- C. kollari Gistl, C. confluentesignata

242

Freitag and Barnes

Horn, C. morio Klug, C. marquardti Horn; friedenreichi group- C. piligera Horn, C.
obsoletesignata Horn, C. friedenreichi Dokhtouroff. Seven new synonymies are
proposed with the senior synonym of each combination listed first: C. argentata
argentata Fabricius = Brasiliella pseudoargentata Mandl; C. obscurella Klug = C.
obscurella constricta Rivalier = Brasiella chrysocollis Mandl = Brasiella pallidipes
Mandl; C. suturalis Fabricius = C. trifasciata boliviana Mandl.

A reconstructed phytogeny of Cicindela subgenera which have Brazilian species
indicates the following relationships: Brasiella as sister group of

Gaymara/Cylindera lineage, Gaymara as sister group of Plectographa/Cylindera
lineage, and Plectographa as sister group o/Cylindera.

Relationships among species groups in each subgenus are indicated as follows:
for Brasiella, argentata group sister of ancestral lineage of sister groups
viridicollis/hemichrysea, aureola group sister of cubana/pretiosa lineage, misella
group sister of cubana/horioni lineage, monobasic stamatovi group taxon of
uncertain relationships, and minarum group sister of cubana/venezuelensis lineage;
for Gaymara, chlorosticta group sister of monobasic anulipes group; for
Plectographa, monobasic halophila group taxon of uncertain relationships, suturalis
group sister o/melaleuca group, monobasic nigrovittata group as taxon of uncertain
relationships, apiata group sister of siccalacicola/nigrovittata lineage; and for
Cylindera, morio group as sister of friedenreichi group.

Relationships among species of Brasiella are: viridicollis group, C. cubana Leng
and Mutchler sister of ancestral lineage of sister species C. acuniai Mutchler/C.
viridicollis Dejean, C. wickhami Horn sister of cubana/viridicollis lineage;
hemichrysea group, C. hemichrysea Chevrolat sister of ancestral lineage of sister
species C. mendicula Rivalier! C. sphaerodera Rivalier; argentata group, C. argentata
Fabricius sister of venustula/pretiosa lineage, C. venustula Gory sister of ancestral
lineage of sister species C. obscurella Klug/C. pretiosa Dokhtouroff; aureola group-
C. rivalieri Mandl sister of amaenula/horioni lineage, C. amaenula Chaudoir sister
of ancestral lineage of sister species C. aureola Klug/C. horioni Mandl; misella
group, ancestral lineage of sister species C. dolosulaffinis Mandl/C. tippmanni
Mandl sister of dolosula/venezuelensis lineage, C. venezuelensis Mandl sister of
ancestral lineage of sister species C. dolosula Rivalier! C. misella Chaudoir,
stamatovi group, C. stamatovi Sumlin relationships uncertain, minarum group,
hamulipenis/banghaasi lineage sister of minarum/balzani lineage, C. banghaasi
Horn sister of ancestral lineage of sister species C. hamulipenis Horn/C. brevipalpis
Horn, C. balzani Horn sister of minarum/insularis lineage, C. minarum Putzeys
sister of nebulosa/insularis lineage, C. nebulosa Bates sister of ancestral lineage of
sister species C. mandli Brouerius van Nidek/C. insularis Brouerius van Nidek.

Relationships among Gaymara species are: anulipes group, C. anulipes Horn
sister to chlorosticta/paranigroreticulata lineage; chlorosticta group, ancestral
lineage of sister species C. chlorosticta Kollar/C. staudingeria Horn sister of
ancestral lineage of sister species C. nigroreticulata Horn/C. paranigroreticulata

Brazilian Species of Cicindela

243

n.sp.

Relationships among Plectographa species: halophila group, C. halophila Sumlin
relationships uncertain; suturalis group, C. siccalacicola Sumlin sister of
sinuosa/nahuelbutae lineage, ancestral lineage of sister species C. sinuosa Brulle/C.
suturalis Fabricius sister of hirsutifrons/nahuelbutae lineage, ancestral lineage of
sister species C. hirsutifrons Sumlin/C. nivea Kirby sister of ancestral lineage of
sister species C. ramosa Brulle/C. nahuelbutae Pena; melaleuca group, C. mixtula
Horn relationships uncertain, ancestral lineage of sister species C. ritsemai Horn/C.
drakei Horn sister o/melaleuca/chiliensis lineage, ancestral lineage of sister species
C. melaleuca Dejean/C. patagonica Brulle sister of ancestral lineage of sister
species C. gormazi Reed/C. chiliensis Audouin and Brulle; nigrovittata group, C.
nigrovittata Horn relationships uncertain; apiata group, C. eugeni Castelnau sister
of C. apiata Dejean.

Relationships among South American Cylindera species: morio group, ancestral
lineage of sister species C. kollari Gistl/C. malaris Horn sister of
confluentesignata/marquardti lineage, ancestral lineage of sister species C.
confluentesignata HornIC. granulipennis Bates sister of C. morio Klug/C.
marquardti Horn; friendenreichi group, C. friedenreichi Dokhtouroff sister of
ancestral lineage of sister species C. piligera Horn/C. obsoletesignata Horn.

The subgenera Brasiella, Gaymara, Plectographa and Cylindera are hypothesized
to have originated in western Gondwana prior to the formation of South America.
Neotropical diversification within subgenera occurred mainly in the northern half of
South America during the Tertiary and Pleistocene. Major centers of species
concentration of Brasiella, Gaymara, and Cylindera are in open country in the
eastern and central Brazilian highlands and that of Plectographa in northern
Argentina. These centers appear to have been long-standing and the chief sources of
dispersal during hospitable climatic and geophysical periods, and into which taxa
retreated during climatically hostile periods. Important centers of taxa
diversification are eastern and southeastern Brazil, northern Argentina and the
Amazon River basin. Vegetational changes in the Amazon basin during the
Pleistocene appear to have been the major causes of taxa formation.

RESUME

Les coleopteres Bresiliens de la tribue des Cicindelini, Ctenostomatini, et Megacephalini, du sous-tribue
des Cicindelina, des Odontocheilina et des Prothymina, et de la taxa du genre Cicindela sont classes en
groupes distinctes. Les diagnoses, bases sur les caracteristiques des adultes, males et femelles inclus, sont
nouveaux pour la plupart des especes et seront donnes pour quatre sous-especes, 12 groupes d’ especes et 29
especes de Cicindela Bresiliennes. Le classement des especes est base, sur une nomenclature, une
identification, des listes de synonymes et de types, une description, une variation geographique, les
relations, 1’ habitat et les periodes d’activites, la distribution geographique , les differentes localisations des
especes examinees, une carte des repartitions, et des chiffres illustrant de fagon taxonamicale les
importantes structures exterieurs et interieurs.

Une classification de 61 especes de Cicindela Bresiliennes et Neotropicales apparentees est etablie
grace a une phylogenie reconstitute en utilisant les methodes de Hennig. Les taxa Bresiliennes sont

Quaest. Ent., 1989, 25 (3)

244

Freitag and Barnes

regrouptes selon la phytogeny reconstitute. Les quartre sous-genres (groupes d especes), et les
organisations des especes sefont comme suit: sous-genre Brasiella Rivalier, (groupe argentataj, C. argentata
Fabricius, C. obscurella Klug, C. pretiosa Dokhtouroff, ( groupe aureola), C. amaenula Chaudoir, C. aureola
Klug, (groupe misella), C. dolosula Rivalier, C. misella Chaudoir, (groupe minarum), C. hamulipenis Horn,
C. brevipalpis Horn, C. banghaasi Horn, C. minarum Putzeys; nouveau sous-genre Gaymara (type
d’ especes.— C. chlorosticta Kollar), (groupe chlorosticta), C. chlorosticta Kollar, C. staudingeria Horn, C.
nigroreticulata Horn, la nouvelle espece C. paranigroreticulata (type localite.-Tramandai, Rio Grande do
Sul), (groupe anulipes), C. anulipes Horn; sous-genre Plectographa Rivalier, (groupe sutural is), C. suturalis
Fabricius, C. nivea Kirby, (groupe melaleuca), C. melaleuca Dejean, C. patagonica Brullt, (groupe
nigrovittata), C. nigrovittata Horn, (groupe apiata), C. apiata Dejean; sous-genre Cylindera Westwood,
(groupe morio), C. kollari Gistl, C. confluentesignata Horn, C. morio Klug, C. marquardti Horn, (groupe
friedenreichi), C. piligera Horn, C. obsoletesignata Horn, C. friedenreichi Dokhtouroff. Sept nouveaux
sy steme s de synonymes sont proposes avec I’ancien synonyme de chaque combinaisons donates auparavent:
C. argentata argentata Fabricius = Brasiliella pseudoargentata Mandl; C. obscurella Klug = C. obscurella
constricta Rivalier = Brasiella chrysocollis Mandl = Brasiliella pallidipes mandl; C. suturalis Fabricius = C.
trifasciata boliviana Mandl.

Une phylogtnie reconstitute des sous-genres Cicindela qui comprant les especes Brtsilienne indique les
relations suivantes: Brasiella comme groupe soeur de la lignte Gaymara/Cylindera, Gaymara comme groupe
soeur de la lignte Plectographa/Cylindera, et Plectographa comme group soeur de Cylindera.

Les relations entre les groupes d’ especes dans chaque sous-genres sont indiqutes comme suit: la
Brasiella, le groupe argentata soeur de la lignte ancestrale des groupes soeurs viridicollis/hemichrysea, le
groupe aureola soeur de la lignte cubana/pretiosa, le groupe misella soeur de la lignte cubana/horioni, le
groupe monobasique stamatovi taxon de relations incertaines, et le groupe minarum apparentt de la lignte
cubana/venezuelensis; la Gaymara, le groupe chlorosticta soeur du groupe monobasique anulipes; la
Plectographa, le groupe monobasique halophila taxon de relations incertaines, le groupe suturalis soeur du
groupe melaleuca, le groupe monobasique nigrovittata taxon de relations incertaines, le groupe apiata soeur
de la lignte siccalacicola/nigrovittata; et la Cylindera, le groupe morio soeur du groupe friedenreichi.

Les relations entre les especes Brasiella sont: (groupe viridicollis), C. cubana Leng et Mutchler soeur de
la lignte ancestrale des especes soeurs C. acuniai Mutchler/C. viridicollis Dejean, C. wickhami Horn soeur
de la lignte des cubana/viridicollis; (groupe hemichrysea), C. hemichrysea Chevrolat soeur de la lignte
ancestrale des especes soeurs C. mendicula Rivalier! C. sphaerodera Rivalier; (groupe argentata), C.
argentata Fabricius soeur de la lignte des venustula/pretiosa, C. venustula Gory soeur de la lignte
ancestrale des especes soeurs C. obscurella Klug/C. pretiosa Dokhtouroff; (groupe aureola, C. rivalieri
Mandl soeur de la lignte des amaenula/horioni, C. amaenula Chaudoir soeur de la lignte ancestrale des
especes soeurs C. aureola Klug/C. horioni Mandl; (groupe misella), la lignte ancestrale des especes soeurs
C. dolosulaffinis Mandl! C. tippmanni Mandl soeur de la lignte des especes soeurs C. dolosula Rivalier! C.
misella Chaudoir, (groupe stamatovi), C. stomatovi Sumlin relations incertaines, (groupe minarum), la
lignte hamulipenis/banghaasi soeur de la lignte des minarum/balzani, C. banghaasi Horn soeur de la lignte
ancestrale des especes soeurs C. hamulipenis Horn/C. brevipalpis Horn, C. balzani Horn soeur de la lignte
des minarum/insularis, C. minarum Putzeys soeur de la lignte nebulosa/insularis, C. nebulosa Bates soeur de
la lignte ancestrale des especes soeurs C. mandli Brouerius van Nidek/C. insularis Brouerius van Nidek.

Les relations entre les espece Gaymara sont: (groupe anulipes), C. anulipes Horn soeur de la lignte des
chlorosticta/paranigroreticulata; (groupe chlorosticta, la lignte ancestrale des especes soeurs C. chlorosticta
Kollar/C. staudingeria Horn soeur de la lignte ancestrale des especes soeurs C. nigroreticulata Horn/C.
paranigroreticulata n.sp.

Les relations entre les especes Plectographa; (groupe halophila), C. halophila Sumlin relations
incertaines; (groupe suturalis), C. siccalacicola Sumlin soeur de la lignte des sinuosa/nahuelbutae, la lignte
ancestrale des especes soeurs C. sinuosa Brullt/C. suturalis Fabricius soeur de la lignte des
hirsutifrons/nahuelbutae, la lignte ancestrale des especes soeurs C. hirsutifrons Sumlin! C. nivea Kirby soeur
de la lignte ancestrale des especes soeurs C. ramosa Brullt/C. nahuelbutae Pena; (groupe melaleuca), C.
mixtula Horn relations incertaines, la lignte ancestrale des especes soeurs C. ritsemai HornIC. drakei Horn
soeurs de la lignte des melaleuca/chiliensis, la lignte ancestrale des espece soeurs C. melaleuca Dejean/C.
patagonica Brullt soeurs de la lignte ancestrale des especes soeurs C. gormazi ReedIC. chiliensis Audouin
et Bruellt; (groupe nigrovittata), C. nigrovittata Horn relations incertaines; (groupe apiata), C. eugeni
Castelnau soeur de C. apiata Dejean.

Brazilian Species of Cicindela

245

Les relations entre les Cylindera d’Amerique du sud: (groupe mono, la lignee ancestrale des especes
soeurs C. kollari Gistl/C. malaris Horn soeur de la lignee des confluentesignata/marquardti, la lignee
ancestrale des especes soeurs C. confluentesignata Horn/C. granulipennis Bates soeur des C. mono Klug/C.
marquardti Horn; (groupe friendenreichi), C. friedenreichi Dokhtouroff soeur de la lignee ancestrale des
especes soeurs C. piligera Horn/C. obsoletesignata Horn.

Selon une hypothese les sous-genres Brasiella, Gaymara, Plectographa et Cylindera proviennent de
I’ouest du Gondwana avant la formation de VAmerque du sud. La diversification Neotropicale d’ une meme
sous-genres se trouve surtout dans la partie nord de VAmerique du sud pendant 1’ ere Tertiaire et le
Pleistocene. Les principaux centers de concentration des Brasiella, Gaymara, et Cylindera sont la pleine
campagne et lest et central pays de montagnes Bresiliennes et ceux du Plectographa en Argentina du nord.
Ces centers semblent avoir ete pendant longtemps les principaux points de dispersion des periodes
climatiques et geophysique hospitalieres et de points de retraite pour les taxa durant les periodes hostiles.
D’ important centers de diversifications des taxa se situent a lest et au sud-est du Bresil, en Argentine du
nord et dans le bassin Amazonien. Les transformations de la vegetation du bassin Amazonien pendant le
Pleistocene semblant etre a I’origine de la transformation des taxa.

TABLE OF CONTENTS

Introduction 246

Materials and Methods 246

Classification 248

Notes about taxonomic characteristics 248

Taxa of the Brazilian Cicindelidae 248

Key to adults of tribes of Brazilian Cicindelidae 248

Key to adults of subtribes and subgenera of Cicindela of Brazilian

Cicindelini 248

Key to the Brazilian species of Cicindela 249

Subgenus Brasiella Rivalier 254

Subgenus Gaymara , new subgenus 278

Subgenus Plectographa Rivalier 289

Subgenus Cylindera Westwood 317

History of South American species of subgenera Brasiella, Gaymara,

Plectographa, and Cylindera 332

Introduction 332

Phylogeny 332

Biogeography 346

Expected geographical patterr 350

Distribution and historical interpretation - subgenera 350

Distribution and historical interpretation - species groups and species 35 1

Geographical history 363

Summary 372

Problems and Predictions 372

Acknowledgements 373

References 374

Index 383

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INTRODUCTION

Brazilian species of Cicindela have not been reviewed as a geographical entity,
though they have been treated as part of broad systematic works. Horn (1915,
1926a, 1938) in studies of the world fauna established a preliminary classification
based mainly on form, colouration, and pubescence of the exoskeleton of adults. He
also included synonymies and geographical distributions. In a ‘catalogue’ on the
evolution of Cicindela , Schilder (1953) placed almost all Brazilian species into one
genus, Cicindosa Motschulsky.

Rivalier (1954, 1955) demonstrated relationships among Neotropical species by
comparisons of the internal sac of the median lobe, and divided Brazilian forms of
Cicindela into Brasiella Rivalier and Cylindera Westwood. Vidal Sarmiento (1966a,
b) also conducted studies of female and male genitalia of genera and species of the
Cicindelidae of Argentina, but did include some taxa from Brazil.

More recently in a synopsis of the genera of Neotropical Carabidae Reichardt
(1977) presented a brief systematic history of Cicindela , and recognized nine
‘genera’ of Rivalier, two of which are principally Brazilian. Sumlin (1979) reviewed
Argentine species of Cicindela in which a few Brazilian members were noted and
included in a key.

Our study was conducted to develop a taxonomic system and hi orical synthesis
for Brazilian and related Neotropical species of Cicindela. The fc lowing account
includes identification keys, descriptions of taxa, classification and evolutionary
history. Determinations of species, species groups, and their relationships, are based
on external structures, male genitalia as described by Rivalier (1954, 1955) and
Vidal Sarmiento (1966b), and female genitalia which are described herein for the
first time for most species. Descriptions of male genitalia not studied by Rivalier
(1954, 1955) are included.

MATERIALS AND METHODS

This study was based on adults of Cicindela , those tiger beetles which occupy
open habitats, such as bare patches of soil in grasslands, roads and footpaths, river
banks, margins of standing fresh water, and sea beaches. About 1200 adult
specimens of Neotropical species of Cicindela were examined, including Horn
types. W.G. Graham and the senior author collected some specimens in the Manaus
area. The great majority of specimens were received on loan from private
collections and institutions, for which the following codens after Arnett and
Samuelson (1969) and Heppner and Lamas (1982) are used in the text.

BMNH British Museum (Natural History), Oomwell Road, London, England
SW7 5BD.

BRI Biosystematics Research Institute >..W. Neatby Bldg. Room 3125,

Brazilian Species of Cicindela

247

CASC

HFHC

ICCM

INPA

IOC

IPZE

LEPC

MMKC

MNHP

MNRJ

MZSP

RRMC

USNM

ZSMS

Research Branch, Ottawa, Ontario, Canada K1A 0C6.

California Academy of Sciences, Golden Gate Park, San Francisco,
California, U.S.A. 941 18.

Henry H. Howden Collection, Department of Biology, Carleton
University, Ottawa, Ontario, Canada K1S 5B6.

Carnegie Museum of Natural History, Section of Entomology, 4400
Forbes Avenue, Pittsburgh, Pennsylvania, U.S.A. 15213.

Instituto Nacional de Pesquisas da Amazonia, Caixa Postal 478, Manuas -
69000, Amazonas, Brazil.

Instituto Oswaldo Cruz, Rio de Janeiro, Brazil.

Akademie der Landwirtschaftswissenschaften der DDR, Institut fur
Pflanzenschutzforschung Zweigstelle, Abt. Taxonomie der Insekten,
Kleinmachnow, Bereich Eberswalde, 13 Eberswalde-Finow 1,
Schicklerstrasse 5, Germany.

Luis E. Pena Collection, P.O. Box 2974, Santiago, Chile.

Michael M. Kaulbars Collection, Department of Biology, Carleton
University, Ottawa, Ontario, Canada K1S 5B6.

Museum National d’Histoire Naturelle, Entomologie, 45 rue de Buffon,
75005 Paris, France.

Museu Nacional, Quinta da Boa Vista, Rio de Janeiro, RJ-20,000, Brazil.
Museu de Zoologie, Universidade de Sao Paulo, Avenida Nazare, 481
(Agencia Ipiranga), 04263 Sao Paulo, SP- Brazil.

Robert R. Murray Collection, Fort Worth, Texas, U.S.A. 76107.

United States National Museum of Natural History, Smithsonian
Institution, Washington, D.C., U.S.A. 20560

Zoologische Staatssammlung, Maria-Ward Strasse lb, D-8000 Munchen
19, Germany.

The various methods and concepts applied in this study have been fairly well
established. Structures examined, drawings and measurements, dissections of the
male and female genitalia, distribution maps and localities, criteria for species and
subspecies, diagnosis of the genus Cicindela and reconstruction of phylogenetic
system relationships based solely on derived characteristics, have been described
elsewhere (Freitag, 1979).

We use special terms that have been applied to features of the male and female
genitalia by Rivalier (1954, 1955), Freitag (1965, 1966, 1972), and Vidal Sarmiento
(1966a, b, 1967), as shown in Figures 34, 36, 46, 72, 108. A new term, “right bar”,
is given to a small sclerite which seems to be an extension of the arciform piece on
the right side of the internal sac.

Asterisks (*) listed in the Localities sections indicate place names for which the
localities are unknown, and an exclamation mark (!) notes that several specimens
were examined but that their sex was not determined.

Quaest. Ent., 1989, 25 (3)

248

Freitag and Bames

CLASSIFICATION

Notes about taxonomic characteristics

Characteristics common to adults of Brazilian species of Cicindela are those
typical of the genus (Horn, 1915; Willis, 1968; Freitag, 1979) as well as the
following which characterize most species: scape of antenna with one apical seta;
tooth of mentum well developed; pronotum with lateral and front portions setose;
lateral portions of metastemum and metacoxa setose; procoxa and mesocoxa setose;
and elytra with isodiametric microsculpture and apical microserrulations.

Taxa of the Brazilian Cicindelidae

The following keys to the taxa of Brazilian Cicindelidae are constructed from
personal examinations of specimens, and diagnostic characteristics given in
descriptions, keys and figures by Horn (1915, 1926a, 1938), Rivalier (1954, 1955),
Freitag (1966, 1972, 1979), Vidal Sarmiento (1966a, b), Pena (1969), Reichardt
(1977), and Sumlin (1979).

Key to Adults of Tribes of Brazilian Cicindelidae

1 Metepistemum narrow, sulcate for entire length;

mesepistemum short; lacinia of maxilla without articulated
tooth Ctenostomatini

1' Metepistemum plate-shaped, not entirely sulcate;

mesepistemum elongate; lacinia with articulated tooth 2

2 (T) Pronotum with anterior lateral angles prominent, projected

further forward than anterior margin of prostemum, head wide,
eyes small; maxillary palpamere 4 in most members of most
taxa shorter than maxillary palpamere 3; body without
pubescence Megacephalini

2' Pronotum with anterior lateral angles not prominent; head with

large prominent eyes; maxillary palpamere 4 longer than
maxillary palpamere 3; body with pubescence in members of
most taxa Cicindelini

Key to Adults of Subtribes and Subgenera of Cicindela of Brazilian Cicindelini

1 Pubescence generally present on head, thorax, and abdomen in

members of most taxa; or posterior 0.33 of elytron with pale
markings subtribe Cicindelina; genus Cicindela 3

T Pubescence generally absent from head, thorax, and abdomen

in members of most taxa; posterior 0.33 of elytron without pale
markings 2

2 (1') Elytral epipleuron and free lateral margin of hind coxa glabrous

Brazilian Species of Cicindela

249

subtribe Prothymina

2' Elytral epipleuron glabrous or pubescent, and free lateral

margin of hind coxa pubescent subtribe Odontocheilina

3 (1) Pale maculations of elytra various, from complete and narrow

to reduced and incomplete; labrum tridentate in most species;

median lobe of male with apical hook in specimens of most species 4

3' Pale maculations of elytra broad to extensive in specimens of

most species, or depressed, or absent, or discontinuous giving
spotted appearance; labrum unidentate or multidentate in
specimens of most species; median lobe of male tapered,
without hook 5

4 (3) Body size small, most adults less than 7.5 mm long; labrum

with five to 10 (most commonly eight) submarginal setae;
pronotum narrow; membrane in place of oviduct sclerite;

ventral sclerite of bursa without posterior projections

subgenus Brasiella Rivalier, p. 254

4' Body size large, most adults 8.0-12.0 mm long (C. anulipes is

7. 0-7. 5 mm); labrum with four to six (seven in a few
specimens) submarginal setae; pronotum broad; oviduct sclerite
present; ventral sclerite of bursa with setose lateral posterior
projections subgenus Gaymara n. subg., p. 278

5 (3') Pale maculations of elytra broad to extensive, or depressed, or

absent; head glabrous in specimens of all but one species (gena
sparsely pilose in C. confluentesignata)’, thoracic pleuron and
sternum glabrous or moderately pilose; labrum unidentate or
multidentate; internal sac of median lobe of male lacking dorsal

spatulate sclerite subgenus Cylinder a Westwood, p. 317

5' Pale maculations of elytra well developed in specimens of most

species, tendency to be discontinuous, appearance spotted; head
with setae (glabrous in C. suturalis)\ thoracic pleuron including
proepistemum and sternum densely pilose; internal sac of

median lobe of male with dorsal spatulate sclerite

subgenus Plectographa Rivalier, p. 289

Key to the Brazilian species of Cicindela

1 Head with setae on either the vertex, frons, clypeus or genae,

fine in some specimens and not obvious or abraded 2

1' Head glabrous (except for one or two supraorbital setae near

the inner margin of each eye) 8

2 (1) Dense appressed setae on vertex, frons, clypeus, genae and

most of the rest of the body; elytra completely or almost

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250

Freitag and Bames

without ground colour (Figs. 91a, b); body size 9.0-13.5 mm;

distributed from Espirito Santo, Brazil, south to Argentina

C. nivea Kirby, p. 300

2' Combination of characters not as above 3

3 (2') Body length 8.5-1 1.0 mm; elytra with ground colour very dark
brown to black, dull, maculations broad, continuous away from
lateral margin, row of large foveae with umbilicate centers near
suture (Fig. 92); distributed in coastal regions in southernmost

portion of Rio Grande do Sul, Argentina, and Chile

C. melaleuca Dejean, p. 302

3' Combination of characters not as above 4

4 (3') Labrum of most specimens with seven to nine submarginal

setae 5

4' Labrum of most specimens with fewer than seven submarginal

setae 6

5 (4) Body length 6. 5-7. 5 mm; elytra with ground colour brown, dull,

marginal band and apical lunule broad, middle band broad,
oblique (Fig. 32); coupling sulcus of mesepistemum of female
deep round pit; distributed in Matto Grosso

C. banghaasi Horn, p. 276

5' Body length 9.0-10.0 mm; elytra with ground colour brown,

glossy, maculations broad, continous along lateral margin
(Figs. 93a, b); coupling sulcus deep groove; distributed from

Rio Grande do Sul south to Patagonia

C. patagonica Brulle, p. 303

6 (4') Dorsum coppery brown, slightly glossy; elytra with broad and

continuous pale maculations (Figs. 130a-c); genae sparsely
setose, frons and clypeus glabrous; antennae with articles 5-1 1
pale; labrum with margin irregular, unidentate in most
specimens; in some, seven dentate, with margin incised by
setiferous punctures (Figs. 116a-c); distributed in Minas Gerais,

Uruguay, Argentina, Paraguay

C. confluentesignata Horn, p. 319

6' Combination of characters not as above 7

7 (6') Ground colour dull (matte) black; one or two small pale spots

on elytra (Fig. 94); vertex, frons, and genae sparsely setose;
coupling sulcus of mesepistemum of female shallow, wide,

almost absent; distributed in Minas Gerais

C. nigrovittata Horn, p. 304

7' Ground colour black (not matte); maculations of elytra reduced

(Figs. 65a-c); vertex, frons clypeus, and genae sparsely setose;
tuft of setae on front inner margin of each eye; coupling sulcus

Brazilian Species of Cicindela

251

elongate groove with deep middle; distributed in Rio Grande do
Sul C. paranigroreticulata new species, p. 287

8 (1') Labrum edentate or with very small single tooth (Figs. 2, 3, 5) 9

8' Labrum either unidentate or multidentate 1 1

9 (8) Labrum with anterior margin broadly protruded in center (Figs.

3a-c); elytra dark brown, purplish reflections in few specimens,
with short wide transverse middle band, small subapical spot of
humeral lunule present in most specimens, and with subapical

spot, (Figs. 25a-c); distributed in Amazon basin

C. pretiosa Dokhtouroff, p. 269

9' Labrum with anterior margin straight (Figs. 2, 5); pattern of

elytral markings as in Figures 24, 27 10

10 (9') Dorsum bright to dark red-coppery, elytral pattern with humeral
spot, middle band and subapical spot (Fig. 27); distributed in

inland southern Brazil C. aureola Klug, p. 271

10' Dorsum black to dark brown; elytral pattern almost effaced

(Figs. 24a, b); distributed in northern Argentina, Bolivia,

Paraguay, Uruguay, southern Brazil

C. obscurella Klug, p. 267

1 1 (8') Labrum at least 5-dentate 12

1 1 ' Labrum at most tridentate 1 5

12 (11) Body length 6.0 mm; body dull black; labrum indistinctly five

dentate with six submarginal setae, with middle four setae close
to margin (Fig. 9); elytron without shoulder, pattern absent

(Fig. 31); distributed in Matto Grosso,

C. brevipalpis Horn, p. 276

12' Combination of characters not as above 13

13 (12') Ground colour dull brown to black with green head and

pronotum; elytra with broad apical and humeral lunules (Figs.

132a, b); tooth of mentum short; labrum 5-dentate (Figs. 118a,

b); distributed in Matto Grosso C. marquardti Horn, p. 321

13' Ground colour black; elytra various, from immaculate to fully

maculate (Figs. 131, 133); labrum seven to 12 dentate (Figs.

1 17, 1 19); tooth of mentum well developed 14

14 (13') Elytra with maculations reduced to three spots, each in

depression, large punctures along suture, large depression in
basal 0.33 (Figs. 133a, b); articles 5-6 of antenna dark; labrum
seven dentate with six marginal setae (Figs. 119a, b); coupling
sulcus of mesepistemum of female in form of groove with

central pit; distributed in Minas Gerais

C. piligera Horn, p. 322

14' Elytra various, from immaculate to fully maculate (Figs.

Quaest. Ent., 1989, 25 (3)

252

Freitag and Bames

15'

16 (15')

16'

17 (16)

131a-e); articles five or six of antenna pale; labrum elongate,
seven to 12 dentate with eight to 10 marginal setae (Figs.

117a-e); coupling sulcus broad groove; distributed from

Amazon River to Matto Grosso C. morio, Klug, p. 320

15 (IF) Elytra with dark lines in form of reticulated pattern against
lighter brown and coppery ground colour, humeral lunule short,
middle band markedly curved (Figs. 64a, b); labrum unidentate
or weakly tridentate with four submarginal setae (Figs. 54a, b);

distributed in inland Rio Grande do Sul

C. nigroreticulata Horn, p. 286

Combination of characters not as above 16

Head and pronotum deeply rugose and bright coppery with
faint green reflections in some specimens; elytra with middle

band narrow and strongly oblique as in Figures 62, 63 17

Head and pronotum not deeply rugose and bright coppery;

elytra with middle band not oblique 18

Body length 8. 0-8. 5 mm; pattern of elytral maculations as in
Figures 62a, b; distributed in central and southeastern Brazil,

northeastern Argentina, Paraguay

C. chlorosticta Kollar, p. 284

17' Body length 10.0-12.0 mm; pattern of elytral maculations as in

Figures 63a, b; distributed in southeastern Brazil, northern
Argentina, Uruguay C. staudingeria Horn, p. 285

18 (16') Labrum elongate, convex, with three small teeth, and four

submarginal setae (Fig. 56); proepistemum with deep and
evenly distributed punctures; elytra with reduced maculations
(Fig. 66); distributed in Goias, Matto Grosso, Minas Gerais,

Rio Grande do Sul C. anulipes Horn, p. 288

18' Combination of characters not as above

19 (18') Labrum distinctly tridentate (Figs. 1, 4, 6, 7)

19' Labrum unidentate or obscurely tridentate

20 (19) Labrum distinctly narrow throughout, central portion of front

margin not protruded, central tooth small (Figs. 7a, b); humeral
spot of elytron small, on latero-ventral side of shoulder, not
evident in dorsal view; distributed in southern Matto Grosso
near the Bolivian border, Central America and northwestern
South America (probably east of the Andes) south to

southwestern Brazil and Argentina

C. misella Chaudoir, p. 274

20' Combination of characters not as above 21

21 (20') Elytra slightly glossy brown, maculations broad, middle band

slightly oblique (Fig. 26); labrum with broadly rounded teeth.

19

20
23

Brazilian Species of Cicindela

253

21'

22 (21’)

22'

23 (19’)

23'

24 (23')

24'

25 (24')

25’

26 (25’)
26'

27 (26)

27'

eight submarginal setae (Fig. 4); distributed inland from Matto
Grosso north to Amazon River ....C. amaenula Chaudoir, p. 270
Elytra dull, brown to black, maculations narrower or more

reduced; teeth of labrum narrower (Figs. 1 , 6)

Elytron with humeral spot, subhumeral spot distinct (Figs.
23a-e); small bell-shaped unpigmented area on posterior margin
of sternum 5 of females distributed from Brazil north to

Mexico, south to Argentina C. argentata Fabricius , p. 265

Elytron with humeral spot absent, subhumeral spot tiny (Fig.
28); bell-shaped unpigmented area on posterior margin of
sternum 5 of females absent; distributed from eastern Brazil

west to Colombia C. dolosula Rivalier, p. 273

Elytra with pale maculations widely expanded, continuous in
some specimens, humeral lunule oblique, punctation very large
and deep (Figs. 129a, b); articles 5-11 of antennae pale; body
pubescence moderately dense; labrum distinctly unidentate
with five or six sub-marginal setae (Figs. 115a, b); distributed

in central Brazil C. kollari Gistl, p. 317

Combination of characters not as above

Elytra black, maculations obsolete and depressed, with broad
punctures near median suture and shoulders (Fig. 134); labrum
unidentate, dark at base, six submarginal setae (Fig. 120);

distributed in Santa Catarina, northern Argentina

C. obsoletesignata Horn, p. 323

Combination of characters not as above

Body length 6.0 mm; elytra dull dark brown, maculations
largely effaced (Fig. 30); labrum unidentate with five to seven

submarginal setae (Fig. 8); distributed in Goias

C. hamulipenis Horn, p. 275

Combination of characters not as above

Elytra with maculations complete and continuous or broad

(Figs. 90, 95)

Elytra with maculations reduced (Figs. 33, 121)

Elytra with ground colour dull dark brown, maculations broad,
lacking marginal band (Figs. 95a-d); labrum unidentate, tooth
prominent, or obscurely tridentate, seven to 13 submarginal
setae (Figs. 83a-d); coupling sulcus of mesepistemum of female
broad groove; distributed in coastal and inland Rio de Janeiro,
Minas Gerais, Matto Grosso, Rio Grande do Sul, northern

Argentina C. apiata Dejean, p. 305

Elytra with ground colour glossy, with coppery, green and blue
(in some specimens) reflections, maculations complete and

22

24

25

26

27

28

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254

Freitag and Bames

continuous, middle band sinuate (Figs. 90a-e); labrum
unidentate, with 8-10 submarginal setae (Figs. 78a-e); coupling
sulcus deep sinuate groove; distributed from southern Brazil

north to Trinidad and southern Caribbean Islands

C. suturalis Fabricius, p. 298

28 (26') Elytra dull, black, apical lunule with recurved hook at distal
end (Figs. 33a, b); labrum short almost tridentate, with seven to
12 submarginal setae (Figs. 11a, b); coupling sulcus of
mesepistemum of female deep groove with central pit;
distributed in Espirito Santo, Minas Gerais, Matto Grosso, Sao

Paulo C. minarum Putzeys, p. 277

28' Elytra slightly glossy, black, middle band, especially apical

end, depressed (Figs. 135a, b); labrum unidentate with seven of
eight setae almost marginal (Figs. 121a, b); coupling sulcus
long sinuate groove; distributed in Rio Grande do Sul, Santa
Catarina C.friedenreichi Dokhtouroff, p. 324

Subgenus Brasiella Rivalier

Genus Brasiella Rivalier, 1954: 261 (TYPE SPECIES, Cicindela argentata, by iginal designation).

Rivalier, 1955: 79. Reichardt, 1977: 374.

Brasiliella Mandl, 1963: 581; 1973: 270 (incorrect subsequent spelling).

Recognition. — Adults of this subgenus are distinguished by the five characters
given in couplet 4 of the key. In addition the head is glabrous (except for C.
banghaasiy, labrum tridentate, unidentate, edentate, or indistinctly five dentate; pale
maculations of elytra are complete and narrow in most species or reduced or
completely absent; middle of abdominal sterna pubescent; apex of median lobe of
the male is hooked in most species, and central plate and flagellum are absent in the
internal sac; the spermatheca and duct of the female together are approximately
1.0- 1.5 mm long.

Species groups. — This subgenus has seven species groups that include 29
species. Six groups are found in South America, of which four are in Brazil,
argentata group, aureola group, misella group, and minarum group.

Geographical distribution. — The geographical range of this subgenus extends
from northern Argentina northward to southwestern United States and the West
Indies. In Brazil most taxa in this subgenus are concentrated south of the Amazon
River in the Brazilian Highlands and southern parts of the Amazon basin.

Phylogenetic relationships. — Subgenus Brasiella is sister to the lineage that
gave rise to subgenera Gaymara, Plectographa , and Cylindera.

Figs. 1-11. Labium, dorsal aspect. 1, Cicindela argentata Fabricius: (a) female, Estac, S.P.; (b) female,
Ypiranga, S.P.; (c) male, Arinos, M. Gerais; (d) male, 40 km w. Manaus, Am.; 2, C. obscurella Klug,
Montevideo, Uruguay: (a) female; (b) male; 3, C. pretiosa Dokhtouroff, Manaus, Am.: (a) female; (b), (c)
male; 4, C. amaenula Chaudoir, female, near Amazon River; 5, C. aureola Klug, female: (a) Sao Paulo, S.P.;
(b) Vacaria, M. Grosso; 6, C. dolosula Rivalier, male, Chapada, Go.; 7, C. misella Chaudoir, Bugaba,
Colombia: (a) female; (b) male; 8, C. hamulipenis Horn, male, Dianopolis, Go.; 9, C. brevipalpis Horn,
male, Vacaria, M. Grosso; 10, C. banghaasi Horn, female, Cuyaba, M. Grosso; 1 1, C. minarum Putzeys: (a)
female, Vacaria, M. Grosso; (b) male, Espirito Santo. Figs. 12-22. Pronotum, dorsal aspect. 12, C. argentata
Fabricius: (a) female, Ypiranga, S.P.; (b) male, Arinos, M. Gerais; (c) male, 40 km w. Manaus, Am.; 13, C.
obscurella Klug, Montevideo, Uruguay: (a) female; (b) male; 14, C. pretiosa Dokhtouroff, Manaus, Am.: (a)
female; (b), (c) male; 15, C. amaenula Chaudoir, female, near Amazon River; 16, C. aureola Klug, female;
(a) Sao Paulo, S.P.; (b) Vacaria, M. Grosso; 17, C. dolosula Rivalier, male Chapada, Go.; 18, C. misella
Chaudoir, Bugaba, Columbia: (a) female; (b) male; 19, C. hamulipenis Horn, male, Dianopolis, Go.; 20, C.
brevipalpis Horn, male, Vacaria, M. Grosso; 21, C. banghaasi Horn, female, Cuyaba, M. Grosso; 22, C.
minarum Putzeys; (a) female, Vacaria, M. Grosso; (b) male, Espirito Santo.

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Figs. 23-33. Elytron, dorsal aspect. 23, Cicindela argentata Fabricius; (a) female, Estac, S.P., apical lunule
(. al ), humeral spot (hs), middle band (mb), marginal band or lunule (ml), subhumeral spot (sh)\ (b), (c),
female, Ypiranga, S.P.; (d) male, Arinos, M. Gerais; (e) male, 40 km w. Manaus, Am.; 24, C. obscurella
Klug, Montevideo, Uruguay: (a) female; (b) male; 25, C. pretiosa Dokhtouroff, Manaus, Am.: (a) female;
(b), (c) male; 26, C. amaenula Chaudoir, female, near Amazon River; 27, C. aureola Klug, female: (a) Sao
Paulo, S.P.; (b) Vacaria, M. Grosso; 28, C. dolosula Rivalier, male, Chapada, Go.; 29, C. misella Chaudoir,
Bugaba, Colombia: (a) female; (b) male; 30, C. hamulipenis Horn, male, Dianopolis, Go.; 31, C. brevipalpis
Horn, male, Vacaria, M. Grosso; 32, C. banghaasi Horn, female, Cuyaba, M. Grosso; 33, C. minarum
Putzeys: (a) female, Vacaria, M. Grosso; (b) male, Espirito Santo.

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257

Figs. 34—37. Female genitalia: (a) sternum 8 ( s8 ), second gonocoxa (sg), second gonapophyses (sgp), ventral
aspect; (b) syntergum 9 and 10 ( t9&10 ), dorsal aspect; (c) bursa copulatrix (be), median ridge (mr), oviduct
sclerite (os), oviduct ( ov ), spermatheca ( sp ) and duct ( sd ), ventral sclerite (vs), ventral aspect; (d) bursa
copulatrix left lateral aspect; (e) bursa copulatrix dorsal aspect. Cicindela argentata Fabricius: 34, Estac,
S.P.; 35, Ypiranga, S.P.; 36, 37, C. obscurella Klug, Montevideo, Uruguay.

Quaest. Ent., 1989, 25 (3)

Figs. 38 — 4 1 . Female genitalia: (a) sternum 8, second gonocoxa, second gonapophyses, ventral aspect; (b)
syntergum 9 and 10, dorsal aspect; (c) bursa copulatrix, median ridge, oviduct sclerite, oviduct, spermatheca
and duct, ventral sclerite, ventral aspect; (d) bursa copulatrix left lateral aspect. 38, Cicindela pretiosa
Dokhtouroff, Manaus, Am; 39, C. amaenula Chaudoir, near Amazon River; C. aureola Klug, 40, Sao Paulo,
S.P.; 41, Vacaria, M. Grosso.

Brazilian Species of Cicindela

259

Figs. 42-44. Female genitalia: Female genitalia: (a) sternum 8, second gonocoxa, second gonapophyses,
ventral aspect; (b) syntergum 9 and 10, dorsal aspect; (c) bursa copulatrix, median ridge, oviduct sclerite,
oviduct, spermatheca and duct, ventral sclerite, ventral aspect. 42, Cicindela misella Chaudoir, Bugaba,
Columbia; 43, C. banghaasi Horn, Cuyaba, M. Grosso; 44, C. minarum Putzeys, Vacaria, M. Grosso. Figs.
45 — 46. Male genitalia: median lobe (a) right lateral aspect; (b) dorsal aspect; (c) left lateral aspect; (d)
internal sac, arciform piece (ap), setal brush ( sb ), shield ( sh ), stylet (st), tooth (to). 45, C. hamulipenis Horn,
Dianopolis, Go.; 46, C. brevipalpis Horn, Vacaria, M. Grosso.

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Freitag and Bames

Fig. 47. Map showing the geographical distribution of the species Cicindela argentata Fabricius. Open
circles represent state records.

Brazilian Species of Cicindela

261

Fig. 48. Map showing the geographical distribution of the species Cicindela obscurella Klug (•), and C.
pretiosa Dokhtouroff (■). Open circles represent state records of C. obscurella Klug.

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Freitag and Bames

Fig. 49. Map showing the geographical distribution of the species Cicindela amaenula Chaudoir (•), and C.
aureola Klug (■). Open circle and square represent state recor for C. amaenula Chaudoir and C. aureola
Klug respectively.

Brazilian Species of Cicindela

263

Fig. 50. Map showing the geographical distribuiton of the species Cicindela dolosula Rivalier (•), and C.
misella Chaudoir (■). Open circle and square represent state records for C. dolosula Rivalier, and C. misella
Chaudoir respectively.

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Fig. 51. Map showing the geographical distribution of the species Cicindela hamulipenis Horn (•), C.
brevipalpis Horn (■), C. banghaasi Horn (A), and C. minarum Putzeys (▼).

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265

The argentata group

Adults of this group are characterized by elytral foveae not obvious, and
unpigmented bell-shaped spot on abdominal sternum 5 of the female. The group has
four species, three of which are found in Brazil, C. argentata, C. obscurella, and C.
pretiosa. The geographical range of the argentata group extends from northern
Argentina northward to northern South America and the Lesser Antilles. The
argentata group is a close relative of the viridicollis and hemichrysea groups.

Cicindela (Brasiella) argentata Fabricius
Figs. 1, 12, 23,34,47

Cicindela argentata argentata Fabricius, 1801: 242 (TYPE LOCALITY, in America meridionali). - Herbst,
1806: 208. - Dejean, 1825: 147; 1831: 215. Kirsch, 1873: 125. - Bates, 1881: 14. - Horn, 1896a: 354;
1903: 334; 1904: 86; 1906: 87-91; 1915: 406; 1923: 111; 1926a: 308: 1938: 52. Varas Arangua, 1925:
37. - Barattini, 1929: 1218. - Blackwelder, 1944: 17. - Cazier, 1954: 286. - Rivalier, 1954: 261-263;
1955: 79-80; 1970: 857. Brouerius van Nidek, 1956: 320. - Mandl 1956: 389; 1963: 581; 1964: 16;
1967: 437, 439; 1973: 290. Vidal Sarmiento, 1966a: 256-257; 1966b: 32-33. - Balazuc and Chalumeau,
1978: 22. - Sumlin, 1979: 102.

Cicindela guerin Gory, 1833: 178 (TYPE LOCALITY, Cayenne). - Horn, 1896a: 354; 1915: 406; 1926a:
308. - Blackwelder, 1944: 17.

Cicindela lucorum Gistl ,1837: 71 (TYPE LOCALITY, in Cayenna). - Horn, 1915: 406; 1926a: 308. -
Blackwelder, 1944: 17.

Cicindela egaensis Thomson, 1857: 130 (TYPE LOCALITY, Ega, Amazone super.).) - Horn, 1915: 406;
1926a: 308. - Blackwelder, 1944: 17.

Brasiliella pseudoargentata Mandl, 1963: 582 (TYPE LOCALITY, Jacare P.N. Xingu, M. Grosso, Bras.).
NEW COMBINATION AND SYNONYMY.

Cicindela argentata pallipes Fleutiaux and Salle, 1889: 359 (TYPE, sex undetermined, in the MNHP general
collection bearing the following label: “Guadeloupe Delauney/C. argentata F., v. pallipes (ch2)/Museum
Paris Box collection Fleutiaux/Type” (black letters on red label); (TYPE LOCALITY, Guadeloupe).
-Horn, 1903: 334; 1915: 406; 1926a: 308. - Leng and Mutchler, 1916: 696. - Blackwelder, 1944: 17. -
Rivalier, 1955: 80.

Cicindela argentata umbrogemmata Horn , 1906: 87 (TYPE LOCALITY, Posorja, Ecuador (Campos));

1915: 407; 1926a: 308; 1938: 52. -Blackwelder, 1944: 17.

Cicindela argentata macella Rivalier , 1955: 80 (TYPE, a male in the MNHP general collection bearing the
following label: “Museum Paris de la Mana Leschor/ penis 873 Rivalier /argentata s.s. macella mihi. E.
Rivalier det/TYPE” (black letters on red label); TYPE LOCALITY, Bresil).

Cicindela argentata semicircumscripta Mandl , 1958: 23 (TYPE LOCALITY, Santiago del Estero, El
Pinto). - Sumlin, 1979: 103.

Cicindela argentata ecuadorensis Mandl , 1973: 290 (TYPE LOCALITY, Ecuador, San Anton. Curaray).

Recognition. — Specimens of C. argentata are distinguished from those of the
similar species C. obscurella by a combination of the following characteristics: in
most specimens maculations of elytra well developed with complete marginal band
and humeral spot, subapical spot and apical lunule (Figs. 23a-e); labrum of average
length and obsoletely to strongly tridentate with middle tooth well developed in
most specimens (Figs, la-d); sides of inside of bursa copulatrix in females lightly
sclerotized and brushes of setae absent (Figs. 34c, 35c). In addition, the shapes of
the sclerites of the internal sac of the median lobe of the two above species differ
(Rivalier, 1955: 81,84; Vidal Sarmiento, 1966b: 33). Also see Recognition section

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for C. misella and C. dolosula.

Synonyms and Types. — We have not seen types of the C. argentata complex
except for that of C. argentata macella Rivalier and C. argentata pallipes Fleutiaux
and Salle. Names are based upon comparison of original descriptions with
specimens on loan. Rivalier (1955: 80) is correct in stating that C. taitiensis
Boheman is a cicindelid from Tahiti which is not an element of the American fauna.
The name C. pseudoargentata Mandl has been assigned to small specimens of C.
argentata.

Description. —

Body length. 6.0-8.0 mm M and F.

Body color. Head, pronotum and elytra dull to slightly glossy, black to coppery brown, elytra of some
specimens with green reflections. Venter, pleuron and sides of head with green, blue, and coppery
reflections.

Body setae. Proepistemum and mesepistemum sparsely setose, metepistemum more densely setose;
lateral margins of abdominal sterna 1-6 setose.

Other external features. Labrum of average length, tridentate, with eight submarginal setae, ranging in
number from five to 12 (Figs. la-d). Pronotum narrow, sutures shallow (Figs. 12a-c). Coupling sulcus of
female shallow to moderately deep groove, with deeper pit in middle. Apex of front trochanters with one
seta. Apices of elytra slightly to not recurved. Pattern of elytral maculations with humeral lunule
discontinuous, humeral spot on shoulder in most specimens, humeral subapical spot present; marginal band
in most specimens; middle band complete or discontinuous; apical lunule complete or discontinuous (Figs.
23a-e). Punctures of elytra large, shallow, with green (mainly) and coppery reflections.

Female genitalia. Sternum 8 with deep and broad V-shaped posterior emargination; apices moderately
rounded, each with three short stout setae (Figs. 34a, 35a). Second gonapophyses broad, medial portion
almost as long as lateral portion (Figs. 34a, 35a). Syntergum 9 and 10 as in Figures 34b, 35b. Ventral sclerite
broad, lightly sclerotized, median ridge absent; inside lateral walls of bursa slightly sclerotized (Figs. 34c,
35c). Thick membrane in place of oviduct sclerite (Figs. 34c, 35c). Length of spermatheca and duct ca 1.0
mm. See also Vidal Sarmiento (1966a; 256).

Male genitalia. Male genitalia have been described by Rivalier (1955: 81) and Vidal Sarmiento (1966b:
33).

Geographical Variation and Subspecies. — Variable elytral maculations include
the humeral spot almost absent or very well marked, and marginal band absent or
reduced (few specimens) or well marked (most specimens). The middle band is
discontinuous in some specimens. A complete apical lunule is the general condition,
though a few specimens with a discontinous one are present throughout the species
range. Geographical patterns in these characteristics are not clearly discernible,
though maculations of the elytra are broader and more frequently complete in
specimens from northern Brazil. Specimens in southern Brazil tend to have
discontinuous or absent portions of the elytral maculations. Specimens of C. a.
macella Rivalier have elytral maculations much reduced. Whether or not they are
the predominant form in well defined geographical populations remains to be seen.
We follow recent authors in recognizing subspecies of non-Brazilian forms.

Relationships. — Cicindela argentata is sister to the lineage that gave rise to C.
venustula Gory (northern S. America) and sister species C. obscurella and C.
pretiosa.

Habitat and Period of Activity. — Label data indicate a period of activity from
December to April. Adults live in moist open areas in grassy vegetation. They

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267

occasionally occur along river beaches but usually near clumps of grass (Pearson,
1984).

Geographical Distribution, Localities, Examined Specimens. — Inland and near
coasts, from northern Argentina, Uruguay to northern Brazil and French Guiana
(Fig. 47), and Guadeloupe.

Argentina. Cordoba : no locality, 1M, MZSP. Formosa : Guaycolec, 2F, MZSP. La Rioja : Patquia, 1M,
IF, MZSP. Tuouman : no locality, 1M, IF, MZSP.

Bolivia. Prov. del Sara*, 6M, 9F, ICCM; Santa Cruz de la Sierra Bol., 1M, ICCM.

Brazil. Amazonas : Arima (on Rio Purus), 1M, ICCM; Beruri (on Rio Purus), 4M,5F,MZSP; Caninde (50
km e), 4M, 3F,MZSP; Hyutanaham (on Rio Purus), 31M, 18F, ICCM; Itacoatiara (30 km w. on Rio Urubu),
2F, ICCM; Manaus, 3M, 4F, INPA; Manaus (Reserva Ducke, km 26 on Itacoatiara Hwy), 1M, BRI; Manaus
(30 km ne), 3M, INPA; Manaus (40 km w. at Lago Janauaca), 1M, BRI: Manaus (60 km n), 1M INPA; Rio
Cavaburi, 3F, MZSP; Rio Preto, 1M, MZSP; Santo Antonio do Iga, IF, MNRJ; Sao Gabriel, 1M 2F, IOC;,
Tapuruquara (on Rio Negro), IF, MZSP; Tefe, IF IOC. Ceara : Aracati, IF, MZSP; IOC, 1M, IF, MZSP;
Jaquaribe, 1M, MZSP; Russas, 1M, 2F, MZSP . Espirito Santo : Guandu*, I'M, IF, IOC; no locality IF,
MNRJ. Goias: Cabeceiras (Lagos Formosa), 1M, 2F, MZSP; Chapada, 6M, 6F, ICCM; Jatai, 17M, 6F,
MZSP; Pirineus*, IF, MZSP; Vianopolis, IF, MZSP. Matto Grosso : Barra do Tapirape, 3M, IF, MZSP;
Bodoquena, !, IOC; Camisao*, IF, MZSP; Corumba, 1M, MZSP; Jacare Pq. Nac. Xingu, 1M, MZSP;
Murtinho*, 1M, MZSP; Vacaria, 6M, IF, MZSP; Salobra (Rio?), !, IOC; Tres Lagoas, 1M, MZSP; Urucum,
IF, MZSP, 1M, MNRJ; Xingu, 1M, MZSP. Minas Gerais: Buritis, 1M, IF, MZSP; Ipatinga, 1M, MZSP;
Mar de Espanha*, 1M, MNRJ; no locality, IF, MZSP; Serra Caraca, IF, MZSP. Para: Cachimbo, !, IOC,
12!, MZSP; Oriximina, 2M, 2F, MZSP; Santarem, 7M, 5F, ICCM. Paraiba: Corema, 1M, MZSP. Parana:
Ponta Grossa, 1M, IF, MZSP. Santa Catarina: Joinville, 1M, MZSP; Nova Teutonia, 2M, IF, MZSP. Sao
Paulo: Alto da Serra*, 1M, MZSP, Avanhand (Garbe Lake)*, 1M, MZSP; Barueri, 4M, 5F, MZSP;
Boraceia*, 1M, 3F, MZSP; Campos do Jordao, IF, MZSP; Cantareira, IF, MZSP; Caraquatatuba, IF,
MZSP; Embu, IF, MZSP; Estac, IF, MZSP; Iporanga, 3F, MZSP; Itu, 2M, MZSP; Mairipora, 1M, MZSP;
1M, MZSP; Onda Verde*, !F, MZSP; Pindamonhangaba, 1M, IF, MZSP. Pirassununga, 1M, MZSP;
Rincao, 1M, MZSP; Sao Paulo, 2M, 2F, MZSP; Sapuchui, 1M, MZSP; Ypiranga, !, MZSP.

French Guiana. Oiapoque River, 3M, 5F, ICCM.

Paraguay. Asuncion, 2F, ICCM.

Peru. Huancayo, !, IOC; Rio Ampiacu*, 1M, IF, MZSP.

Uruguay. Paysandu (on Rio Uruguay), 1M, MNRJ.

(Note: Ecuador; C. a. umbrogemmata Horn, 1906: 87.

Venezuela; C. argentata and C. a. pallipes Horn, 1903: 334.

Cicindela (Brasiella) obscurella Klug
Figs. 2, 13, 24, 36, 37, 48

Cicindela obscurella Klug, 1829: 3 (TYPE LOCALITY, Sud-Brasilien). - Dejean, 1831: 268. - Horn, 1891:
324; 1906: 89; 1915: 407; 1926a: 308; 1938: 52. - Barattini, 1929: 1219. - Fernandez, 1936: 105. -
Blackwelder, 1944: 17. - Rivalier, 1954: 263; 1955: 82. Vidal Sarmiento, 1966a: 256-257; 1966b: 32.
Sumlin, 1979: 103.

Cicindela tripunctata Dejean , 1831: 267 (TYPE LOCALITY, les parties meridionales du Bresil). - Horn,
1915: 407; 1926a: 308. Blackwelder, 1944: 17. - Vidal Sarmiento, 1966b, 32.

Cicindela celeripedestris Horn, 1896b: 357 (TYPE LOCALITY, Minas Geraes); 1938: 52. Blackwelder,
1944: 17. - Rivalier, 1955: 84. Vidal Sarmiento, 1966b: 32.

Cicindela obscurella constricta Rivalier, 1955: 83 (TYPE, a male in the MNHP general collection bearing
the following labels: “Bresil (Minas) Sertao de Diamantina Faz Das Melancias E. Gounelle 10-11
1902/Museum Paris Bresil coll. E. Gounelle 1913/ B. obscurella ssp. constricta mihi E. Rivalier
det/TYPE” (black letters on red label)/ “penis 974 Rivalier”; TYPE LOCALITY, Minas Geraes). NEW
SYNONYMY.

Brasiliella chrysocollis Mandl, 1963: 585 (TYPE LOCALITY, Jacare P.N. Xingu, M. Grosso, Bras.). NEW
COMBINATION AND SYNONYMY.

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Brasiliella pallidipes Mandl, 1963: 589 (TYPE LOCALITY, Sta. Catarina, Brasil). NEW COMBINATION

AND SYNONYMY.

Recognition. — The elongate and edentate labrum (Figs. 2a, b) combined with
the almost effaced elytral maculations. (Figs. 24a, b) separates C. obscurella from
other species of the argentata group. Adults of C. obscurella are generally larger
than adults of its sister species C. argentata. Two dark finger-like brushes of setae in
the bursa copulatrix of females also distinguish C. obscurella from other related
species (Figs. 36d, 37d, e). In addition the shapes of sclerites in the internal sac of
the median lobe of males characterize this species (Rivalier, 1955: 84).

Synonyms and Types. — Except for the holotype and allotype of C. o. constricta
Rivalier, we have not seen the types of this complex. The names are based on
comparison of original descriptions with specimens on loan.

In treating C. celeripedestris Horn as a junior synonym we follow Horn (1938:
53) and Vidal Sarmiento (1966b: 32). We have examined specimens from Uruguay
which conform to the description of C. o. constricta Rivalier. They do not appear to
form a single geographical population. Brasiliella chrysocollis Mandl appears to be
a coloured form of C. o. constricta Rivalier. We consider Brasiliella pallidipes
Mandl to be a small form of C. obscurella in view of the fact that the labrum is
edentate and that it is found within the range of C. obscurella.

Description. —

Body length. 8.0 mm M, 8.5 mm F.

Body colour. Head and pronotum slightly glossy, black with coppery reflections; elytra dull, black with
coppery reflections, some specimens with green, blue or purple reflections. Venter glossy, black with green,
blue, purple and coppery reflections; pleuron mainly coppery, green and blue.

Body setae. Pronotum and proepistemum sparsely setose, mesepistemum with a few setae at ventral end,
metepistemum more densely setose; abdominal sterna one to six setose mainly on lateral margins.

Other external features. Labrum elongate edentate, median portion almost tooth-like in some females,
with eight submarginal setae, ranging in number from six to 10 (Figs. 2a, b). Pronotum narrow, sutures
shallow (Figs. 13a, b). Coupling sulcus of mesepistemum of female a long and sinuate groove. Apex of front
trochanters with one sensory seta, middle of trochanters glabrous. Elytra with recurved apices; maculations
discontinuous or almost effaced; punctures large, shallow, with green (mainly) and coppery reflections (Figs.
24a, b); a few erect setae present near shoulder; microsculpture isodiametric, bead-like; apical
microserrulations very small, almost obsolete.

Female genitalia. Sternum 8 with wide V-shaped shallow emargination in apical end; apices broadly
rounded, each with a group of four short stout setae (Fig. 36a). Second gonocoxa with setae along medial
margin (Fig. 36a). Second gonapophyses with medial portion 2/3 length of lateral portion (Fig. 36a).
Syntergum 9 and 10 rectangular (Fig. 36b). Ventral sclerite strongly sclerotized, median ridge well
developed with two finger-like brushes on inside of bursa (Figs. 36c, d, 37c-e). Membrane in place of
oviduct sclerite (Figs. 36c, 37c). Spermatheca and duct ca 1.5 mm.

Male genitalia. Male genitalia have been described by Rivalier (1955: 84).

Geographical Variation. — Within populations, the elytral maculations,
especially the middle band, vary from diffuse to almost effaced (Figs. 24a, b). A few
adults with almost complete middle bands are present in Sao Paulo and can be
confused with C. argentata adults.

Relationships. — Cicindela obscurella and C.pretiosa are sister species.

Habitat and Period of Activity. — Adults have been collected in December in
Brazil, and in February in Argentina, Paraguay and Uruguay. Habitat unknown.

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269

Geographical Distribution, Localities, Examined Specimens. — Inland northern
Argentina, Bolivia, Paraguay and Uruguay to south Brazil (Fig. 48).

Argentina. Salta: San Lorenzo*, 1M, IF, MNRJ. Tucuman: no locality, 1M, 2F, IOC.

Brazil, no locality, IF, BMNH, IF, USNM. Minas Gerais : Pocos de Caldas, 6F, RRMC. Santa
Catarina: Nova Teutonia, 6!, RRMC. Sao Paulo: Campos do Jordao, 3M, 10F, MZSP.

Paraguay. Loma*, IF, IOC; no locality, 2M, IF, IOC, Puerto Bertoni*, 1M, IF, IOC, Villarrica, 1M,
IOC.

Uruguay. Maldonado, 1M, MNRJ; Montevideo, 4F, CASC, 1M, MNRJ, 4M, 7F, 1!, USNM; Rocha,
1M, MNRJ; Tacuarembo, IF, MNRJ.

Cicindela (Brasiella) pretiosa Dokhtouroff
Figs. 3, 14, 25,38, 48

Cicindela pretiosa Dokhtouroff, 1882: 276 (TYPE LOCALITY, l’Amazone). Horn, 1915: 407; 1926a, 309;

1938: 52. Blackwelder, 1944: 19. Rivalier, 1954: 263; 1955: 97.

Recognition. — The protruded central portion of the front margin of the labrum
(Figs. 3a-c) together with pattern of elytral maculations distinguish adults of C.
pretiosa from those of all other South American tiger beetles.

Synonyms and Types. — We have not seen the type specimen of C. pretiosa. The
name is based on comparison of specimens on loan with the original description and
the drawings of the elytra by Horn (1938: 52) and Rivalier (1955: 97).

Description. —

Body length. 7.0 mm M, 9.0 mm F.

Body colour. Head and pronotum slightly glossy, dark brown to black with coppery reflections; elytra
dull, dark brown with coppery reflections. Venter black with blue, green, and coppery reflections.

Body setae. Pronotum with front and lateral margin moderately setose, and a few setae medially;
proepistemum and mesepistemum with a few setae near ventral margin; metepistemum moderately setose
throughout; metastemum and abdominal sterna setose laterally.

Other external features. Labrum edentate (or obsoletely tridentate), with front margin broadly
protruding in center (Figs. 3a-c), and eight to 10 submarginal setae. Pronotum broader anteriorly (Figs.
14a-c). Coupling sulcus of mesepistemum of female a deep sinuate groove. Elytra apices slightly recurved
with small apical spine; humeral lunule represented by a small subapical spot, with a small humeral spot
present in some specimens, middle band short, sinuate and broad, and apical lunule reduced (Figs. 25a-c);
punctation broad, shallow with green and coppery reflections; microsculpture isodiametric, bead-like.

Female genitalia. Sternum 8 with broad V-shaped posterior emargination, apices each with a group of
three stout setae (Fig. 38a). Second gonocoxa with medial setae. Second gonaphophyses with medial portion
0.75 length of lateral portion (Fig. 38a). Syntergum 9 and 10 as in Figure 38b. Ventral sclerite broad, slightly
sclerotized, with two apical setiferous brushes; median ridge absent. Membrane in place of oviduct sclerite
(Fig. 38c). Spermatheca and duct ca 1.5 mm in length.

Male genitalia. Unknown.

Geographical Variation. — The pattern of the elytral maculations varies,
particularly in the shape of the middle band and the presence or absence of the
humeral and subhumeral spots (Figs. 25a-c).

Relationships. — Cicindela pretiosa and C. obscurella are sister species.

Habitat and Period of Activity. — Adults have been collected in January and
February. Habitat unknown.

Geographical Distribution, Localities, Examined Specimens. —

Amazon Basin (Fig. 48).

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270

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Brazil. Amazonas : Itacoatiara (km 244), 1M, INPA; Manaus (1 km w. Taruma Falls), 1M, IF, ICCM;
Manaus (30 km. n), 1M, INPA; Manaus (60 km n.), 2M, IF, INPA; Manaus (Reserva Ducke), 2M, IF,
INPA.

The aureola group

Adults of this group are characterized by bright coppery with some green on the
head, pronotum and elytra, and unpigmented bell-shaped spot in abdominal sternum
5 and setae present in the bursa copulatrix of the female. The group has four species,
two of which are found in Brazil, C. amaenula and C. aureola . The geographical
range of the aureola group extends from northern Argentina northward to
Venezuela. This group is a distant relative of the argentata, hemichrysea and
viridicollis groups.

Cicindela ( Brasiella ) amaenula Chaudoir
Figs. 4, 15, 26, 39,49

Cicindela amaenula Chaudoir , 1854: 120 (TYPE LOCALITY, les rives du fleuve des Amazones). - Horn

1906: 88; 1915: 407; 1923: 112; 1926a: 308; 1938: 52. Blackwelder, 1944: 17. - Rivalier, 1954: 263;

1955: 89. Mandl, 1963: 588, 591.

Recognition . — Specimens of C. amaenula are characterized by teir very broad
elytral maculations (Fig. 26). As Rivalier (1955: 90) observed so e C. amaenula
specimens have narrow maculations and may be confused witii C. argentata
specimens, but the former are distinguished by their more obliquely directed middle
band (Fig. 2 cf Fig. 23a) combined with poorly developed marginal teeth of the
labrum (Fig. 4). Female genitalia (Fig. 39) and male genitalia (Rivalier, 1954: 262;
1955: 90) are also specifically distinct.

Synonyms and Types. — The name C. amaenula is based on comparisons of the
original description with specimens on loan.

Description. —

Body length. 8.0 mm F.

Body colour. Head and pronotum slightly glossy brown to black with green and coppery reflections;
elytra slightly glossy brown with coppery reflections. Venter glossy, black with coppery green and blue
reflections pleura with mainly coppery and green reflections.

Body setae. Pronotum moderately setose, proepistemum sparsely setose; mesepistemum with a few
setae at ventral end; metepistemum more densely setose; metastemum setose laterally; abdominal sterna 1-6
setose, more densely so on lateral margin.

Other external features. Labrum obsoletely tridentate with eight submarginal setae (Fig. 4). Posterior
end of pronotum narrow (Fig. 15). Coupling sulcus of mesepistemum of female a long sinuate moderately
deep groove. Apices of front trochanters with one seta. Elytra of female with slightly recurved apices, apical
spine not well developed; humeral lunule reduced to shoulder and apical spots, middle band, marginal band
and apical lunule well developed and broad (Figs. 26); punctation broad, shallow and with green (mainly)
and coppery reflections.

Female genitalia. Sternum 8 with shallow broad V-shaped posterior emargination; apices broadly
rounded each with a group of four stout setae (Fig. 39a). Second gonocoxa moderately setose near medial
margin (Fig. 39a). Second gonapophyses with medial porti 0.75 length of lateral portion (Fig. 39a).
Syntergum 9 and 10 as in Figure 39b. Ventral sclerite bro<r pically, very narrow basally; median ridge
absent. Membrane in place of oviduct sclerite (Fig. 39c). Sp itheca and duct ca 1.0 mm in length.

Brazilian Species of Cicindela

271

Male genitalia. Male genitalia have been described by Rivalier (1954: 262; 1955: 90).

Geographical Variation. — Even though considerable variation is evident in the
pattern of the elytral maculations (Rivalier, 1955: 90) they are broad in most
specimens. A geographical pattern could not be discerned as too few specimens
were available for examination. Specimens examined from Matto Grosso were
typically brown with coppery reflections, although a single green specimen was
seen.

Relationships. — C. amaenula is sister to the lineage that gave rise to sister
species C. aureola and C. horioni Mandl (Bolivia).

Habitat and Period of Activity. — Adults have been collected in November; they
are probably riparian.

Geographical Distribution, Localities, Examined Specimens. — Inland, ranging
from Matto Grosso north to the Amazon River (Fig. 49).

Bolivia. Prov. de Sara*, IF, ICCM.

Brazil. Amazonas : Amazon River, IF, BMNH, 2M, MZSP. Goias: Chapada, 4M, 4F, ICCM. Matto
Grosso: Corumba, 6M, IF, MZSP; Cuyaba, 1M, RRMC, Sao Luiz de Caceres, 3M, 2F, IOC, 1M, MNRJ.

Cicindela ( Brasiella ) aureola Klug
Figs. 5, 16, 27, 40,41,49

Cicindela aureola aureola Klug, 1834: 35 (TYPE LOCALITY, sudlichen Brasilien). Horn, 1906,: 88; 1915:
407; 1926a,: 308; 1938: 52. Varas Arangua, 1925: 37. Blackwelder, 1944: 17. Rivalier, 1954: 263;
1955: 89. Vidal Sarmiento, 1966b: 34. Sumlin, 1979: 103.

Cicindela alboguttata Audouin and Brulle, 1839: 137 (not Klug) (TYPE LOCALITY, Bresil. Province de
Campos-Geraes). Horn, 1896a: 353; 1915: 407; 1926a: 308. Blackwelder, 1944: 17. Vidal Sarmiento,
1966b: 34.

Cicindela argyrosticta Gemminger and Harold, 1868: 9 (replacement name for C. alboguttata Audouin and
Brulle). Horn, 1892b: 213; 1915: 407; 1926a: 308. Blackwelder, 1944: 17. Vidal Sarmiento, 1966b: 34.
Cicindela cyanitarsis Kollar, 1836: 332 (TYPE LOCALITY, in Brasiliae provincia Ypanema). Hom, 1891:
324; 1892a: 95; 1915: 407; 1926a: 308; 1938: 52. Blackwelder, 1944: 17. Rivalier, 1955: 91. Mandl,
1960: 279; 1963: 587. Vidal Sarmiento, 1966b: 34.

Cicindela aureola jatahyana Rivalier, 1955: 91 (TYPE, a male in the MNHP general collection bearing the
following label: “Jatahy Etat de Goyas ch. Pujol 1895-96/ Museum Paris 1952 coll. R. Oberthiir/penis
945 Rivalier/B. aureola s.sp. jatahyana mihi Rivalier det./ TYPE” (black letters on red label); TYPE
LOCALITY, Jatahy (etat de Goyaz)). Mandl, 1963: 587. Vidal Sarmiento, 1966b: 34.

Brasiliella aureola alverengai Mandl, 1963: 586 (TYPE LOCALITY, Jacare P.N. Xingu, M. Grosso, Bras.).

Recognition. — The combination of edentate labrum (Figs. 5a, b), bright or dark
red coppery or brown dorsum with green reflections, and reduced pattern of elytral
maculations (Fig. 27) characterize C. aureola. Female genitalia (Figs. 40, 41) and
male genitalia (Rivalier, 1954: 263; 1955: 90) also distinguish this species.

Synonyms and Types. — The names of the taxa in this complex are based on
comparison of original descriptions with examined specimens and examination of
the type of C. aureola jatahyana Rivalier. We follow Hom (1938: 52, PL 85) in
treating C. cyanitarsis as a morph of C. a. aureola distinguished by a defined middle
band. Members of C. aureola jatahyana Rivalier resemble the C. cyanitarsis form,
with green punctation on the dorsum, which is the prevalent condition in C. aureola.
Cicindela aureola alvarengai Mandl is a dark coppery-brown-red form of C.

Quaest. Ent., 1989, 25 (3)

272

Freitag and Bames

aureola with a slender middle band.

Description. —

Body length. 8.5 mm M, 8. 5-9.0 mm. F.

Body colour. Head and pronotum slightly glossy, elytra dull, almost matte. Dorsum brown with bright
red-coppery (mainly) and green reflections, some specimens with dark dorsum or pronotum darker than
elytra. Venter glossy with coppery, green, and blue reflections.

Body setae. Pleuron sparsely to moderately setose; mesepistemum with a few setae on ventral end only;
metastemum setose laterally; abdominal sterna 1-6 setose mainly on lateral margins.

Other external features. Labrum edentate, middle portion protrudes in female with eight submarginal
setae (Figs. 5a, b). Coupling sulcus of mesepistemum of female in form of sinuate groove. Apex of front
trochanters each with one sensory seta, middle trochanters glabrous. Elytra with slight apical sinuation and
apex recurved; apical spine small; maculations reduced, with one humeral spot, middle band, and subapical
spot, or almost effaced (Fig. 27); punctures very shallow with green reflections; microsculpture isodiametric
and bead-like. Pronotum as in Figure 16.

Female genitalia. Sternum 8 with wide and very shallow V-shaped posterior emargination; each apex
with three very thick setae (Figs. 40a, 41a). Second gonocoxa with several setae on medial margin (Figs.
40a, 41a). Medial portion of second gonapophyses as long as or almost as long as lateral portion (Figs. 40a,
41a). Syntergum 9 and 10 as in Figures 40b, 41b. Ventral sclerite of bursa with posterior end emarginated
and curved dorsally, setae present on lateral margins, median ridge absent (Figs. 40c, d, 41c, d). Membrane
in place of oviduct sclerite (Figs. 40c, 41c). Length of spermatheca and duct unknown (lost in dissection).

Male genitalia. Male genitalia have been described by Rivalier (1954: 263; 1955: 90).

Geographical Variation and Subspecies. — Specimens with discontinuous or
almost effaced maculations on the elytra seem to be predominant in the southern
populations of Paraguay. Darker specimens appear to be more common in the
populations of Matto Grosso, Brazil. Firm application of subspecific names (see
Synonyms and Types section) will depend on whether these characteristics are clinal
in nature or mark geographically distinct populations. Nonetheless the subspecific
names C. a. jatahyana Rivalier and C. a. alvarengai Mandl are probably valid,
representing populations in the northern parts of the species range and Matto Grosso
respectively.

Relationships. — Cicindela aureola and C. horioni are sister species.

Habitat and Period of Activity. — Specimens have been captured in December in
Matto Grosso. Habitat unknown.

Geographical Distribution, Localities, Examined Specimens. — Mainly inland,
southern Brazil (Fig. 49).

Brazil. Amazonas : Manaus, 2F, MZSP. Matto Grosso : Vacaria, 5M, 3F, MZSP. Sao Paulo : Sao Paulo,
IF, BMNH, 1M, MZSP.

Paraguay, no locality, IF, MZSP.

The misella group

Adults of this group are characterized by a broad and shallow apical
emargination in abdominal sternum 8 and narrow oviduct sclerite in the female, and
one or two sagittal sclerotized plates in the internal sac of the male. The group has
five species, of which two are found in Brazil, C. dolosula and C. misella. The
geographical range of the misella group extends northward from northern Argentina
to Venezuela and Guatemala, and is centered in southeastern Brazil and eastern

Brazilian Species of Cicindela

273

Paraguay. The misella group is a distant relative of the aureola, argentata,
hemichrysea and viridicollis groups.

Cicindela (Brasiella) dolosula Rivalier
Figs. 6, 17, 28, 50

Brasiella dolosula Rivalier, 1955: 95 (TYPE, a male in the MNHP general collection bearing the following

labels: “1114 86” (label green on one side)/ “Salobro prov. de Bahia L. Gounelle 6.7.1885”/(extracted

male genitalia)/”#, dolosula (type) mihi E. Rivalier det ./ Museum Paris Gounelle 1114-86/TYPE”

(Black letters on red label); TYPE LOCALITY, Salobro, province de Bahia).

Recognition. — Specimens of C. argentata and C. dolosula are nearly identical
except for striking differences in the internal sac of the male genitalia (female
genitalia are unknown). Most specimens of C. dolosula are distinguished as follows:
pronotum with sides feebly convex (less convex than that of C. argentata
specimens); humeral spot almost always absent and the subhumeral spot tiny and
scarcely visible; male genitalia resemble that of C. misella (Rivalier, 1955: 95). See
also Recognition section of C. misella.

Description. —

Body length. 7.0 - 7.5 M, female unknown.

Body colour. Head and pronotum dull to slightly glossy with coppery tint; elytra dull, coppery brown
with green punctuation; pleuron coppery-green; venter with green, blue, coppery reflections.

Body setae. Pronotum sparsely setose on lateral and front margins and a few setae on front central
portions; pleuron, and lateral portions of abdominal sterna 1-6 moderately setose; mesepistemum with a few
ventral setae.

Other external features. Labrum slightly projected in front, tridentate, with eight submarginal setae (this
number varies from six to 10) (Fig. 6). Pronotum elongate, sides feebly convex, sutures shallow (Fig. 17).
Apex of front trochanters with one sensory seta. Elytra obliquely rounded behind, slightly recurved to small
apical spine. Humeral spot absent from shoulder (or almost absent), subhumeral spot very tiny, almost
effaced; marginal band short, middle band and apical lunule complete; punctures large, shallow, with green
reflections (Fig. 28); fine erect setae present near shoulder.

Genitalia. Female specimens have not been examined. Male genitalia have been described by Rivalier
(1955: 95).

Geographical Variation. — The following notes on variation are based on
observations by Rivalier (1955: 95-97). The number of submarginal setae on the
labrum is mainly 8 but varies from 6-10; and the humeral spot on the shoulder of the
elytra is almost always absent. No geographical pattern in the variation of these
characteristics has been found.

Relationships. — Cicindela dolosula and C. misella are sister species.

Habitat and Period of Activity. — Adults have been collected in November.
Habitat unknown.

Geographical Distribution, Localities, Examined Specimens. — Widespread
inland, from the eastern Brazilian highlands and northern Argentina to northwestern
Brazil and the Canal Zone (Fig. 50).

Argentina. Formosa: Guaycolec, IF, MZSP. Jujuy: Dique La Dienaga, 3M, 5F, MZSP.

Brazil. Bahia: Salobro* (type), 1M, MNHP. Espirito Santo: Linhares, 3M, 3F, RRMC; Rio Itabapoana,
1M, MZSP. Goias: Chapada, 2M, ICCM; Deodoro (Federal District), 1M, MZSP; Dianopolis, 6M, 3F,
MZSP. Matto Grosso: Barra do Tapirape, IF, MZSP; Tres Lagoas, 1M, MZSP; Xingu, 1M, MZSP. Minas
Gerais: Unai, 1M, MZSP. Para: Cachimbo, 16!, MZSP. Roraima: Rio Mucajai south of Boa Vista, 1M,

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Freitag and Bames

MZSP. Rio de Janeiro : Campos, 1M, RRMC; no locality, 1M, 2F, ICCM. Sao Paulo : Barueri, 2M, IF,
MZSP; Campos do Jordao, 8M, 8F, MZSP; Estrada Rio*, (km 47), 1M, IF, MZSP; Ilha da Vitoria*, IF,
MZSP; I tu, 2M, 3F, MZSP; Nova Europa*, IF, MZSP.

Cicindela (Brasiella) misella Chaudoir
Figs. 7, 18, 29, 42, 50

Cicindela misella misella Chaudoir, 1854: 121 (TYPE, a female in MNHP general collection bearing the
following label: “Columbi e/misella Chaud./ Museum Paris type de Chaudoir ex collection
Chaudoir/TYPE” (black letters on red label); TYPE LOCALITY, la Columbie). Bates, 1881: 14. Horn
1915: 406; 1926a: 308. Blackwelder, 1944: 17. Rivalier, 1954: 263; 1955: 93. Vidal Sarmiento, 1966b:
34. Sumlin, 1979: 104.

Cicindela misella transversalis Rivalier, 1955: 95 (TYPE, a male in the MNHP general collection bearing
the following labels: “Guatemala” (green label)/ /’’Museum Paris ex. coll. M. Maindron coll. G. Babault
1930/penis 924 Rivalier /misella m. transversalis mihi type E. Rivalier det.”; TYPE LOCALITY,
Guatemala).

Recognition. — A tridentate labrum with a non-protruding front margin (Figs. 7a,
b), and small humeral spot hidden from dorsal view but present on the latero-ventral
side of the elytral shoulder together distinguish adults of C. misella from those of C.
argentata and C. dolosula. Because these characteristics vary, a decisive
identification can only be made on the basis of the male genitalia.

Description. —

Body length, ca 6.5 mm M and F.

Body colour. Head and pronotum slightly glossy black with green and coppery reflections, elytra dull,
dark brown, pleuron and venter glossy to slightly glossy black with coppery, green, and blue reflections.

Body setae. Head glabrous. Pleuron setose, mesepistemum with a few setae near ventral end. Lateral
(mainly) portions of abdominal sterna 1-6 (M,F) with appressed setae.

Other external features. Labrum narrow, tridentate, with lateral teeth broadly rounded, with basically
eight (five to eight) submarginal setae (Figs. 7a, b). Pronotum narrow, broadened anteriorly (Figs. 18a, b).
Coupling sulcus of mesepistemum of female with deep elongate central pit. Apex of front trochanters with
one seta. Elytra with apex not recurved in males, slightly recurved in females, small apical spine (Figs. 29a,
b). Pattern of elytral maculations with small humeral spot hidden on the lateroventral side of shoulder, large
subhumeral spot, broadened complete middle band and apical lunule, marginal band short (Figs. 29a, b).
Punctation of elytra green.

Female genitalia. Sternum 8 with shallow, very broad apical emargination, apices with four or five stout
setae (Fig. 42a). Second gonocoxa and second gonapophyses as in Figure 42a. Syntergum 9 and 10
broadened apically (Fig. 42b). Ventral sclerite flat, elongate, lightly sclerotized (Fig. 42c). Oviduct sclerite
narrow (Fig. 42c). Spermatheca and duct ca 1.0 mm long.

Male genitalia. Described and figured by Rivalier (1954: 263; 1955: 93-95) and Vidal Sarmiento
(1966b: 34).

Geographical Variation. — Maculations of the elytra vary but a geographical
pattern is not evident. Adults of Rivalier’ s C. misella transversalis from Guatemala
have a distinct transverse middle band.

Relationships. — Cicindela misella and C. dolosula are sister species.

Habitat and Period of Activity. — Adults have been collected in March. Habitat
unknown.

Geographical Distribution, Localities, Examined Specimens. — Central America
and northwestern South America (probably east of the Andes) south to southwestern
Brazil and Argentina (Fig. 50).

Brazilian Species of Cicindela

275

Bolivia. Chiquitos* (300 m), 4M, IF, ICCM.

Brazil. Matto Grosso. Corumba, 1M, ICCM.

Colombia. Bugaba (or Buga) (800-1500'), 2M, 2F, BMNH.

Canal Zone. Barro Colorado, IF, MZSP.

The minarum group

Adults of this group are characterized by a median ridge on the ventral sclerite of
the bursa copulatrix of the female, and a short tooth and absence of small stiffening
rib in the internal sac of the male. The group has eight species four of which are
found in Brazil, C. hamulipenis, C, brevipalpis, C. banghaasi, and C. minarum. The
geographical range of the minarum group extends from the Brazilian Highlands
northward to Venezuela and southern Mexico. The minarum group is not closely
related to any other group in subgenus Brasiella.

Cicindela (Brasiella) hamulipenis Horn
Figs. 8, 19, 30, 45,51

Cicindela hamulipenis Horn, 1938: 14, 52, PI. 85, Fig. 1 (HOLOTYPE, a male, in the IPZE collection

bearing the following label: “Goyas/Type W. Hom/Holotypus” (black letters on red label); TYPE

LOCALITY, Goyas (Brasilia center.)). - Blackwelder, 1944: 18.

Recognition. — The combination of protruding unidentate labrum with six or
seven submarginal setae (Fig. 8), reduced pattern of the elytral maculations (Fig.
30), and small body size characterizes this species. The apical end of the median
lobe is broad (Fig. 45a, b) and the internal sac contains sclerites with specific shapes
(Fig. 45d).

Description. —

Body length. 6.0 mm M.

Body colour. Head pronotum and elytra dull, dark brown, with coppery (especially the pronotum) and
green reflections. Pleuron with coppery and green reflections. Venter with green and blue-green reflections.

Body setae. Lateral margins of pronotum and pleuron moderately setose, mesepistemum of male mainly
glabrous except for the ventral end. Abdominal sterna 1-6 setose, mainly on lateral margins.

Other external features. Labrum convex and elongate, unidentate, with five to seven submarginal setae
(Fig. 8). Pronotum widest in anterior 0.5 (Fig. 19). Apex of front trochanters with one seta. Apices of elytra
recurved to very small spine; maculations largely effaced showing remains of subapical humeral spot,
middle band and apical lunule (Fig. 30); punctures of elytra shallow and green, a few larger punctures are
present along median suture and on shoulder; microsculpture isodiametric and bead-like.

Female genitalia. Females were not available for examination.

Male genitalia. The median lobe is like that of other species in Brasiella but broader in the apical end
(Figs. 45a, b); sclerites of the internal sac are specific for C. hamulipenis. Shield bifid apically. Stylet and
arciform piece elongate, slender. Tooth slender, apex pointed. Right bar absent. Flagellum lacking (Fig.
45d).

Relationships. — Cicindela hamulipenis and C. brevipalpis are sister species.

Habitat and Period of Activity. — Adults have been collected in January. Habitat
unknown.

Geographical Distribution, Localities, Examined Specimens. — Only the
Holotype (IPZE) and one other male were examined. Both specimens were collected

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276

Freitag and Bames

in Goias, the non-Holotype in Dianopolis, 1 1-14.1.1962 (Fig. 51).

Brazil. Goias : Dianopolis, 1M, MZSP; no locality, 1M IPZE.

Cicindela ( Brasiella ) brevipalpis Horn
Figs. 9, 20,31,46,51

Cicindela brevipalpis Horn, 1926b: 77 (TYPE LOCALITY, Vacarias von Sud-Matto Grosso); 1926a: 308:

1938: 52. Blackwelder, 1944: 17.

Recognition. — The character state of small body size, ca 6.0 mm long, elytra
that lack both a distinct shoulder at the anterior end and maculations (Fig. 31), and
an indistinct 5-dentate labrum and 6 submarginal setae (Fig. 9) taken together are
diagnostic for adults of C. brevipalpis.

Synonyms and Types. — The name C. brevipalpis Horn is based on comparison
of the original description with a male specimen on loan from IPZE labelled: “Matto
Grosso Vacaria XII-22/Type W. Hom/Syntypes” (black letters on a red label).

Description. —

Body length, ca 6.0 mm M.

Body colour. Body dull, black, with some coppery and green reflections on the head, thorax and elytra.

Body setae. Pronotum with a few setae on lateral margins and central disc. Pleuron and lateral portions
of the metastemum with appressed setae, only the ventral end of the mesepistemum setose; abdominal sterna
3-6 inclusive sparsely setose.

Other external features. Head with protruding eyes. Labrum indistinctly five dentate, six submarginal
setae, with the middle four close to the margin (Fig. 9). Pronotum narrow, broader at the anterior end (Fig.
20). Tibia rufotestaceous. Apex of front trochanters with one sensory seta, middle trochanters glabrous.
Elytra lacking distinct shoulder, apices not recurved, apical spine well developed; maculations absent;
punctures very shallow and marked with green (mainly) and coppery spots (Fig. 31); microsculpture
isodiametric, bead-like. Flight wings absent.

Female genitalia. Females were not available for dissection.

Male genitalia. Apex of median loabe with short protruding tip; not hooked. Internal sac with at least
one apical setal brush. Flagellum lacking. Sclerites of internal sac very well sclerotized; the shield bears two
sharp apices, arciform piece long and strongly oblique, stylet obliquely truncated at apex, tooth of moderate
size and very thin in the apical 0.5 (Figs. 46a-c).

Relationships. — - Cicindela brevipalpis and C. hamulipenis are sister species.

Habitat and Period of Activity. — The syntypes were found running quickly
between grass in sparse grasslands in December 1922 (Horn, 1926b: 78).

Geographical Distribution, Localities, Examined Specimens. — The syntypes,

collected in Vacaria, Matto Grosso (Fig. 51), are the only known specimens.

Brazil. Matto Grosso: Vacaria, 1M, IPZE.

Cicindela (Brasiella) banghaasi Horn
Figs. 10,21,32, 43,51

Cicindela banghaasi Horn, 1907: 24 (TYPE LOCALITY, Cuyaba, Matto-Grosso); 1915: 408; 1923: 112;

1926a: 310; 1938: 53. Blackwelder, 1944: 17. Rivalier, 1954: 263; 1955: 98.

Recognition. — The character state combination of the sparsely setose frons and
gena, unidentate labrum with 8 setae, pattern of elytral maculations with a broad
oblique middle band (Fig. 32), round pit-like coupling sulcus of the mesepistemum

Brazilian Species of Cicindela

277

of the female, and structure of female genitalia (Figs. 43a-c) distinguishes C.
banghaasi from other Brazilian species of Cicindela. Also structures of the internal
sac of the male are specific (Rivalier, 1955:98).

Synonyms and Types. — The name C. banghaasi is based upon comparison of the
original description with six syntypes, one female and five males, on loan from the
IPZE, each labelled as follows: “Staudinger Cuyaba Matt. Gr./ Type! Dr. W.
Hom/Syntypes”; and with one large group label “ Cicindela Banghaasi H.6”.

Description. —

Body length. 6.5-7.0 mm M, 7.5 mm F.

Body colour. Dorsum dull; head and pronotum black or very dark brown with some coppery and green
reflections; elytra very dark brown with coppery reflections and large green puncture spots. Pleuron coppery
(mainly), blue and green. Venter black with blue-green (mainly) and some coppery reflections.

Body setae. Head with a few setae between the eyes, on the frons, clypeus in some specimens, and
genae. Pronotum and pleuron moderately setose. All abdominal sterna setose, more densely so laterally.

Other external features. Labrum unidentate to tridentate (indistinct) with eight submarginal setae (Fig.
10). Pronotum broader anteriorly (Fig. 21). Coupling sulcus of mesepistemum of female a deep round pit.
Apex of front trochanters with one sensory seta. Elytra with recurved apex to very small spine; humeral spot
and subhumeral spot small, marginal band and apical lunule broad, middle band broad and oblique;
punctation green, with a row of much larger green umbilicate punctures along with the median sutures, and
some intermediate in size clustered near the shoulder (Fig. 32); a few erect setae present near shoulder;
apical microserrulations very small.

Female genitalia. Sternum 8 with wide V-shaped emargination in apical end, apices each with three
stout setae and two smaller setae (Fig. 43a). Second gonocoxa with a few setae near apical end of medial
margin; and second gonapophyses as shown in Figure 43a. Syntergum 9 and 10 as shown in Figure 43b.
Ventral sclerite with a round darkened basal end and central broad median ridge well developed (Fig. 43c).
Membrane in place of oviduct sclerite (Fig. 43c). Spermatheca and duct (broken in dissection), short,
approximately 1 .0 mm.

Male genitalia. Male genitalia have been described by Rivalier (1955: 98).

Relationships. — C. banghaasi is sister to the lineage that gave rise to sister
species C. hamulipenis and C. brevipalpis.

Habitat and period of activity. — Unknown.

Geographical Distribution, Localities, Examined Specimens. —

Cuyaba, Matto Grosso (Fig. 51).

Brazil. Matto Grosso : Cuyaba, IF, 5M, IPZE, IF, RRMC.

Cicindela (Brasiella) minarum Putzeys
Figs. 11,22, 33,44,51

Cicindela minarum Putzeys, 1845: 369 (TYPE LOCALITY, Bresil. Province des Mines.). Horn, 1915: 406;

1926a: 307; 1938: 52. Blackwelder, 1944: 18. Rivalier, 1954: 263; 1955: 92,98,99.

Recognition. — The character combination of the short unidentate labrum with
7-12 submarginal setae (Figs. 11a, b), oblique truncation of the apical end of the
elytra and colour pattern of elytral maculations (Figs. 33a, b) distinguishes C.
minarum from other species of Cicindela.

Description. —

Body length, ca 8.0 mm M, ca 8.5 mm F.

Body colour. Dorsum dull, black with copper and green reflections. Pleuron copper (mainly) and some
green. Venter black with mainly blue and purple reflections.

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Body setae. Pronotum with a few setae along lateral margin and middle suture. Proepistemum and
mesepistemum with a few sparse setae mainly in ventral 0.5; other pleural sclerites, the lateral 0.5 of the
metastemum and abdominal sterna 1 -6, moderately setose.

Other external features. Labrum short and unidentate, almost tridentate, with seven (M) to 12 (F)
submarginal setae (Figs. 11a, b). Pronotum square-shaped with parallel sides. Coupling sulcus of
mesepistemum of female a deep groove with a pit in the center. Apex of front trochanters with one seta.
Apical end of elytra obliquely truncated and recurved to small apical spine (Figs. 33a, b). Pattern of elytral
maculations reduced with subapical spot present or absent, marginal band short, discontinuous sinuate
middle band, and complete apical lunule with recurved hook-shape at distal end (Figs. 33a, b). Punctation
large, shallow, blue and green, and sparsely distributed. Microsculpture isodiametric, bead-like.

Female genitalia. Sternum 8 with broad V-shaped apical emargination, apices each with a group of three
stout setae (Fig. 44a). Second gonocoxa with several setae on medial margin (Fig. 44a). Gonapophyses
short, stout (Fig. 44a). Syntergum 9 and 10 as in Figure 44b. Ventral sclerite darkly sclerotized; median
ridge well developed (Fig. 44c). Membrane in place of oviduct sclerite (Fig. 44c). Spermatheca and duct
length unknown (broken in dissection).

Male genitalia. Male genitalia have been described by Rivalier (1955: 99).

Relationships. — Cicindela minarum is sister to the lineage that gave rise to C.
nebulosa Bates (Nicaragua to Colombia and Ecuador) and sister species C. mandli
Brouerius van Nidek (Chiapas, Mexico) and C. insularis Brouerius van Nidek
(Trinidad).

Habitat and Period of Activity. — Specimens of C. minarum have been collected
in December and April. Habitat unknown.

Geographical Distribution, Localities, Examined Specimens- Geographical

distrubution mainly inland in southern Brazil (Fig. 51).

Brazil. Espirito Santo: no locality, IF, IOC, 1M, MZSP. Matto Grosso: Vacaria, 1 2F, MZSP. Minas

Gerais: Mar de Espanha*, 1M, IF, IOC, 1M, IF, MZSP, IF, RRMC; PassaQuatro, 1M, xF, IOC.

Subgenus Gaymara new subgenus

Type species. — Cicindela chlorosticta Kollar, 1836: 332 (here selected).

Derivation of name. — From first given names of Gayla E. Freitag, Gavin W.
Freitag, Margot E. Freitag and Barbara L. Bames (junior author), who have assisted
the senior author in studies on tiger beetles.

Recognition. — Adults of this subgenus are distinguished by the five characters
given in couplet 4 of the key. In addition the head is mainly glabrous, with elongate
tridentate labrum; elytral maculations are partially reduced; the apex of the median
lobe of the male consists of a short ventral hook (straight in C. anulipes ), and a
flagellum is absent from the internal sac; the spermatheca and duct together are
approximately 1.0 - 2.0 mm long. (See the Recognition section of subgenus
Brasiella for similarities between it and the subgenus Gaymara.

Species groups. — This subgenus has two groups, the chlorosticta group and
anulipes group, that together include five species. Both groups are found in eastern
South America mainly in southeastern Brazil.

Geographical distribution. — The range of this subgenus extends from northern
Argentina northward to southern parts of the Amp n basin.

Brazilian Species of Cicindela

279

Figs. 52-56. Labrum, dorsal aspect. 52, Cicindela chlorosticta Kollar: (a) female, Esperanca do Sul, S.P.; (b)
male, Jatai, Go.; 53, C. staudingeria Horn: (a) female, Jatai, Go.; (b) male, Vacaria, M. Grosso; 54, C.
nigroreticulata Horn, Ronda Alta, R.G. do Sul: (a) female; (b) male; 55, C. paranigroreticulata n.sp.,
Tramandai, R.G. do Sul: (a), (b) female; (c) male; 56, C. anulipes Horn, female, Utiariti, Rio Papagaio, M.
Grosso. Figs. 57-61. Pronotum, dorsal aspect. 57, Cicindela chlorosticta Kollar: (a) female; (b) male; 58, C.
staudingeria Horn: (a) female; (b) male; 59, C. nigroreticulata Horn: (a) female; (b) male; 60, C.
paranigroreticulata Horn n. sp.: (a), (b) female; (c) male; 61, C. anulipes Horn, female.

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Figs. 62-66. Elytron, dorsal aspect. 62, Cicindela chlorosticta Kollar: (a) female; (b) male; 63, C.
staudingeria Horn: (a) female; (b) male; 64, C. nigroreticulata Horn: (a) female; (b) male; 65, C.
paranigroreticulata n. sp.: (a), (b) female; (c) male; 66, C. anulipes Horn, female.

Brazilian Species of Cicindela

281

Figs. 67-71. Female genitalia: (a) sternum 8, second gonocoxa, second gonapophyses, ventral aspect; (b)
syntergum 9&10, dorsal aspect; (c) bursa copulatrix, oviduct sclerite, oviduct, spermatheca and duct, ventral
sclerite, ventral aspect. 67, Cicindela chlorosticta Kollar; 68, C. staudingeria Horn; 69, C. nigroreticulata
Horn; 70, C. paranigroreticulata n. sp.; 71, C. anulipes Horn.

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Figs. 72-76. Male genitalia: median lobe (a) right lateral aspect; (b) dorsal aspect; (c) internal sac, arciform
piece ( ap ), central plate ( cp ), right bar (rb), spiny rod (sr), stylet (st), right lateral aspect; (d) left lateral
aspect. 72, Cicindela chlorosticta Kollar; 73, C. staudingeria Horn; 74, C. nigroreticulata Horn; 75, C.
paranigroreticulata n. sp.; 76, C. anulipes Horn. Localities for Figures 57-76 as for 52-56.

Brazilian Species of Cicindela

283

Fig. 77. Map showing the geographical distribution of the species Cicindela chlorosticta Kollar (•); C.
staudingeria Horn (■) C. nigroreticulata Horn (A); C. paranigroreticulata n. sp. (♦); C. anulipes Horn (+).
Open squares and open circle with a cross represent state records for C. staudingeria Horn and C. anulipes
Horn respectively.

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Phylogenetic relationships. — Subgenus Gaymara is sister to the lineage that
gave rise to Plectographa and Cylindera.

The chlorosticta group

Adults of this group are characterized by two swirls of rugosity between eyes,
broad pronotum, and large elytral foveae. The group has four species all of which
are found in Brazil, C. chlorosticta, C. staudingeria, C, nigroreticulata, and C.
paranigroreticulata. The geographical range of the chlorosticta group extends from
southeastern Brazil into southern parts of the Amazon basin. The chlorosticta and
anulipes groups are sister groups.

Cicindela ( Gaymara ) chlorosticta Kollar
Figs. 52, 57, 62, 67, 72, 77

Cicindela chlorosticta Kollar, 1836: 332 (TYPE LOCALITY, in Brasiliae provincia Ypanema). Horn, 1915:

404; 1926a: 306; 1938: 52. Blackwelder, 1944: 17. Rivalier, 1954: 264; 1955: 99. Vidal Sarmiento,

1966a: 257; 1966b: 35. Sumlin, 1979: 105.

Cicindela chlorosticta smaragdina Horn, 1893: 198 (TYPE LOCALITY, Sao Paulo); 1915: 404; 1926a:

306. Blackwelder, 1944: 17.

Recognition. — Adults of the sister species of C. chlorosticta and C. staudingeria
are distinguished from other Brazilian tiger beetles by a deeply rugose head and
pronotum, bright coppery dorsum and pleuron, and pattern of elytral maculations, in
particular the oblique middle band, and row of large foveae along the median suture
of elytra (Figs. 62, 63). Adults of C. chlorosticta are distinguished from those of C.
staudingeria by the smaller body size; a relatively narrow pronotum (Fig. 57 cf. Fig.
58); four setae on apex of each lobe of sternum 8 (Fig. 67a cf. Fig. 68a); very
elongate pair of posterior extension of the ventral sclerite of the bursa copulatrix and
narrow oviduct sclerite (Fig. 67c cf. Fig. 68c) in the female; and relatively shorter
tooth (spiny field) and stylet (spiny rod) of the internal sac of the male (Figs. 72c, d
cf. Figs. 73c, d).

Synonyms and Types. — Adults of C. smaragdina are blue or green members of
C. chlorosticta found in Sao Paulo. We follow Horn, 1926a: 404 in treating C.
smaragdina as a synonym of C. chlorosticta. These colour morphs appear to be rare.

Description. —

Body length. 8.0-8. 5 mm M, ca 8.5 mm F.

Body colour. Head and pronotum deeply rugose, bright coppery and with some green. Elytra matte light
brown or coppery, combined with a little green in some specimens. A few adults with blue or green dorsum.
Pleuron glossy and bright coppery. Venter mainly coppery with some green, purple and blue; center of
abdomen black with mainly purple reflections.

Body setae. Head glabrous. Pleuron sparsely to moderately setose. Lateral portions of abdominal sterna
1-5 (F) and 1-6 (M) with appressed setae.

Other external features. Vertex of head with two wide shallow pits (swirls of rugosity) between hind
margins of eyes. Labrum elongate tridentate, with four to six submarginal setae (Figs. 52a, b), Pronotum
broadened anteriorly, sides not protruding (Figs. 57a, b). Coupling sulcus of mesepistemum of female a

Brazilian Species of Cicindela

285

long, moderately deep groove. Apex of front trochanters with one seta. Elytra with apex slightly recurved to
a well developed spine in females (Figs. 62a, b). Pattern of elytral maculations with discontinuous humeral
lunule and oblique complete or discontinuous middle band (Figs. 62a, b). Punctation of elytra green; with a
row of foveate punctures along median suture and a cluster near shoulder (Figs. 62a, b).

Female genitalia. Sternum 8 with shallow and broad apical emargination, apices broadly rounded, each
apex with four stout setae (Fig. 67a). Second gonocoxa with four or five setae in medial margin (Fig. 67a).
Second gonapophyses short (Fig. 67a). Syntergum 9 and 10 as in Figure 67b. Ventral sclerite with two
elongate posterior projections; row of fine setae on anterior margins (Fig. 67c). Oviduct sclerite narrow (Fig.
67c). Spermatheca and duct probably short (broken in dissection).

Male genitalia. Apical end of median lobe narrowly tapered (Figs. 72a, b). Internal sac without
flagellum. Large tooth in the form of an elongate spiny field consisting of two lobes. Stylet joined to base of
tooth short with pointed apex. Spiny rod extended beyond apical end of tooth. Arciform piece moderately
broad. Central plate large (Figs. 72c, d).

Geographical Variation. — The dorsal colour and pattern of elytral maculations
vary slightly but a geographical pattern is not evident.

Relationships. — Cicindela chlorosticta and C. staudingeria are sister species.

Habitat and Period of Activity. — Adults have been collected from October to
March. Habitat unknown.

Geographical Distribution, Localities, Examined Specimens. — Central and

southeastern Brazil to northeastern Argentina and Paraguay (Fig. 77).

Brazil. Goias : Chapada, 2M, ICCM; Jatai, 2M, IOC, 3M, MZSP; Rio Verde, 1M, IOC. Matto Grosso:
Utiariti, 1M, MZSP; Vacaria, 2M, 2F, MZSP. Sao Paulo : Avanhand (Garbe L.)*, 4M, 6F, MZSP; Eng.
Coelho*, 1M, IF, IOC; Esperanca do Sul*, 1M, IF, MZSP; Guatapara*, 1M, 2F, MZSP; Onda Verde*, IF,
MZSP; Piraju, 1M, IF, MZSP.

Paraguay. Caaguazu, 1M, IF, MNRJ.

Cicindela (Gaymara) staudingeria Horn
Figs. 53, 58, 63, 68, 73, 77

Cicindela staudingeri Horn, 1892c: 368 (not Kraatz) (TYPE LOCALITY, Sao Paulo); 1915: 404; 1926a:
306. Blackwelder, 1944: 17. Sumlin 1979: 105.

Cicindela staudingeria Horn, (replacement name for Cicindela staudingeri Horn), 1915: 404; 1926a: 306;
1938: 52. Blackwelder, 1944: 17. Rivalier, 1954: 264; 1955: 99. Sumlin, 1979: 105

Recognition. — Adults of C. staudingeria are distinguished from other Brazilian
tiger beetles by character combination of the large body size, ca 10.0-12.0 mm long,
markedly rugose head and pronotum, two wide fairly deep pits between the hind
margins of the eyes, bright coppery reflections from the dorsum and pleuron,
particularly from the proepistemum, and pattern of elytral maculations (Figs. 63a,
b). See also Recognition section for C. chlorosticta.

Synonyms and Types. — The name C. staudingeria is based upon comparison of
the original description of C. staudingeri Horn with four males and one female in
the IPZE, each labelled as: “Staudinger, Sao Paulo/Type ! Coll. W. Hom/Syntypus
Cicindela staudingeri Horn 5”.

Description. —

Body length. 10.0-1 1.0 mm M, 12.0 mm F.

Body colour. Head and pronotum deeply rugose, bright coppery and some green; elytra matte light
brown and coppery and faintly green in some specimens. Pleuron glossy and bright coppery. Venter mainly

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coppery with some green, blue and purple reflections; center of abdomen black with some blue, green or
purple.

Body setae. Pleuron sparsely setose. Lateral margins of abdominal sterna 1-5 (F) and 1-6 (M) covered
with appressed setae.

Other external features. Vertex of head of most adults with two wide shallow pits between hind margins
of eyes. Labrum elongate, tridentate, weakly so in some specimens, with four to seven (most commonly six)
submarginal setae (Figs. 53a, b). Pronotum broad, sides ampliate, in most specimens sutures moderately
deep (Figs. 58a, b). Coupling sulcus of mesepistemum of female a long and moderately deep groove. Apex
of front trochanters with one sensory seta. Elytra with apex slightly recurved to a well developed spine (Figs.
63a, b). Pattern of elytral maculations with discontinuous to almost effaced humeral lunule, middle band
oblique, marginal band and apical lunule complete (Figs. 63a, b). Punctation green. Large foveate punctures
along median suture and a cluster of smaller ones near middle of shoulder (Figs. 63a, b).

Female genitalia. Sternum 8 with shallow and broadly rounded V-shaped apical emargination, each
apex with five stout fairly long setae (Fig. 68a). Second gonocoxa with several setae in medial margin (Fig.
68a). Second gonapophyses elongate (Fig. 68a). Syntergum 9 and 10 as in Figure 68b. Ventral sclerite with
two posterior projections, recurved apex and row of fine setae on apical end (Fig. 68c). Oviduct sclerite short
and broad (Fig. 68c). Spermatheca and duct probably short (broken in dissection).

Male genitalia. Apical end of median lobe narrowly tapered (Figs. 73a, b). Internal sac without
flagellum. Very long bilobed spiny field in place of the large tooth. Stylet joined to base of tooth, short, apex
sharp, bent. Slender spiny rod extended far beyond apical end of tooth. Arciform piece long; central plate
large. Right bar wide (Figs. 73c, d).

Geographical Variation. — Dorsal colour, pattern of elytral maculations, and
number of submarginal setae of the labrum vary within populations, though
geographical patterns of variation are not apparent.

Relationships. — Cicindela staudingeria and C. chlorosticta are s ter species.

Habitat and Period of Activity. — Adults have been collected \ >m October to
March. Habitat unknown.

Geographical Distribution, Localities, Examined Specimens. — Southeastern

Brazil to northern Argentina and Uruguay (Fig. 77).

Brazil. Golds'. Chapada, 6M, 5F, ICCM: Jatai, IF, IOC, IF, MZSP. Matto Gro sso :Murtinho*, 1M, IOC;
no locality, 2M, 3F, MZSP. Sao Paulo : Batatais, 1M, IOC; no locality, 1M, IOC, 1M, IF, MZSP.

Uruguay. Rivera, 2M, MNRJ; Tacuarembo, IF, MNRJ.

Cicindela (Gaymara) nigroreticulata Horn
Figs. 54, 59, 64, 69, 74, 77

Cicindela nigroreticulata Horn, 1927: 139 (TYPE, a female in IPZE bearing the following label: “Cochilha
Grande R. Grande d. Sul/Type W. Horn/ Holotypus (black letters on red label)/ nigroreticulata”-, TYPE
LOCALITY, Cochilha Grande, Rio Grande do Sul); 1938: 52. Blackwelder, 1944: 19.

Recognition. — Adults of C. nigroreticulata are distinguished from those of other
Brazilian species by the colouration of the elytra and pattern of elytral maculations.
Many dark brown lines form a reticulated pattern against the lighter brown and
coppery ground colour, and the middle band is strongly curved (Figs. 64a, b). In
addition, the labrum is unidentate or weakly tridentate with 4 submarginal setae
(Figs. 54a, b), and a pair of wide pits are located between the hind margins of the
eyes.

Description. —

Body length. 9.0 mm M, ca 9.0-9.5 mm F.

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287

Body colour. Dorsum of head and pronotum slightly glossy, with bright coppery and green reflections;
elytra dull brown coppery or green, with a reticulated pattern formed by many dark brown lines (Figs. 64a,
b). Venter glossy, with green and coppery reflections.

Body setae. Pleuron, lateral portions of abdominal sterna with sparsely to moderately dense appressed
setae, most of the mesepistemum glabrous.

Other external features. Vertex of head and pronotum deeply rugose. Two wide pits located on vertex
between hind margins of eyes. Tooth of men turn elongate. Labrum unidentate or weakly tridentate, four
submarginal setae (Figs. 54a, b). Pronotum broad, sides ampliate, sutures deep (Figs. 59a, b). Coupling
sulcus of mesepistemum of female a groove with center depressed. Legs red-coppery throughout. Apex of
front trochanters with one seta. Elytra tapered apically, apex recurved to short apical spine. Maculations of
elytra separate and clearly defined; humeral lunule short not reaching central disc, middle band strongly
curved (Figs. 64a, b); punctation almost effaced, large shallow punctures near shoulders and anterior
portions of disc beside median suture.

Female genitalia. Sternum 8 with shallow V-shaped posterior emargination; apices broadly rounded
each with four to six short thick setae (Fig. 69a). Second gonocoxa with a few setae on median edge (Fig.
69a). Second gonapophyses short and broad at base (Fig. 69a). Syntergum 9 and 10 broadened posteriorly
(Fig. 69b). Ventral sclerite bulbous, with two elongate posterior projections, row of fine setae on anterior
dorsal margin (Fig. 69c). Oviduct sclerite shield-like, well sclerotized (Fig. 69c). Spermatheca and duct
elongate (broken in dissection), greater than 1.5 mm in length.

Male genitalia. Apical end of median lobe slender, apex with sharp ventral emargination (Figs. 74a, b).
Internal sac without flagellum. Stylet short sinuate, apex sharp, joined to base of tooth. Elongate
unsclerotized spiny field-like bilobed tooth and thin rod each of equal length. Two basal membranous lobes
present. Arciform piece slender; central plate large. Right bar narrow, elongate (Figs. 74c, d).

Geographical Variation. — The pattern of reticulations on the elytra varies
appreciably within and among populations.

Relationships. — Cicindela nigroreticulata and C. paranigroreticulata are sister
species.

Habitat and Period of Activity. — Adults have been collected in February.
Habitat unknown.

Geographical Distribution, Localities, Examined Specimens. — Inland Rio
Grande do Sul, Brazil (Fig. 77).

Brazil. Rio Grande do Sul: Cochilha Grande, IF (holotype), IPZE; Ronda Alta*, 1M, 2F, MNRJ; Santo
Augusto*, 2M, IF, ICCM.

Cicindela (Gaymara) paranigroreticulata new species
Figs. 55, 60, 65, 70, 75, 77

Type Specimens. — HOLOTYPE, a female in the collection of MNRJ labelled:
“Brasil Rio Gde. do Sul Tramandai 14-11-64 C.S. Carbonell A. Mesa y M.A.
Monne/Holotype Cicindela paranigroreticulata R. Freitag, B.L. Barnes” (red label).
PARATYPES, one male and one female in the MNRJ labelled same as Holotype,
but type label is: “Paratype Cicindela paranigroreticulata R. Freitag and B.L.
Barnes” (blue label). TYPE LOCALITY, Tramandai Rio Gde do Sul.

Recognition. — The character state combination of black body, large body size,
8. 0-9.0 mm long, extensive head and body pubescence, setose elytra, and pattern of
elytral maculations (Figs. 65a-c) distinguish adults of Cicindela paranigroreticulata
from those of other Brazilian species of Cicindela.

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Description. — Body length. 8.0 mm M, ca 9.0 mm F.

Body colour. Head, pronotum, pleuron and venter slightly glossy; elytra dull. Body black with obscure
green and coppery reflections on the elytra and pleuron. Four basal articles of the antenna coppery and
green; articles 5-11 moderate pale rufopiceous. Legs with coppery and green reflections.

Body setae. Vertex, frons, clypeus and genae with scattered setae, and a tuft of setae on the front inner
margins of each eye. Pronotum setose, except for central disc. Elytra sparsely setose mainly on basal 0.5,
medial along suture and lateral margins. Pleuron, lateral portion of metastemum, and later portions of
abdominal sterna moderately setose. Dorsal 0.5 of mesepistemum glabrous.

Other external features. Vertex of head deeply rugose. Two obsolete wide pits (swirls of rugosity) on
vertex between hind margins of eyes. First antennal article with one long apical seta and one short medial
seta. Tooth of mentum elongate. Labrum elongate tridentate; six to eight submarginal setae (Figs. 55a-c).
Pronotum broad expanded in apical 0.5 (Figs. 60a-c). Coupling sulcus of mesepistemum of female an
elongate groove with deep middle. Procoxa, mesocoxa and lateral portion of metacoxa moderately setose.
Apex of front trochanters with one seta. Femora, tibia, and tarsi moderately setose. Apex of elytra recurved
to small spine; maculations of elytra reduced, humeral lunule short, middle band discontinuous not reaching
lateral margin of elytra, apical lunule elongate (Figs. 65a-c). Elytral punctations shallow marked by green
and coppery metallic spots; a row of larger punctures near median suture and another short row inside the
shoulder; microsculpture mainly isodiametric. Apices of elytra with microserrulations.

Female genitalia. Sternum 8 with shallow U-shaped posterior emargination; apices broadly rounded
each with five short thick setae (Fig. 70a). Second gonocoxa with a few setae on medial edge (Fig. 70a).
Second gonapophyses elongate (Fig. 70a). Syntergum 9 and 10 broadened posteriorly (Fig. 70b). Ventral
sclerite wide, lightly sclerotized, two elongate posterior projects slightly curved medially, row of fine setae
on dorsal anterior margin (Fig. 70c). Oviduct sclerite lightly sclerotized, almost transparent, shield-shaped
with lateral flanges (Fig. 70c). Length of spermatheca and duct ca 2.0 mm.

Male genitalia. Apical end of median lobe slender, apex sharply hooked posteriorly (Figs. 75a, b).
Internal sac was accidentally everted. Flagellum absent. Stylet joined to base of tooth very short, stout with
pointed apex. Spiny field-like bilobed tooth and two rod-like extensions of equal length. Arciform piece
slender; central plate large. Right bar was not seen (Figs. 75c, d).

Relationships. — Cicindela paranigroreticulata and C. nigroreticulata are sister
species.

Habitat and Period of Activity. — Type collected on or beside ocean shore,
February or November, 1964.

Geographical Distribution, Localities, Examined Specimens. — Known only
from Tramandai, Rio Grande do Sul, Brazil (Fig. 77).

Brazil. Rio Grande do Sul: Tramandai, IF (holotype), 1M, IF, (paratypes), MNRJ.

The anulipes group

Adults of this group are characterized by some or all setae of labrum positioned
very near the anterior margin, and long mentum tooth. The group has only one
species, C. anulipes, which is found in southeastern Brazil and the Brazilian
Highlands. This group and the chlorosticta group are sister groups.

Cicindela (Gaymara) anulipes Horn
Figs. 56,61,66,71,76, 77

Cicindela anulipes Horn, 1897a: 255 (TYPE, a male in the IPZE bearing the following labels: “Staudinger,
Minas Geraes/Type !, Dr. W. Horn/ Holotypus” (black letters on red label) “ Cicindela anulipes Horn”;
TYPE LOCALITY, Minas Geraes); 1915: 405; 1923: 112; 1926a: 307; 1938: 58. Blackwelder, 1944:
16. Rivalier, 1954: 264; 1955: 99.

Brazilian Species of Cicindela

289

Recognition. — The character state combination of the elongate and weakly
tridentate labrum, deep and evenly distributed punctures of the proepistemum,
pattern of the elytral maculations reduced in extent (Fig. 66), sclerotized oviduct
sclerite and shape of ventral sclerite of the bursa copulatrix distinguishes C. anulipes
from other species of Cicindela in Brazil. The internal sac of the male contains a
unique bifid stylet (Fig. 76c).

Synonyms and Types. — The name is based upon examination of the type
specimen and other specimens on loan.

Description. —

Body length. 7. 0-7. 5 mm M, 7.5 mm F.

Body colour. Dorsum dull, black; elytra matte and slightly velvety. Venter and pleuron balck with green,
coppery and blue reflections; legs with testaceous sections.

Body setae. Margins of pronotum, proepistemum, mesepistemum and lateral margins of abdominal
sterna 1-6 with appressed setae. Elytra with a few erect setae near shoulder.

Other external features. Tooth of mentum long. Labrum elongate, convex, slightly tridentate; four
submarginal setae (Fig. 56). Pronotum narrowed at posterior end, sutures shallow (Fig. 61). Punctures of
proepistemum deep and evenly distributed. Coupling sulcus of mesepistemum of female a broad and
moderately deep groove. Apex of front trochanters each with one seta. Elytra with apical end tapered and
slightly recurved to a small spine (Fig. 66). Pattern of elytral maculations with humeral spot on shoulder,
middle spot on disc, and marginal spot near the apex (Fig. 66). A small portion of the marginal band may or
may not be present. Punctures of elytra large gradually becoming deeper near shoulder. Elytra with a few
erect setae near shoulder.

Female genitalia. Sternum 8 with deep and narrow emargination in apical end, apices each acutely
rounded, and with 5 thick setae (Fig. 71a). Second gonapophyses elongate, narrow, median portion almost as
long as lateral portion (Fig. 71a). Syntergum 9 and 10 as in Figure 71b. Ventral sclerite with wide apical end
spatulate and deflected dorsally (Fig. 71c); two elongate posterior projections (Fig. 71c). Oviduct sclerite
short (Fig. 71c). Spermatheca and ducts ca 1.0 mm long.

Male genitalia. Apical end of median lobe slender, apex not hooked (Figs. 76a, b). Internal sac without
flagellum. Stylet unique bifid, joined to base of tooth, elongate. Tooth spiny field with long stylet-like piece.
Arciform piece very wide; central plate large; right bar present (Figs. 76c, d).

Geographical Variation. — Patterns of geographical variation in examined adult
characteristics were not apparent. Some adults had a marginal spot on the elytra.

Relationships. — Cicindela anulipes is sister to the lineage that gave rise to sister
species C. chlorosticta and C. staudingeria, and sister species C. nigroreticulata and
C. paranigroreticulata.

Habitat and Period of Activity. — Unknown.

Geographical Distribution, Localities, Examined Specimens. — Goias, Matto Grosso,
Minas Gerais, and Rio Grande do Sul (Fig. 77).

Brazil. Goias: Chapada, 2M, 5F, ICCM; Jatai, IF, IOC, 3M, 1 IF, MZSP; Jolaby*, IF, MZSP. Matto
Grosso: Utiariti, 9M, 10F, MZSP. Minas Gerais: no locality, 1M, IPZE, IF, MZSP. Rio Grande do Sul:
Tramandai, 3M, 2F, MNRJ.

Subgenus Plectographa Rivalier

Subgenus Plectographa Rivalier, 1954: 265 (TYPE SPECIES, Cicindela gormazi Reed, by original
designation).

Recognition. — Adults of this subgenus are distinguished by the four characters
given in couplet 5 of the key. In addition the labrum is basically unidentate with
4-1 1 submarginal setae; apical end of the median lobe is tapered and not hooked; in

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290

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Figs. 78-83. Labrum, dorsal aspect. 78, Cicindela suturalis Fabricius: (a) female, Maturaca, Am.; (b)
female, (c) male, Russas, Ce.; (d) female, (e) male, Sao Luis; Mar.; 79, C. nivea Kirby: (a) female, (b) male,
Florianopolis, Sta. Cat.; (c) female, Vacaria, M. Grosso; 80, C. melaleuca Dejean: (a) female, Rio Grande do
Sul; (b) female, Buenos Ayres, B.A.; 81, C. patagonica Brulle: (a) C. p. cherubim Chevrolat, female,
Montevideo, Uruguay; (b) C. p. bergiana Horn, male, Rio Grande do Sul; 82, C. nigrovittata Horn, female,
Minas Gerais; 83, C. apiata Dejean: C. a. apiata, (a) female, (b) male, Rio de Janiero, R.d.J.; C. a. claussenii
Putzeys, (c) female, (d) male, Rio Grande do Sul. Figs. 84-89. Pronotum, dorsal aspect. 84, Cicindela
suturalis Fabricius: (a) female, Maturaca, Am.; (b) female, (c) male, Russas, Ce.; (d) female, (e) male, Sao
Luis, Mar.; 85, C. nivea Kirby: (a) female, (b) male, Florianopolis, Sta. Cat.; (c) female, Vacaria, M. Grosso;
86, C. melaleuca Dejean: (a) female, Rio Grande do Sul; (b) female, Buenos Ayres, B.A.; 87, C. patagonica
Brulle: (a) C. p. cherubim Chevrolat, female, Montevideo, Uruguay; (b) C. p. bergiana Horn, male, Rio
Grande do Sul; 88, C. nigrovittata Horn, female, Minas Gerais; 89, C. apiata Dejean: C. a. apiata (a)
female, (b) male, Rio de Janeiro, R.d.J.; C. a. claussenii Putzeys, (c) female, (d) male, Rio Grande do Sul.

Brazilian Species of Cicindela

291

Figs. 90-95. Elytron, dorsal aspect. 90, Cicindela suturalis Fabricius: (a) female, Maturaca, Am.; (b) female,
(c) male, Russas, Ce.; (d) female, (e) male, Sao Luis, Mar.; 91, C. nivea Kirby: (a) male, Florianopolis, Sta.
Cat.; (b) female, Vacaria, M. Grosso; 92, C. melaleuca Dejean, female, Rio Grande do Sul; 93, C.
patagonica Brulle: (a) C. p. cherubim Chevrolat, female, Montevideo, Uruguay; (b) C. p. bergiana Horn,
male, Rio Grande do Sul; 94, C. nigrovittata Horn, female, Minas Gerais; 95, C. apiata Dejean: C. a. apiata
(a) female, (b) male, Rio de Janeiro, R.d.J.; C. a. claussenii Putzeys, (c) female, (d) male, Rio Grande do
Sul.

Quaest. Ent., 1989, 25 (3)

Figs. 96-100. Female genitalia: (a) sternum 8, second gonocoxa, second gonapophyses, ventral aspect; (b)
syntergum, 9 & 10, dorsal aspect; (c) bursa copulatrix, oviduct sclerite, oviduct, spermatheca and duct,
ventral sclerite, ventral aspect. Cicindela suturalis Fabricius: 96, Maturaca, Am.; 97, Russas, Ce.; 98, Sao
Luis, Mar.; C. nivea Kirby: 99, Florianopolis, Sta. Cat.; 100, Vacaria, M. Grosso.

Figs. 101-106. Female genitalia: (a) sternum 8, second gonocoxa, second gonapophyses, ventral aspect; (b)
syntergum, 9 & 10, dorsal aspect; (c) bursa copulatrix, oviduct sclerite, oviduct, spermatheca and duct,
ventral sclerite, ventral aspect. C. melaleuca Dejean: 101, Rio Grande do Sul; 102, Buenos Ayres, B.A.; 103,
C. patagonica cherubim Chevrolat, Montevideo, Uruguay; 104, C. nigrovittata Horn, Minas Gerais; 105, C.
apiata apiata Dejean, Rio de Janeiro, R.d.J.; 106, C. a. claussenii Putzeys, Rio Grande do Sul.

Quaest. Ent., 1989, 25 (3)

294 Freitag and Bames

Figs. 107-111. Male genitalia: median lobe (a) right lateral aspect, (b) dorsal aspect; (c) internal sac,
arciform piece, central plate (cp), flagellum (/?), right bar, shield ( sh ), spatulate field ( sf ), small stiffening rib
(sr), stylet, tooth, right lateral aspect; (d) left lateral aspect. Cicindela suturalis Fabricius: 107, Russas, Ce.;
108, Sao Luis, Mar.; 109, C. nigrovittata Horn, Minas Gerais; 1 , C. apiata apiata Dejean, Rio de Janiero,

R.d.J.; 1 1 1, C. a. claussenii Putzeys, Rio Grande do Sul.

Brazilian Species of Cicindela

295

Fig. 112. Map showing the geographical distribution of the species Cicindela suturalis Fabricius (•), C.
nivea nivea Kirby (■), and C. n. orbignyi Guerin-Meneville (A). Open circles represent state records, and
those with an adjacent point indicate records taken from the literature for C. suturalis Fabricius.

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296

Freitag and Bames

Fig. 1 13. Map showing the geographical distribution of the species Cicindela melaleuca Dejean (•), and C
patagonica Brulle (■).

Brazilian Species of Cicindela

297

Fig. 1 14. Map showing the geographical distribution of the species Cicindela nigrovittata Horn (♦), C.
apiata apiata Dejean (•), and C. a. claussenii Putzeys (■). Open circles are state or country records for C.
apiata apiata Dejean taken from the literature.

Quaest. Ent., 1989, 25 (3)

298

Freitag and Barnes

females the ventral sclerite of the bursa copulatrix is glabrous and has two elongate
posterior projections; the spermatheca and duct together are approximately 2.5 - 4.5
mm long.

Species groups. — This subgenus has five species groups that include 18 species.
Four groups, suturalis group, melaleuca group, nigrovittata group, and apiata
group, are found in Brazil.

Geographical distribution. — The geographical range of this subgenus extends
from temperate Chile and Patagonia northward to the West Indies, and is centered in
northern Argentina.

Phylogenetic relationships. — Plectographa and Cylindera are sister subgenera.

The suturalis group

Adults of this group are characterized by bright coppery and some green body
colouration. The group has seven species, two of which, C. suturalis and C. nivea
are found in Brazil. The geographical range of the suturalis group extends from
central Chile and Patagonia northward to the West Indies, and is centered in
northern Argentina. The suturalis and melaleuca groups are sister groups.

Cicindela ( Plectographa ) suturalis Fabricius
Figs. 78, 84, 90, 96, 97, 98, 107, 108, 1 12

Cicindela suturalis Fabricius, 1798: 62 (TYPE LOCALITY, in America St. Thomas); 1801: 242-Herbst,
1806: 207. -Dejean, 1825: 129. -Horn, 1915: 403; 1926a: 305; 1938: 52,-Leng and Mutchler, 1916:
693 -Blackwelder, 1944: 20.-Jonge Poerink, 1953: 133. -Rivalier, 1954: 266-Balazuc and Chalumeau,
1978: 22-25 -Ivie, 1983: 195.

Cicindela trifasciata Dejean, 1825: 85(not Fabricius) (TYPE LOCALITY, Cayenne).-Hom, 1915: 403;
1926a: 306-Blackwelder, 1944: 20.

Cicindela hebraea Klug, 1834: 20 (TYPE LOCALITY, Cayenne).-Putzeys, 1874: 117,-Hom, 1915: 403;
1926a: 306; 1938: 52. -Leng and Mutchler, 1916: 694,-Varas Arangua, 1930: 154.-Blackwelder, 1944:
20.-Jonge Poerink, 1953: 135. -Rivalier, 1954: 266-Mandl, 1963: 578-Balazuc and Chalumeau, 1978:
25.

Cicindela hieroglyphica Klug, 1834: 30 (TYPE LOCALITY, Para in Brasilien).-Hom, 1915: 403; 1926a:
306,-Blackwelder, 1944: 20.

Cicindela chlorocephala Mannerheim, 1837: 17 (TYPE LOCALITY, in insula Porto Rico).-Hom, 1915:
403; 1926a: 306,-Blackwelder, 1944: 20.

Cicindela insidiosa Gistl, 1837: 48 (TYPE LOCALITY, in America septentrionali, in Antillis).-Hom, 1915:
403; 1926a: 306.

Cicindela nocturna Steinheil, 1875: 96 (TYPE LOCALITY, Magdalenae fluminis, Colombia).-Hom, 1915:
403; 1926a: 306; 1938: 52 - Leng and Mutchler, 1916: 694. -Blackwelder, 1944: 20.-Jonge Poerink,
1953: 137-138.-Rivalier, 1954: 266-Mandl, 1958: 26.-Balazuc and Chalumeau, 1978: 26.

Cicindela hebraea guadeloupensis Fleutiaux and Salle, 1889: 358 (TYPE LOCALITY, Guadeloupe).-Hom,
1915: 403; 1926a: 306-Leng and Mutchler, 1916: 694. -Blackwelder, 1944: 20.-Jonge Poerink, 1953:
137-Balazuc and Chalumeau, 1978: 25.

Cicindela suturalis helvaea Horn, 1903: 333 (not Klug) (TYPE LOCALITY, Ciudad Bolivar, Venezuela).

Cicindela trifasciata boliviano Mandl, 1956: 386 (TYPE LOCALITY, Santa Cruz); 1958; 26; 1964: 16.
NEW SYNONYMY.

Brazilian Species of Cicindela

299

Recognition. — Narrow to broad distinctive elytral maculations (Figs. 90a-e) and
a unidentate labrum bearing 8-10 submarginal setae (Figs. 78a-e) distinguish
specimens of this species.

Synonyms and Types.— Names were based upon comparison of original
descriptions with specimens on loan.

Description. —

Body length. 6. 5-8.0 mm M, 6. 5-9.0 mm F.

Body colour. Head, pronotum and elytra slightly glossy with coppery, green, and less frequently blue,
reflections. Pleuron and venter mainly coppery, and green.

Body setae. Dense appressed setae on lateral portions of pronotum, pleuron, ventral 0.33 of
mesepistemum, and abdominal sterna 1-6.

Other external features. Labrum unidentate with eight to 10 (more frequently eight) submarginal setae
(Figs. 78a-e). Pronotum with parallel sides, sutures moderately deep (Figs. 84a-e). Coupling sulcus of
mesepistemum of female a deep sinuate groove. Apex of front trochanters with a single sensory seta. Elytra
with long apical spine, (longer in C. hebraea form specimens); maculations complete and joined together,
broadened in C. suturalis fomi specimens); punctations generally deep, subsutural row of large punctures
absent (Figs. 90a-e); microsculpture isodiametric; apical serrulations present.

Female genitalia. Sternum 8 with deep V-shaped apical emargination, apices each with three (less
frequently two) medium length setae (Figs. 96a-98a); second gonocoxa with a few long and short setae along
medial margin (Figs. 96a-98a); second gonapophyses as in Figures 96a-98a; syntergum 9 and 10 as in
Figures 96b-98b; bursa with right lateral extension; ventral sclerite lightly sclerotized with posterior lateral
extensions, median ridge absent (Figs. 96c-98c) oviduct sclerite with well developed lateral extensions (Figs.
96c-98c); length of spermatheca and duct 3. 00-3. 25mm.

Male genitalia. Median lobe with short blunt apex (Figs. 107a, b-108a, b). Internal sac with flagellum
1.5 loops at base. Stylet present next to tooth, the large spatulate apex of tooth characteristic of Plectographa
forms not elongate. Arciform piece, right bar, central plate, shield and small stiffening rib all well developed
(Figs. 107c, d- 108c, d).

Geographical Variation and Subspecies. — Populations composed of specimens
with narrow to broad but well defined elytral maculations are widespread in Brazil.
At least one identified specimen with continuous pale maculations (C. suturalis
form) has been collected near Manaus. In most specimens the maculations are broad
(C. hebraea form). They may be continuous in part (Figs. 90d, e) or the lunules may
be clearly discernible (Figs. 90a-c). Specimens with narrow elytral maculations (C.
nocturna, C. guadeloupensis or C. boliviana forms) are less frequent. The form of
the elytral maculations seems to be variable in local populations, and geographical
patterns are not evident. In addition the female genitalia tend to be uniform within
and among populations (Figs. 96-98).

Morphologically distinct geographical populations, in Brazil at least, apparently
do not occur, and consequently we have not applied subspecific names. A varied
elytral pattern and other discordant characteristics predominate in populations of C.
suturalis in South America north of Brazil and the West Indies. A thorough revision
is required as regional studies of this species (complex) have not led to taxonomic
stability (Leng and Mutchler, 1916: 693-695; Jonge Poerink, 1953: 133-138; Mandl,
1958: 24-28; Balazuc and Chalumeau, 1978: 23).

Relationships. — Cicindela suturalis and C. sinuosa Brulle (n. Argentina,
Paraguay and Uruguay) are sister species.

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300

Freitag and Barnes

Habitat and Period of Activity. — Specimens have been collected on sea beaches
and on extensive sand beaches along rivers (Pearson, 1984) during February, May,
June, September and December.

Geographical Distribution, Localities, Examined Specimens. — Islands of the
Caribbean from Jamaica and Hispaniola south to Trinidad, northern half of South

America south to southeastern Brazil (Fig. 1 12).

Bolivia. Cochabamba, IF, MZSP.

Brazil. Amazonas'. Arima (near Rio Purus), IF, ICCM, Barcelos, 1M, MZSP; Manaus, 1M, INPA, 1M,
MZSP; Manaus (Uha de Marchantaria), 2M, IF, INPA; Maturaca (Alto Rio Cauaburi), IF, MZSP; Nova
Olinda, 7F, ICCM; Tapuruquara, IF, MZSP. Bahia : Canavieiras, 1M, IOC; Encruzilhada (980m), 1M,
RRMC; Joazeiro, IF, ICCM; Rio Sao Francisco*, 2F, IOC; Villa Nova*, 2M, MZSP. Ceara: Aracaty, IF,
MZSP; Barreiras*, 1M, MZSP; Ceara Beach *, 5M, 2F, ICCM; Russas, 1M, IF, MZSP. Espirito Santo: IF,
MZSP. Maranhao: no locality, 2M, 4F, ICCM; Sao Luis, 4M, 6F, MZSP. Matto Grosso: Pimentel*, 1M, 2F,
MZSP; S. Domingos, 1M, MZSP; Sinop*, IF, RRMC. Para: Belem, 1M, MZSP; Cachimbo, !, IOC;
Morroda*, IF, ICCM; no locality, IF, ICCM, Santarem, 2M, 2F, ICCM. Paratba: Coremas, 1M, 2F, MZSP.
Santa Catarina: Imbituba, 1M, MNRJ.

Peru. Chancayo*, 2M, MNRJ.

Cicindela (Plectographa) nivea Kirby
Figs. 79, 85,91, 100, 112

Cicindela nivea nivea Kirby, 1818: 376 (TYPE LOCALITY, in Brasilia).- Dejean, 1825: 128,-Hom, 1915:
409; 1926a: 311; 1938: 53. -Fernandez, 1936: 109-Blackwelder, 1944: 19.-Rivalier, 1954:
266,-Mandl, 1963: 578,-Vidal Sarmiento, 1966b: 41.-Sumlin, 1979: 115.

Cicindela conspersa Dejean, 1825: 127 (TYPE LOCALITY, Tile de Sainte-Catherine, au Bresil).-Brulle,
1837: 9. -Horn, 1915: 409; 1926a: 311; 1938: 53,-Barattini, 1929: 1221. -Fernandez, 1936:
109,-Blackwelder, 1944: 19.-Vidal Sarmiento, 1966b: 43.

Cicindela niver orbignyi Guerin-Meneville, 1839: 296 (TYPES, a male and female in the MNHP general
collection bearing the following labels: “intricata” (green label, on male pin only)/Museum Paris
Patagonie (Patagones) d’Orbigny 1834/ 34 6073” (on a folded round label green on one side)/”TYPE”
(black letters on red label); TYPE LOCALITY, la Patagonie). -Horn, 1915: 409; 1926a: 311; 1938:
53.-Femandez, 1936: 109-Blackwelder, 1944: 19. Rivalier, 1954: 266-Vidal Sarmiento, 1966b:
43,-Sumlin, 1979:115.

Cicindela intricata Brulle, 1837: 7 (TYPE LOCALITY, File aux Cachons, dans la baie de San-Blas
(Patagonie), au bord de la mer). -Horn 1895a: 173; 1926a: 3 1 1 -Blackwelder, 1944: 19-Sumlin, 1979:
115.

Recognition. — The character state combination of large body size 9.0-13.5 mm),
elytra completely pale or with almost effaced maculations (Figs. 91a, b), dense
appressed white setae covering most of the body, and unidentate labrum
distinguishes adults of C. nivea.

Description. — Body length. Approximately 9.0-13.5 mm M and F.

Body colour. Head and pleuron slightly glossy, coppery and coppery green reflections. Elytra entirely
pale or with scattered coppery-green patches. Venter black with coppery or green reflections.

Body setae. Dense white appressed setae on head, frons, clypeus, genae, dorsal side or antennal articles
1,3,4 pronotum (mainly front and lateral portions), pleuron, prostemum, lateral portions of metastemum and
metacoxae, legs (sparser on tarsus), and abdominal sterna 1-6 (males), 1-5 (females).

Other external features. Tooth of mentum long. Labrum unidentate with four to 1 1 (most commonly
eight) submarginal setae (Figs. 79a-c). Pronotum broad, hind angles protruding posteriorly, sutures deep
(Figs. 85a-c). Coupling sulcus of mesepistemum of female a deep groove. Apex of front trochanters with
one sensory seta, middle trochanters glabrous. Elytra with apices slightly recurved, apical spine obsolete;
maculations effaced, ground colour completely absent or diffuse; punctation generally small, few large green

Brazilian Species of Cicindela

301

or coppery punctures near shoulder and median suture (Figs. 91a, b); microsculpture irregularly
isodiametric; apical serrulations small.

Female genitalia. Sternum 8 with shallow V-shaped apical emargination; apices rounded, each with five
to seven long thick setae (Figs. 99a, 100a); second gonocoxa with several long and short setae along medial
margin (Figs. 99a, 100a); second gonapophyses elongate as in Figures 99a, 100a; syntergum 9 and 10
broadened apically (Figs. 99b, 100b); ventral sclerite lightly sclerotized with posterior lateral projections,
median ridge absent (Figs. 99c, 100c); oviduct sclerite wide, with little or no anterior emargination (Figs.
99c, 100c); spermatheca and duct fused to bursa at base, spermatheca and duct ca 3. 5-4.0 mm (Figs. 99c,
100c).

Male genitalia. Male genitalia have been described by Vidal Sarmiento (1966b: 41).

Geographical Variation and Subspecies. — Most adults in northern localities of
Brazil have entirely pale elytra; those in southern localities, in Argentina, have
elytra with pigmentation forming a diffuse pattern of elytral maculations.
Intermediate populations consist of a majority of adults with varying degrees of
pigmentation in the elytra. This characteristic seems to be a reasonable basis for the
recognition of two subspecies because a north-south irregular cline is evident. The
names C. nivea nivea and C. nivea orbignyi Guerin-Meneville are retained for
northern (Brazil, Uruguay) and southern (Argentina) populations respectively (Fig.
113). The name C. conspersa, which was assigned to intermediate forms is not
recognized.

Relationships. — Cicindela nivea and C. hirsutifrons (n. Argentina) are sister
species.

Habitat and Period of Activity. — The activity period is from September to June,
though most adults examined were collected in October, November, December,
January, and February. Populations are probably found close to sea beaches as
indicated by the distribution of the species (Fig. 1 12).

Geographical Distribution, Localities, Examined Specimens. — East coast of

South America from Espirito Santo, Brazil south to Patagonia (Fig. 112).

Cicindela nivea nivea Kirby, Brazil. Espirito Santo : Gurapari, 13M, 6F, MZSP. Rio de Janeiro :
Atafona, 1M, IF, MNRJ; Barra de Marua, Praia de Itapuacu*, 3M, IF, MZSP; Cidade, 2M, MZSP;
Copacabana, IF, IOC, 2M, 2F, MZSP; Itacuruga*, 1M, IOC; Leblon, 1M, 5F, MNRJ; Niteroi, 3M, IF, IOC,
IF, MZSP, 2M, 2F, MNRJ; no locality, 12M, 6F, ICCM; Praia do Sai\ 1M, IF, MNRJ; Recreio dos
Bandeirantes*, 1M, 3F, MNRJ; Rio de Janeiro, 1M, 3F, MZSP, IF, MNRJ; Tijuca, 7M, 4F, MZSP. Rio
Grande do Sul: Torres, 3M, IF, MZSP. Santa Catarina : Florianopolis, 9M, 3F, MZSP; Itaiaby*, IF, MZSP;
no locality, IF, IOC, 4M, MZSP. Sao Paulo : Campos do Jordao, 1M, 3F, MZSP; Iguape, IF, MZSP;
Itanhaem, 1M, 5F, MZSP; Peruibe, 1M, IF, MZSP; Santos, 3M, 6F, IOC, 2F, MZSP, 1M, IF, MNRJ; Sao
Vincente, 1M, 3F, MZSP.

Uruguay. Atlantida, 1M, MZSP; Montevideo, 2M, IF, MZSP.

Cicindela nivea orbignyi Guerin-Meneville. Argentina. Buenos Aires: Buenos Aires, 1M, 3F, IOC; La
Plata IF, MZSP.

The melaleuca group

Adults of this group are characterized by eight to fewer than eight submarginal
setae on labrum, and very large foveae on elytra. The group has seven species, two
of which are found in Brazil, C. melaleuca and C. patagonica. The geographical
range of the melaleuca group is centerd in northern Argentina and extends

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northward to southeastern Brazil, western Paraguay and eastern Bolivia, and
southward to temperate Patagonia and Chile. The melaleuca and suturalis groups
are a sister pair.

Cicindela (Plectographa) melaleuca Dejean
Figs. 80, 86, 92, 101, 102, 113

Cicindela melaleuca Dejean, 1831: 238 (TYPE LOCALITY, pres de Buenos-Ayres, sur les bords de la

rivieres de La Plata, Argentina). -Horn, 1915: 409; 1926a: 310; 1938: 53.- Barattini, 1929: 1219.-

Femandez, 1936: 107- Blackwelder, 1944: 18. -Rivalier, 1954: 266- Vidal Sarmiento, 1966b: 40;

1967: 213 - Sumlin, 1979: 110.

Recognition. — The character state combination of the very dark brown to black
body colour, broad pronotum, (Figs. 86a, b), setose vertex, frons, clypeus and genae,
and pattern of elytral maculations (Fig. 92) distinguishes specimens of C. melaleuca
from other Brazilian tiger beetles.

Description. —

Body length, ca 8.5-10.5 mm M, ca 9.0-1 1.0 mm F.

Body colour. Head and pronotum slightly glossy, elytra dull. Dorsum very dark brown or black. Pleuron
and sternum black with slight reflections of copper, blue, green, and purple. Abdominal sterna black with
slight metallic reflections from lateral margins.

Body setae. Head with vertex, frons, clypeus, and genae setose. Pleuron generally setose, mesepistemum
with a few setae near ventral end. Lateral portions of abdominal sterna 1-5 (females' nd 1-6 (males) with
appressed setae.

Other external features. Labrum elongate, unidentate, almost tridentate, with six 10 (most commonly
eight) submarginal setae (Figs. 80a, b). Pronotum very wide, especially in females, broadened in anterior
0.33 (Figs. 86a, b). Coupling sulcus of mesepistemum of female a straight, long, shallow, broad groove.
Apex of front trochanters with one seta. Elytra each with apex recurved to a small spine (Fig. 92). Pattern of
elytral maculations with connected humeral lunule and middle band, marginal lunule reduced, apical lunule
complete (Fig. 92). Punctation of elytra green, with a row of large foveae, green -blue with bright coppery
umbilicate centers, near median suture and a cluster near shoulder (Fig. 92).

Female genitalia. See also Vidal Sarmiento (1967: 209). Sternum 8 with very broad and shallow apical
emargination, apices each with five to nine stout setae (Figs. 101a, 102a). Second gonocoxa and second
gonapophyses as in Figures 101a, 102a. Syntergum 9 and 10 as in Figures 101b, 102b. Ventral sclerite with
two elongate posterior projections (Figs. 101c, 102c). Oviduct sclerite wide (Figs. 101c, 102c). Spermatheca
and duct at least 4.5 mm long (broken in dissection).

Male genitalia. Male genitalia have been described by Vidal Sarmiento (1966b: 40).

Geographical Variation. — Body colour, pattern of the elytral maculations and
genitalic characteristics vary considerably in Argentina, and distinct geographical
populations occur there. Cicindela melaleuca is widespread ranging from Patagonia
to Rio Grande do Sul. The C. reedi Horp form is a conspecific member or close
relative that occurs in Patagonia. Horn (1938: 53) is followed by Sumlin (1979: 111)
in treating C. reedi as a subspecies of C. melaleuca though Rivalier (1954: 266)
considers it to be a member of C. drakei Horn.

Relationships r — Cicindela melaleuca and C. patagonica, are sister species.

Habitat and Period of Activity. — Adults have been found beside salt marshes in
Rio Grande do Sul, at the mouth of the Rio de La Plata, and inland in dry areas
(Brulle, 1837: 8), in January and February.

Brazilian Species of Cicindela

303

Geographical Distribution, Localities, Examined Specimens. — Coastal southern

Brazil, Uruguay, and Argentina (Fig. 1 13).

Argentina. Buenos Aires: Buenos Aires, 1M, IF, MZSP.

Brazil. Rio Grande do Sul: salt marshes, 3F, ICCM.

Uruguay. Montevideo, 3M, 2F, ICCM, IF, MNRJ.

Cicindela (Plectographa) patagonica Brulle
Figs. 81,87, 93, 103, 113

Cicindela patagonica patagonica Brulle, 1837: 7 (TYPE, a male in the MNHP general collection bearing the
following labels: (green disc)/ “Museum Paris/Patagonie d’Orbigny/Patagonica/TYPE” (black letters on
a red label); TYPE LOCALITY, sur les bords du Rio Negro (Patagonie)). -Horn, 1915: 409; 1926a:
311; 1938: 53- Barattini, 1929: 1222- Fernandez, 1936: 108. -Black welder, 1944: 19- Rivalier, 1954:
266 - Vidal Sarmiento, 1966b: 43.- Sumlin, 1979: 114.

Cicindela patagonica cherubim Chevrolat, 1858: 315 (TYPE LOCALITY, des environs de

Montevideo).-Hom, 1915: 409; 1926a: 311; 1938: 53.- Barattini, 1929: 1222. -Fernandez, 1936: 108.-
Blackwelder, 1944: 19.- Rivalier, 1954: 266.- Vidal Sarmiento, 1966b: 43.- Sumlin, 1979: 114.
Cicindela patagonica bergiana Horn, 1985a: 174 (TYPE LOCALITY, Montevideo); 1915: 409; 1926a: 311;
1938: 53.- Fernandez, 1936: 108. -Blackwelder, 1944: 19.- Rivalier, 1954: 266.- Sumlin, 1979: 114.

Recognition. — The broadly fused pale maculations along lateral margin of the
elytron (Figs. 93a, b) combined with a setose clypeus, unidentate labrum bearing
seven to nine (eight basic) submarginal setae, large body size (approximately
9.0-10.0 mm), and restricted distribution in southern Rio Grande do Sul distinguish
specimens of C. patagonica from those of other Brazilian species.

Description. —

Body length. 9.0 mm M, 10.0 mm F.

Body colour. Dorsum ground colour brown. Head, pronotum and elytra with coppery and blue-green
reflections.

Body setae. Frons and clypeus setose. Pleuron setose. Proepistemum densely setose; setae on lateral
margin of abdominal stemites 1-5.

Other external features. Labrum unidentate with seven to nine (eight basic) submarginal setae (Figs.
81a, b). Pronotum broad with posteriorly protruding hind angles (Figs. 87a, b). Coupling sulcus of
mesepistemum of female a deep groove. Apex of front trochanters with a single seta. Elytra somewhat
flattened; apices recurved; apical spine small (Figs. 93a, b). Pattern of elytral maculations broad, continuous
along lateral margin, but all lunules present and distinct (Figs. 93a, b). Punctation sparse and shallow with
row of larger punctures along median suture.

Female genitalia. No females of C. p. patagonica or C. p. bergiana were available for study thus only
genitalia of C. p. cherubim were examined. Sternum 8 with V-shaped emargination, apices broadly rounded
each with seven or eight long thick setae (Fig. 103a). Second gonocoxa with long setae on apical end (Fig.
103a). Second gonapophyses with medial and lateral portions of equal length (Fig. 103a). Syntergum 9 and
10 as in Figure 103b. Ventral sclerite of bursa broad and lightly sclerotized, median ridge absent (Fig. 103c).
Oviduct sclerite with well developed lateral flanges (Fig. 103c). Length of spermatheca and duct
approximately 3.5 mm.

Male genitalia. Male genitalia of C. p. patagonica and C. p. cherubim have been described by Vidal
Sarmiento, 1966b.

Geographical Variation and Subspecies. — Cicindela patagonica bergiana Horn,
a brown form, has been recorded from northern Argentina, Uruguay and Rio Grande
do Sul, Brazil by Horn (1915: 409), but no details about the latter record were given.
Cicindela p. cherubim Chevrolat is a green to blue-green from similar to C. p.
bergiana in the very broad continuous elytral maculations and occupies Uruguay

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304

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and the region of northeastern Argentina near Buenos Aires. The nominate from, C.
p. patagonica, occurs southward to Patagonia.

Relationships. — Cicindela patagonica and C. melaleuca are sister species.

Habitat and Period of Activity. — Specimens have been collected in December
and February on the seacoast.

Geographical Distribution, Localities , and Examined Specimens. — Coastal
from Rio Grande do Sul to Patagonia (Fig. 113). The subspecies C. p. bergiana

occurs in northern most portions of the range.

Argentina. Buenos Aires: Buenos Aires, 3M, IOC. Santa Cruz: Santa Cruz, 2M, IOC.

Brazil. Rio Grande do Sul: no locality, 1M, BMNH.

Uruguay. Aramincla, IF, MNRJ; Cuchilla Alta, 1M, MNRJ; Montevideo, 2F, IOC, 3F, MZSP; Rocha,
IF, MNRJ.

The nigrovittata group

Adults of this group are characterized by deep rugosity in form of two swirls on
head between the eyes. The group has only one species, C. nigrovittata , which is
found in the Brazilian Highlands. Phylogenetic relationships of the nigrovittata
group are uncertain.

Cicindela (Plectographa) nigrovittata Horn
Figs. 82, 88,94, 104, 109, 114

Cicindela nigrovittata Horn, 1896c: 172 (SYNTYPES, two females and one male in the IPZE labelled:

“Nonfried Minas Geraes/Type ! Dr. W. Horn (1M IF), Type ! Coll. W. Horn (1F)/Syntypes” (black

letters on red label); TYPE LOCALITY, Minas Geraes); 1915: 405; 1926a: 307; 1938: 52.-

Blackwelder, 1944: 19.

Recognition. — The character state combination of the large body size, black
body colour, large deep punctures with sharp edges in the anterior 0.5 of the elytra
(Fig. 94) and pronotum (Fig. 88), and one or two small pale spots, and velvety
patterns on the elytra (Fig. 94) distinguish adults of C. nigrovittata from those of
other Brazilian species of Cicindela. The female genitalia are diagnostic (Figs.
104a-c).

Description. —

Body length, ca 7.5 mm M, ca 8.0-9. 0 mm F.

Body colour. Dorsum dull, black, matte and velvety patterns on the elytra. Venter slightly glossy, black
faint purple reflections from proepistemum and mesepistema. Legs black, with hint of coppery and green
reflections.

Body setae. Fine sparse white pubescence present on: vertex, frons, and genae of head; all margins and
beside longitudinal suture of pronotum, pleuron, and lateral portions of abdominal sterna 1-5. Few short fine
setae arise from deep punctures on basal end of elytra.

Other external features. Vertex of head deeply rugose. Two wide pits on vertex between hind margins
of eyes. Tooth of mentum elongate, well developed. Preapical articles of labial palpi broad. Labrum convex,
elongate, tridentate, with four to six submarginal setae (Fig. 82). Pronotum broadened anteriorly, sides not
rounded; large, deep punctures on lateral portions (Fig. 88). Coupling sulcus of mesepistemum of female
shallow, wide, virtually absent. Apex of front trochanters with one seta. Elytra broadened in apical 0.33;
apices slightly recurved to small spine (Fig. 94). Pattern of elytral maculations with one or two small pale

Brazilian Species of Cicindela

305

spots, and velvety and matte motif (Fig. 94). Punctures of elytra large, deep, with sharp edges (not raised) in
basal 0.33, shallow and indistinct in apical 0.66 (Fig. 94). Microsculpture of elytra basically isodiametric,
unclear in matte areas. Apices of elytra almost smooth, microserrulations tiny.

Female genitalia. Sternum 8 with deep moderately wide apical emargination, each apex with five stout
setae (Fig. 104a). Second gonocoxa with two or three setae on medial margin (Fig. 104a). Second
gonapophyses elongate (Fig. 104a). Syntergum 9 and 10 as in Figure 104b. Bursa copulatrix with pair of
lateral extensions (Fig. 104c). Ventral sclerite of bursa with two lateral posterior projections (Fig. 104c).
Oviduct sclerite small, shield-shaped (Fig. 104c). Spermatheca and duct at least 2.0 mm long (broken in
dissection).

Male genitalia. Median lobe stout, apex short slightly curved ventrally (Figs. 109a, b). Internal sac with
flagellum 1.5 loops at base. Tooth dark elongate, narrow, spatulate apex; stylet absent. Small stiffening rib
present; shield small. Arciform piece broad; central plate square-shaped at its base; right bar narrow (Figs.
109a-d).

Geographical Variation. — The pale spots on the elytra vary from one to two,
and the extent of the velvety pattern on the elytra differs in the few specimens
examined. A geographical pattern in these characteristics is not evident.

Relationships. — Cicindela nigrovittata is not closely related to any other species
of subgenus Plectographa.

Habitat and Period of Activity. — Unknown.

Geographical Distribution, Localities, Examined Specimens. — Inland (Fig.
114).

Brazil. Minas Gerais : Nonfried*, 1M, 2F (syntypes), IPZE; San Antonio Barranca*, IF, IPZE.

The apiata group

Adults of this group are characterized by stylet tooth long and pointed, and shield
reduced or absent from the internal sac of the male. The group has two species, of
which one, C. apiata, is found in Brazil. The geographical range of the apiata group
is centered in northern Argentina and extends northward to the Brazilian Highlands.
The apiata group is relatively primitive and not closely related to other groups in
subgenus Plectographa.

Cicindela ( Plectographa ) apiata Dejean
Figs. 83,89, 95, 105, 106, 110, 111, 114

Cicindela apiata apiata Dejean, 1825: 86 (TYPE LOCALITY, la partie meridionale du Bresil).- Bruch,
1907: 123.- Horn, 1915:408; 1926a: 310, 1938: 52. -Varas Arangua, 1925: 37.- Barattini, 1929: 1217.-
Femandez, 1936: 107- Blackwelder, 1944: 16. -Rivalier, 1954: 266 - Vidal Sarmiento, 1966a: 256,
259; 1966b: 39.- Sumlin, 1979: 114.

Cicindela apiata claussenii Putzeys, 1845: 365 (TYPE LOCALITY, la province des Mines au Bresil).-
Hom, 1915: 408; 1923: 111; 1926a: 310; 1938: 52. -Blackwelder, 1944: 16.- Rivalier, 1954: 266.-
Vidal Sarmiento, 1966b: 39.

Recognition. — Adults of C. apiata are set apart from those of other Brazilian
species by a combination of the broad elytral maculations lacking a marginal band
(Figs. 95a-d), narrow unidentate labrum with seven to 13 submarginal setae (Figs.
83a-d), and structure of female genitalia (Figs. 105, 106). The male genitalia are
also specific (Figs. 110, 111, and Vidal Sarmiento, 1966b: 33, 39).

Quaest. Ent., 1989, 25 (3)

306

Freitag and Bames

Synonyms. — Names were based upon comparison of specimens on loan with
original descriptions.

Description. —

Body length. 7. 0-9.5 mm M, 7. 5-9.5 mm F.

Body colour. Dorsum dull, dark brown. Head and pronotum greenish in some specimens. Pleuron and
lateral edges of abdomen mainly coppery. Venter black with blue-green reflections.

Body setae. Pleuron moderately setose. Mesepistemum sparsely setose near coxa. Lateral portions of
abdominal sterna 1-6 (males) and 1-6 or 1-5 (females) setose.

Other external features. Labrum unidentate, tooth prominent or obscurely tridentate, seven to 13
submarginal setae (Figs. 83a-d). Pronotum stout broader in anterior 0.5, sides slightly to strongly ampliate
(Figs. 89a-d). Coupling sulcus of mesepistemum of female a broad groove. Apex of front trochanters with
one seta. Profemora and mesofemora with very long fine setae. Apices of elytra not recurved, obliquely
truncated to tapered, apical spine small. Elytra with maculations broad, with or without ragged edges and
appearing as spots or not. Humeral lunule complete, oblique joined or not to complete middle band, apical
lunule complete and separate, marginal band absent (Figs. 95a-d). A few setae present or not on shoulder of
elytra. Punctation green, sparse with larger umbilicate punctures near shoulder and along median suture.
Microsculpture isodiametric bead-like.

Female genitalia. Sternum 8 with moderately deep V-shaped posterior emargination, apices each w:
three to five stout setae (Figs. 105a, 106a). Second gonocoxa with a few setae on medial margin (Figs. 105
106a). Second gonapophyses as in Figures 105a, 106a. Syntergum 9 and 10 elongate, broadened apically
(Figs. 105b, 106b). Ventral sclerite of bursa copulatrix with two posterior lateral extensions (Figs. 105c,
106c). Lateral flanges of oviduct sclerite well developed (Figs. 105c, 106c). Spermatheca and duct ca 2.5
mm long.

Male genitalia. Apex of median lobe with small projecf on (Figs. 110a, b, 111a, b). Internal sac with
flagellum 1.5 loops at base. Stylet and tooth both very long nd pointed. Right bar and small stiffening rib
present; arciform piece moderately wide (Figs. 1 1 lc, d and Vidal Sarmiento, 1966b: 33, 39).

Geographical Variation and Subspecies. — Adults of C. apiata are varied in
body size, in presence or absence of setae on the clypeus, in number of submarginal
setae on the labrum, in shape of pronotum, in the pattern of elytral maculations, in
presence or absence of setae on the base of elytra; and in presence or absence of
setae on sternum 6 of females. Although the specimens used in this study were too
few for statistical analysis, the sparse data assembled indicate geographical patterns
in the variation of these characteristics.

The body length of C. apiata varies from 7.0-7. 5 (male) and 7.5-8.0 mm (female)
in Rio de Janiero to 9.0-9.5 mm (male) d 9.5 mm (female) in Rio Grande do Sul.
In Brazil an uneven southward increa^ 1 adult body length is evident with an
abrupt change in Rio Grande do Sul. t ;ast-west pattern of variation in body
length is not apparent. Adults of both sexes om Argentina are the largest being ca
9.0-10.0 mm long.

Variation in the other observed characteristics roughly complements that of body
length. Most large specimens have a pronotum with ampliate sides, glabrous
clypeus, bear fewer submarginal setae on the labrum (7-8 for males, 8 for females in
Rio Grande do Sul; 10 for males, 9 for females in Rio de Janiero), lack setae on the
base of elytra, and they have expanded elytral maculations with ragged and spotted
edges particularly in the humeral lunule and marginal band which are connected or
almost connected. Females lack setae on the lateral margins of abdominal sternum 6.

In contrast, most small specimens have a less stout pronotum with only slightly
ampliate sides, bear setae on the clypeus, (usually 2, crossed), have a greater number

Brazilian Species of Cicindela

307

of submarginal setae on the labrum (eight or nine for males, 11-12 for females in
Rio de Janiero; 1 1 for males, 13 for females in Sao Paulo), bear setae on the base of
the elytra, and have narrower elytral maculations with even edges and generally
unspotted with unconnected humeral lunule and middle band. Females have a few
setae on the lateral margins of abdominal sternum 6.

Because there is a considerable amount of concordant variation among these
characteristics two subspecies are recognized, C. apiata apiata and C. apiata
claussenii. The former comprises relatively large adults, most of which have a
glabrous clypeus, labrum with seven to 10 (males) and eight or nine (females)
submarginal setae (Figs. 83a, b), broad pronotum with ampliate sides (Figs. 89a, b),
expanded, merged, ragged and spotted elytral maculations (Figs. 95a, b), and
glabrous base of elytra and lateral margin of abdominal sternum 6; the female
genitalia are as in Figures 105a-c. Adults of the latter subspecies are smaller, most
of which have one or two setae on the clypeus, labrum with eight to 1 1 (males) and
11-13 (females) submarginal setae (Figs. 83c, d), pronotum with sides not
noticeably ampliate (Figs. 89c, d), and elytral maculations relatively narrow, with an
even outline, isolated maculations, and few spots (Figs. 95c, d), and a few setae on
the base of elytra and lateral margin of abdominal sternum 6. Female genitalia are as
in Figures 106a-c.

The subspecies C. a. apiata ranges from the northern half of Argenti 1 and
Uruguay north to Rio Grande do Sul, Matto Grosso, Sao Paulo, Minas Gera, and
Rio de Janiero, while C. a. claussenii occurs in Matto Grosso, Sao Paulo, Minas
Gerais, and Rio de Janiero (Fig. 114). Adults with characteristics of both subspecies
are present in Rio de Janiero.

Relationships. — Cicindela apiata and C. eugeni Castelnau (n. Argentina) are
sister species.

Habitat and Period of Activity. — Adults have been collected from October to
May, beside water, at various elevations from sea level to 1700 m. Adults of C. a.
apiata have been found near salt marshes.

Geographical Distribution, Localities, Examined Specimens. — - Coastal and
inland, from the northern half of Argentina north to Rio Grande do Sul, Sao Paulo,
Matto Grosso, Minas Gerais, and Rio de Janiero (Fig. 1 14).

Cicindela apiata apiata Dejean. Brazil. Matto Grosso : Salobra, 1M, MZSP; Tres Lagoas, 5M, MZSP;
Vacaria, 1M, IF, MZSP. Minas Gerais'. Mar de Espanha*, 2F, IOC, 1M, IF, MZSP; Passa Quatro, 1M, 2F,
IOC. Rio de Janeiro: Campo Bello*, IF, MZSP; Itatiaia (700-11 10m), 2M, 7F, IOC; Rio de Janeiro, 1M, IF,
MZSP. Rio Grande do Sul: salt marshes, 4M, IF, ICCM. Sao Paulo: Campos do Jordao, 1M, 3F, MZSP;
Cantareira*, 1M, MZSP; Garca, 6M, IF, MZSP; Onda Verde*, 2F, MZSP; Pindamonhangaba, 1M, IF,
MZSP.

Cicindela apiata claussenii Putzeys. Brazil. Matto Grosso: Vacaria, 4M, 4F, MZSP. Minas Gerais:
Caraga*, 1M, IOC; Pocos de Caldas, 12 !, RRMC: Virginia (1500 m), 1M, MNRJ. Rio de Janeiro: Grajau,
1M, IF, IOC; Itatiaia, 1M, IF, MNRJ; Petropolis, 1M, MNRJ; no locality, 3M, 2F, ICCM. Sao Paulo: Alto
da Serra (mountains between Sao Paulo and Santos), 5M, 6F, MZSP; Serra do Bocaina*, 7M, 4F, MNRJ.

Quaest. Ent., 1989, 25 (3)

308

Freitag and Bames

Figs. 115-121. Labrum, dorsal aspect: 115, Cicindela kollari Gistl, Santa Teresina, M. Grosso: (a) female;
(b) male; 116, C. confluentesignata Horn: (a) female, Minas Gerais; (b) female, (c) male, Rivera, Uruguay;

1 17, C. morio Klug: (a) female, (b) male, Jatai, Go.; (c) female, (d) male, Dianopolis, Go.; (e) female, Para;

118, C. marquardti Horn, Tres Lagoas, M. Grosso: (a) female; (b) male; 119, C. piligera Horn: (a) male,
Minas Gerais; (b) female, Ecuador; 120, C. obsoletesignata Horn, male, Santa Catarina; 121, C.
friedenreichi Dokhtouroff, Rio Grande do Sul: (a) female; (b) male. Figs. 122-128. Pronotum, dorsal aspect.
122, Cicindela kollari Gistl, Santa Teresina, M. Grosso: (a) female; (b) male; 123, C. confluentesignata
Horn: (a) female, Minas Gerais; (b) female, (c) male, Rivera, Uruguay; 124, C. morio Klug: (a) female, (b)
male, Jatai, Go.; (c) female, (d) male, Dianopolis, Go.; (e) female, Para; 125, C. marquardti Horn, Tres
Lagoas, M. Grosso: (a) female; (b) male; 126, C. piligera Horn: (a) male, Minas Gerais; (b) female,
Ecuador; 127, C. obsoletesignata Horn, male, Santa Catarina; 128, C. friedenreichi Dokhtouroff, Rio Grande
do Sul: (a) female; (b) male.

Brazilian Species of Cicindela

309

Figs. 129-135. Elytron, dorsal aspect. 129, Cicindela kollari Gistl, Santa Teresina, M. Grosso: (a) female;
(b) male; 130, C. confluentesignata Horn: (a) female, Minas Gerais; (b) female, (c) male, Rivera, Uruguay;

131, C. mono Klug: (a) female, (b) male, Jatai, Go.; (c) female, (d) male, Dianopolis, Go.; (e) female, Para;

132, C. marquardti Horn, Tres Lagoas, M. Grosso: (a) female; (b) male; 133, C. piligera Horn: (a) male,
Minas Gerais; (b) female, Ecuador; 134, C. obsolete signata Horn, male, Santa Catarina; 135, C.
friedenreichi Dokhtouroff, Rio Grande do Sul: (a) female; (b) male.

Quaest. Ent., 1989, 25 (3)

Figs. 136-139. Female genitalia: (a) sternum 8, second gonocoxa, second gonapophyses, ventral aspect; (b)
syntergum, 9 & 10, dorsal aspect; (c) bursa copulatrix, oviduct sclerite, oviduct, spermatheca and duct,
ventral sclerite, ventral aspect. 136, Cicindela kollari Gistl, Santa Teresina, M. Grosso; 137, C.
confluentesignata Horn: Rivera, Uruguay; C. mono Klug: 138, Jatai, Go.; 139, Dianopolis, Go.;

Brazilian Species of Cicindela

311

141a

Figs. 140-143. Female genitalia: (a) sternum 8, second gonocoxa, second gonapophyses, ventral aspect; (b)
syntergum, 9 & 10, dorsal aspect; (c) bursa copulatrix, oviduct sclerite, oviduct, spermatheca and duct,
ventral sclerite, ventral aspect; (d) dorsal aspect. Cicindela morio Klug: 140, Para; C. marquardti Horn: 141,
Tres Lagoas, M. Grosso; 142, Matto Grosso or Sao Paulo (syntype); C. piligera Horn: 143, Ecuador.

Quaest. Ent., 1989, 25 (3)

Figs. 144—146. Female genitalia: (a) sternum 8, second gonocoxa, second gonapophyses, ventral aspect; (b)
syntergum, 9 & 10, dorsal aspect; (c) bursa copulatrix, oviduct sclerite, oviduct, spermatheca and duct,
ventral sclerite, ventral aspect; (d) dorsal aspect; (e) left lateral aspect; (f) right lateral aspect. 144, Cicindela
piligera Horn, Minas Gerais; 145, C. obsolete signata Horn, Santa Catarina; 146, C. friedenreichi
Dokhtouroff, Rio Grande do Sul.

Figs. 147-154. Male genitalia: (a) dorsal aspect (apex only in some specimens), (b) right lateral aspect, (c)
left lateral aspect; internal sac, arciform piece, central plate, flagellum, right bar, small stiffening rib, stylet,
tooth, (d) right lateral aspect, (e) left lateral aspect. 147, Cicindela kollari Gistl, Santa Teresina, M. Grosso;
C. morio Klug: 148, Jatai, Go.; 149, Dianopolis; C. marquardti Horn: 150, Tres Lagoas, M. Grosso; 151,
Matto Grosso or Sao Paulo (syntype); 152, C. piligera Horn, Minas Gerais; 153, C. obsoletesignata Horn,
Santa Catarina; 154, C.friedenreichi Dokhtouroff, Rio Grande do Sul.

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Freitag and Bames

Fig. 155. Map showing the geographical distribution of the species Cicindela kollari Gistl (■), and C.
confluentesignata Horn (•). Open squares and circle repre state records of C. kollari Gistl and C.
confluentesignata Horn respectively.

Brazilian Species of Cicindela

315

Fig. 156. Map showing the geographical distribution of the species Cicindela morio Klug (•), and C.
marquardti Horn (■). Open circle represents a state record for C. morio Klug.

Quaest. Ent., 1989, 25 (3)

316

Freitag and Bames

Fig. 157. Map showing the geographical distribution of the species of Cicindela friedenreichi Dokhtouroff
(A). Open circles and squares represent state or country records for C. piligera Horn and C. obsolete signata
Horn respectively.

Brazilian Species of Cicindela

317

Subgenus Cylindera Westwood

Genus Cylindera Westwood, 1831: 300 (TYPE SPECIES, Cicindela germanica Linnaeus, by original
designation).-Hom, 1915: 236, 245 and 291. -Rivalier, 1954: 264.- Reichardt, 1977: 374.

Genus Cicindosa Motschulsky, 1864: 173 (TYPE SPECIES, Cicindosa obliquealba Motschulsky, by
original designation). -Horn, 1915: 236, 405.- Reichardt, 1977: 374. NEW SYNONYMY.

Recognition. — Adults of this subgenus are distinguished by the five characters
given in couplet 5 of the key. In addition the labrum has 5-13 submarginal or
marginal setae; elytral maculations are broad in specimens of most species,
extensive in specimens of a few species, and absent in others; apex of the median
lobe of the male is not hooked; females of most species have a bursa copulatrix with
a well developed ventral sac, and ventral sclerite with two anterior lobes and
posterior projections covered with setae; the spermatheca and duct together are
approximately 1. 7-3.0 mm long.

Species Groups. — This subgenus has two South American species groups, mono
group and friedenreichi group, which together have nine species. Both groups are
found in Brazil.

Geographical Distribution. — The geographical range of this subgenus extends
from northern Argentina northward to the Amazon River.

Phylogenetic Relationships. — Cylindera and Plectographa are sister subgenera.

The morio group

Adults of this group are characterized by a broad pronotum and membranous
tooth in the internal sac of the male. The group has six species, four of which are
found in Brazil, C. kollari, C. confluentesignata, C. morio, and C. marquardti. The
geographical range of the morio group extends from eastern Brazil to eastern Peru
and Ecuador, and southern parts of the Amazon basin. The morio group and
friedenreichi group are a sister pair.

Cicindela ( Cylindera ) kollari Gistl
Figs. 115, 122, 129, 136, 147, 155

Cicindela hirticollis Dejean, 1831: 256 (not Say) (TYPE LOCALITY, au Bresil).- Gistl, 1837: 64.- Horn,
1915: 404; 1926a: 306.- Blackwelder, 1944: 18.

Cicindela kollari Gistl, 1837: 64 (TYPE LOCALITY, circa Para in Brasilia).- Horn, 1915: 404; 1926a: 306;

1938: 52. -Blackwelder, 1944: 18.- Rivalier, 1954: 265.

Cicindela scutellaris Gistl, 1837: 87 (not Say) (TYPE LOCALITY, in Brasilia).- Horn, 1915: 404, 1926a:
306.- Blackwelder, 1944: 18.

Cicindela dimidiaticornis Lucas, 1857: 30 (TYPE LOCALITY, le Bresil interieur).- Horn, 1915: 404;
1926a: 306. -Blackwelder, 1944: 18.

Recognition. — The character state combination of the dark head and pronotum
contrasting with the paler elytra, antenna with pale articles 5-11, unidentate short
labrum with 5 or 6 submarginal setae (Figs. 115a, b), single seta on the front and
middle trochanters, widely expanded elytral maculations (Figs. 129a, b), and large

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318

Freitag and Bames

and deep brown punctures of the elytra distinguishes adults of C. kollari from those
of other Brazilian species of Cicindela.

Adults of C. kollari are similar to those of its Peruvian sister species of C.
malaris Horn which is found in the area of the upper Amazon River system from
Pampa del Sacramento to Pebas. Adults of C. malaris have a distinct pattern of
elytral maculations with large patches of dark colour. Males have appressed white
setae on the margins of sterna 1-6. Adults of C. kollari have almost or completely
immaculate elytra, and males have margins of abdominal sterna 1 -5 setose.

Synonyms and Types. — The name C. dimidiaticornis was assigned to male of C.
kollari (see Geographical Variation and Sexual Dimorphism section).

Description. —

Body length, ca 6.8-7. 0 mm M, ca 6.5 mm F.

Body colour. Head green, coppery and purple (especially females). Pronotum coppery and green. Elytra,
mainly ivory, with brown punctation, dark brown patches with hint of blue-green reflections in females.
Venter dark brown coppery and green reflections.

Body setae. Males more densely setose than females. Head glabrous. Pronotum margins densely setose
disc sparsely setose. Pleuron densely setose, mesepistemum with a few setae near mesocoxa. Lateral
portions of abdominal sterna 1-5 in males, 1-6 in females, metastemum and metacoxa with dense appressed
setae.

Other external features. Antennal articles 5-11 pale. Labrum narrow (not elongate), unidentate, five or
six submarginal setae (Figs. 115a, b). Pronotum square, hind angles of females protruded (Figs. 122a, b).
Coupling sulcus of mesepistemum of female an elongate straight groove. Legs, especially the femur, more
densely setose in males. Apices of front and middle trochanters each with one seta. Apex of elytra recurved
to small spine in females, not recurved elongated to long spine in males (Figs. 129a, b). Dark portions of
colour pattern almost effaced or effaced in males (Fig. 129b), patches of dark ground colour indicate oblique
humeral lunule and middle band in females (Fig. 129a). Punctures of elytra large, deep, and brown, a few
large punctures near shoulder and median suture in basal 0.33. Microsculpture of elytra almost effaced.

Female genitalia. Sternum 8 with shallow, broad, V-shaped apical emargination, apices broadly rounded
each with three stout setae (Fig. 136a). Second gonocoxa with several setae on median margin (Figs. 136a).
Second gonapophyses as in Figure 136a. Syntergum 9 and 10 as in Figure 136b. Ventral sclerite lightly
sclerotized with lateral lobes extending posteriorly (Fig. 136c). Oviduct sclerite short and wide (Fig. 136c).
Spermatheca and duct ca 1 .7 mm long.

Male genitalia. Median lobe tapered apically (Figs. 147a, b). Internal sac with flagellum 1.5 loops at
base. Tooth mainly dark elongate membranous field with small sclerite at base. Right bar, stylet and small
stiffening rib well developed; shield membranous Arciform piece moderately wide; central plate lightly
sclerotized (Figs. 147c, d).

Geographical Distribution and Sexuc i morphism . — Insufficient material was

available to determine patterns of get raphical variation, however, within
populations sexual dimorphism occurs. Males have dense body pubescence,
non-protruding hind angles of pronotum, annost effaced or completely effaced
colour pattern of elytra, and elongate, not recurved, apices of elytra. Females have
moderately pubescent bodies, protruding hind angles of pronotum, elytra with
extensive patches of dark ground colour and recurved apices.

Relationships. — Cicindela kollari and C. malaris Horn (Peru) are tentative sister
species.

Habitat and Period of Activity. — Adults have been collected in August on the
Matto Grosso side of Rio Araguaia, Santa Isabel. Habitat unknown.

Brazilian Species of Cicindela

319

Geographical Distribution, Localities, Examined Specimens. — Gentral Brazil
(Fig. 155).

Brazil. Goias: no locality, 3!, MNHP. Matto Grosso: Santa Isabel, 1M, CASC; Santa Teresina, 1M, 2F,
CASC. Para: no locality, 1M, ICCM.

Cicindela (Cylindera) confluentesignata Horn
Figs. 116, 123, 130, 137, 155

Cicindela confluens Horn, 1893: 197 (not Bremi, Fowler, Kraatz); 1894: PI. 3, Fig. 6.- Blackwelder, 1944:
17.

Cicindela confluentesignata Horn, 1915: 407 (replacement name) (HOLOTYPE, a female in the IPZE
collection bearing the following labels: “Minas Geraes, Staudinger/Type ! Dr. W. Hom/Holotypus
(black letters on red \&be\)r Cicindela confluens Horn””; TYPE LOCALITY, Minas Geraes); 1926a:
309; 1938: 52.- Fernandez, 1936: 106.- Blackwelder, 1944: 17.- Rivalier, 1954: 265. -Vidal Sarmiento,
1966a: 259; 1966b: 33, 38 - Mandl, 1973: 296 ( Brasiliella ).- Sumlin, 1979: 105.

Recognition. — The character state combination of pattern of elytral maculations
(Figs. 130a-c) and the shape of the labrum with its nearly marginal five or six setae
(Figs. 116a-c) distinguishes adults of C. confluentesignata from other Brazilian
species of Cicindela.

Synonyms and Types. — Horn (1915: 407) designated the name C.
confluentesignata for the name C. confluens Horn preoccupied by C. confluens
Kraatz.

Description. —

Body length. 8.5 mm M, 8. 5-9.0 mm F.

Body colour. Dorsum slightly glossy; head and pronotum black or dark brown with coppery green
reflections; elytra coppery brown with coppery and green reflections and with a few large green punctures.
Venter and pleuron glossy, black with blue-green and coppery reflections.

Body setae. Genae with a few setae. Proepistemum moderately setose; mesepistemum with a few setae
near ventral end; metepistemum setose. Abdominal sterna 1-6 setose laterally.

Other external features. Antenna with articles 5- 1 1 pale. Labrum convex, basically unidentate but can
be up to seven dentate, with five or six marginal setae (Figs. 116a-c). Pronotum stout widest in front 0.5
(Figs. 123a-c). Coupling sulcus of mesepistemum of female a long and narrow groove. Apex of front
trochanters each with one seta. Elytra with apices recurved to a short spine; maculations broad and more or
less continuous; large and small punctures with green reflections (Figs. 130a-c).

Female genitalia. Sternum 8 with sharp V-shaped emargination in apical end; apices with four or five
stout setae of moderate length (Fig. 137a). Second gonocoxa with setae along median margin (Fig. 137a);
second gonapophyses narrow and elongate (Fig. 137a); syntergum 9 and 10 as in Figure 137b; ventral
sclerite lightly sclerotized with two lateral extensions directed posteriorly, and two anterior flaps covered
with brown setae (Fig. 137c); oviduct sclerite small, rectangular (Fig. 137c); spermathecal duct broken,
length unknown.

Male genitalia. The male genitalia have been described by Vidal Sarmiento (1966b: 33, 38).
Geographical Variation. — There is local and geographical variation in the
pattern of elytral maculations (Figs. 130a-c) and in the shape of the labrum (Figs.
1 16a-c). Distinct geographical patterns are unclear.

Relationships. — - Cicindela confluentesignata and C. granulipennis Bates
(Ecuador) are tentative sister species.

Habitat and Period of Activity. — Adults have been collected in November and
January. Habitat unknown. This species may be riparian.

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320

Freitag and Barnes

Geographical Distribution, Localities, Examined Specimens. — The species C.
confluentesignata ranges from Minas Gerais south to Uruguay, northern Argentina
and Paraguay (Fig. 155).

Brazil. Minas Gerais : no locality, IF (holotype), IPZE. Rio Grande do Sul : Pelotas, IF, MNRJ.

Uruguay. Durazno, 1M, MNRJ; Rivera, 1M, 3F, MNRJ; Tacuarembo, 1M, MNRJ.

Cicindela (Cylindera) morio Klug
Figs. 117, 124, 131, 138, 139, 140, 148, 149, 156

Cicindela denticulata Klug, 1834: 15 (TYPE LOCALITY, aus Brasilien).- Horn, 1892b: 213; 1915: 405;

1923: 112; 1926a: 307; 1938: 52. -Blackwelder, 1944: 18.- Rivalier, 1954: 265.

Cicindela morio Klug, 1834: 16 (TYPE LOCALITY, aus Brasilien).- Horn, 1892b: 213; 1915: 405; 1926a:
307; 1938: 52. -Blackwelder, 1944: 18.- Rivalier, 1954: 265.- Vidal Sarmiento, 1966a: 259; 1966b:
27.- Sumlin 1979: 108.

Cicindela ocskayi Gistl, 1837: 22 (TYPE LOCALITY, in Brasilia).- Horn, 1915: 405; 1926a: 307-
Blackwelder, 1944: 18.

Cicindela acompsa Chaudoir, 1852: 27 (TYPE LOCALITY, les bords du fleuve des Amazones); 1854: 122.

-Horn, 1915: 405; 1926a: 307; 1938: 52.- Blackwelder, 1944: 18.

Cicindela egena Chaudoir, 1854: 123 (TYPE LOCALITY, les contrees riveraines du fleuve des Amazones).

-Horn, 1915:405; 1926a: 307.- Blackwelder, 1944: 18.

Cicindosa obliquealba Motschulsky, 1864: 173 (TYPE LOCALITY, Des rives du fl. des Amazones).- Horn,
1915: 405; 1926a: 307.- Blackwelder, 1944: 18.

Cicindosa inaequalis Motschulsky, 1864: 174 (TYPE LOCALITY, De l’Amerique equatoriale).- Horn,
1915:405; 1926a: 307.- Blackwelder, 1944: 18.

Recognition. — The character state combination of the elongate, convex, and
7-12 dentate labrum (Figs. 1 17a-e), black body, pattern of elytral maculations (Figs.
131a-e), and structure of male and female genitalia distinguishes adults of C. morio
from those of other Brazilian species.

Synonyms and Types. — We have not seen the type of C. morio. The name is
based upon comparison of the original description with specimens on loan.

Description. —

Body length. 6.5-10.0 mm M and F.

Body colour. Dorsum dull, black, metallic reflections faint, matte patterns on elytra. Pleuron and venter
semi-glossy to glossy, black, metallic reflections more obvious.

Body setae. Pleuron setose (appressed mainly). Lateral portions of abdominal sterna one to six in males
and one to five in females with sparse to moderately dense appressed setae.

Other external features. Antennal articles 5-6 paler than 1-4 and 7-11. Labrum elongate, convex seven
to 12-dentate (basically nine); deep marginal notches each with one marginal seta (eight to 10 setae total)
(Figs. 117a-e). Pronotum widest in anterior 0.5, sutures shallow (Figs. 124a-e). Coupling sulcus of
mesepistemum of female a broad groove. Apex of front and middle trochanters with a single sensory seta.
Elytra with small apical spine. Pattern of elytral maculation varies from completely immaculate to extensive,
but with pale maculations lacking on the front part of the shoulder (Figs. 131a-e). Punctures of elytra
shallow, green.

Female genitalia. Sternum 8 with deep V-shaped apical emargination, each apex with two or three thick
setae. Median margin of second gonocoxa with row of setae (Figs. 138a, 139a, 140a). Syntergum 9 and 10 as
in Figures 138b, 139b, 140b. Ventral sclerite in one piece, or two parts, a central one with recurved apex
(anvil-shaped in lateral aspect), and left lateral flattened piece; setose (Figs. 138c, 139c, 140c). Oviduct
sclerite shield-shaped (Figs. 138c, 139c, 140c). Bursa with small ventral sac on right side (Figs. 138c, 139c,
140c). Spermatheca and duct ca 1.75 mm long.

Male genitalia. Median lobe stout with short tapered apex (Figs. 148a-c, 149). Internal sac with
flagellum 1.5 loops at base (Figs. 148d, e). Tooth elongate field. Arciform piece large. Stylet large (Figs.

Brazilian Species of Cicindela

321

148d, e). Right bar and small stiffening rib well developed 9Figs. 148d, e). Two or three apical brush-like
patches (Figs. 148d, e).

Geographical Variation. — A considerable amount of variation occurs in body
size, dentition and number of submarginal setae of the labrum, and pattern of the
elytral maculations, but geographical patterns are unclear. Variation in these
characteristics was considerable in the Santarem population sample. Adults of the C.
acompsa form which have extensive pale maculations on the elytra are not given
subspecific status as they are found with conspecific adults which have various
elytral patterns.

Relationships. — Cicindela morio and C. marquardti are sister species.

Habitat and Period of Activity. — Adults are active beside fresh water from
October to January. One specimen was collected in June.

Geographical Distribution, Localities, Examined Specimens. — Inland, from
Amazon River south to Matto Grosso, Brazil (Fig. 156).

Brazil. Amazonas. Manaus, 1M, INPA, 1M, IF, IOC; Manaus (1 km w. Taruma Falls), IF, ICCM;
Manaus (Bisego L.), 1M, MZSP; no locality, 3M, IF, INPA. Goias: Chapada, 5M, 5F, ICCM; Dianopolis,
8M, 10F, MZSP; Jatai, 1M, IF, BMNH, 2M, 2F, MZSP. Matto Grosso : Barra do Tapirapes, IF, CASC, 1M,
MZSP; Corumba, 1M, MZSP; Utiariti, 1M, MZSP. Para: Belem, 1M, CASC; no locality, IF, BMNH;
Santarem, 4M, 4F, ICCM. Sao Paulo: Avanhand (Garbe L.)*, IF, MZSP.

Cicindela (Cylindera) marquardti Horn
Figs. 118, 125, 132, 141, 142, 150, 151, 156

Cicindela marquardti Horn, 1906: 91 (SYNTYPES, 4 males and 2 females in the IPZE bearing the

following labels: “Staudinger Matto Grosso oder Sao Paulo/Type ! coll. W. Horn/ syntypus” (black

letters on red label); TYPE LOCALITY, Sao Paulo int. aut Matto Grosso); 1915: 28,29,404; 1924: 48;

1926b: 77; 1938: 52,-Blackwelder, 1944: 18.-Rivalier, 1954: 265.

Recognition. — Diagnostic characteristics which distinguish adults of C.
marquardti from those of other Brazilian species are the combination of the greenish
head and pronotum contrasting with dark brown to black dull elytra, complete and
broad apical and humeral lunules (Figs. 132a, b), sparse appressed setae on lateral
portions of abdominal sterna 3,4,5 and several lateral setae on 2 and 6, and female
and male genitalia as shown in Figures 141, 142,150,151.

Description. —

Body length. 7.5-8.0 mm M, ca 8.5 mm F.

Body colour. Head and pronotum are rugose with dark brown ground colour and bright green (mainly)
and coppery reflections. Elytra dull, velvety, dark brown with hint of purple. Pleuron and venter glossy,
testaceous to dark brown with faint metallic reflections. Proepistemum coppery or coppery-green.

Body setae. Proepistemum moderately setose. Dense appressed setae on ventral 0.33 of mesepistemum,
mesepimeron, metepimeron and lateral 0.33 of mesostemum. Sparse appressed setae on lateral portions of
abdominal sterna 3, 4, 5, and 2 and 6 with several setae only.

Other external features. Tooth of mentum short and sharply pointed. Labrum convex, elongate, five
dentate (three prominent anterior, two small lateral), and six marginal setae (four anterior, two lateral) (Figs.
118a, b). Pronotum short, convex widened at anterior end, sutures deep (Figs. 125a, b). Coupling sulcus of
mesepistemum of female a groove with deepened middle portion. Apex of front and middle trochanters with
one sensory seta. Elytra with apices recurved in females, apical spine small. Humeral and apical lunules
broad and complete, middle and marginal bands absent. Punctation shallow, obscure, green, broader
punctures near shoulder and median suture (Figs. 132a, b). A few fine setae near shoulder and median

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322

Freitag and Bames

suture.

Female genitalia. Sternum 8 with narrow, deep V-shaped emargination in the apical end, apices each
with two or three stout setae (Figs. 141a, 142a). Second gonocoxa with setae along medial margin (Figs.
141a, 142a). Second gonapophyses elongate with medial portion slightly shorter than the lateral portion
(Figs. 141a, 142a). Syntergum 9 and 10 as in Figures 141b, 142b. Ventral sclerite in two parts with setae on
lateral margins and anterior ends (Figs. 141c, 142c, d). Oviduct sclerite shield-shaped, or square (Figs. 141c,
142c). Bursa copulatrix with ventral sac on right (Figs. 141c, 142c, d). Spermatheca and duct approximately
2.0-3.0 mm.

Male genitalia. Median lobe stout with short tapered apex (Figs. 150a, c, 151a-c). Internal sac with
flagellum 1.5 loops at base. Tooth dark elongate field. Arciform piece large. Central plate lightly sclerotized.
Stylet large. Right bar and small stiffening rib well developed. Two or three apical finger-like brushes and
small dark sclerite in right-ventral side of sac (Figs. 151a, d, e).

Geographical Variation. — Geographical patterns of variation have not been
determined because few specimens of C. marquardti have been studied. The pattern
of elytral maculations and sclerites of the female genitalia show limited
intrapopulation variation.

Relationships. — Cicindela marquardti and C. morio are sister species.

Habitat and Period of Activity. — Adults have been collected beside rivers in
January.

Geographical Distribution, Localities, Examined Specimens. — The total range

of C. marquardti is restricted to a small area in Matto Grosso, Brazil (Fig. 156).

Brazil. Matto Grosso: no locality, 4M, 2F, IPZE; Tras Lagoas, 1M, 6F, MZSP; Vacaria, 6M, 4F, MZSP.

The friedenreichi group

Adults of this group are characterized by reduced and partially depressed pale
elytral maculations, and elongate apex of the median lobe. The group has three
species, all of which are found in Brazil, C. piligera, C. obsoletesignata and C.
friedenreichi. The geographical range of the friedenreichi group is centered in
southeastern Brazil and extends westward to Ecuador. The friedenreichi and morio
groups are sisters.

Cicindela (Cylindera) piligera Horn
Figs. 119, 126, 133, 143, 144, 152, 157

Cicindela piligera Horn, 1897b: 18 (TYPE LOCALITY, Minas Geraes); 1915: 406; 1926a: 307; 1938: 52.-

Blackwelder, 1944: 19- Rivalier, 1954: 265.

Recognition.— The character state combination of elongate labrum with seven
uneven marginal teeth, elytra with maculations reduced to three depressed spots and
large depression in basal 0.33, and a membranous sac on the ventral right side of the
bursa copulatrix, together are diagnostic of C. piligera.

Synonyms and Types. — An examination of the holotype (IPZE) served to
establish the name of this rare species.

Description. —

Body length. 7.5 mm M, 7. 0-7. 5 mm F.

Brazilian Species of Cicindela

323

Body colour. Dorsum slightly glossy, pleuron and venter glossy. Ground colour black; dorsum with
slight coppery (mainly) and green reflections; pleuron and venter with coppery, green, and blue reflections.

Body setae. Lateral margins of the elytra, pleuron including ventral 0.5 of proepistemum and ventral end
of mesepistemum, and lateral portions of metastemum, metacoxa, and abdominal sterna 1-6 (males) and 1-5
(females), setose.

Other external features. Labrum elongate seven dentate (not well defined in some specimens) with six
marginal setae (Figs. 1 19a, b). Pronotum widest in middle sutures moderately deep (Figs. 126a, b). Coupling
sulcus of mesepistemum of female a groove with central pit. Apex of front and middle trochanters with one
seta. Elytra with recurved apices, apical spine small. Maculations reduced to three depressed spots.
Punctures large green, row or large punctate depressions along median suture, large depression in basal 0.33
(Figs. 133a, b). Microsculpture isodiametric, bead-like.

Female genitalia. Sternum 8 with moderately deep and narrow apical emargination, apices narrow, each
with three long stout setae (Figs. 143a, 144a). Second gonocoxa with a few setae along medial margin (Figs.
143a, 144a). Second gonapophyses and syntergum 9 and 10 as in Figures 143a, b, 144a, b. Bursa with sac on
ventral right side (Figs. 143c, 144c). Ventral sclerite with posterior emargination and lateral projections,
setose apical extension on right dorsal side (Figs. 143c, d, 144c). Oviduct sclerite shield-like (Figs. 143c,
144c). Spermatheca duct fused to bursa at base, spermatheca duct at least 1.25 mm (broken in dissection).

Male genitalia. Cicindela piligera type; median lobe as in Figures 152a-c. Flagellum 1.5 loops at base
(Fig. 152e). Stylet, arciform piece, right bar, small stiffening rib and central plate all well developed (Figs.
152d, e).

Geographical Variation. — Insufficient material for analysis.

Relationships. — Cicindela piligera and C. obsoletesignata are sister species.
Habitat and Period of Activity. — Unknown.

Geographical Distribution, Localities, Examined Specimens. — Two adults have

been collected in Minas Gerais, Brazil and one in Ecuador (Fig. 157).

Brazil, no locality, IF, IPZE. Minas Gerais: no locality, 1M (holotype), IF, IPZE.

Ecuador. No locality, IF, BMNH.

Cicindela (Cylindera) obsoletesignata Horn
Figs. 120, 127, 134, 145, 153, 157

Cicindela obsoletesignata Horn, 1895b: 91 (TYPE LOCALITY, St. Catharina); 1896c: 169; 1915: 406;

1926a: 307; 1938: 52. -Blackwelder, 1944: 19.- Rivalier, 1954: 263.- Vidal Sarmiento, 1966b: 31.-

Sumlin 1979: 105.

Recognition. — The character state combination of the black body; unidentate
labrum, dark at the base, with six almost marginal setae (seven in a few specimens)
(Fig. 120); sparse and fine body setae; and depressed obsolete maculations of the
elytra (Fig. 134) distinguishes adults of C. obsoletesignata from those of other
Brazilian species of Cicindela.

Synonyms and Types. — The name C. obsoletesignata is based upon an
examination of the holotype and three females and one male determined by Horn.

Description. —

Body length. 6.0 mm M, 7.0-7. 5 mm F.

Body colour. Body with dull to slightly glossy dorsum and slightly glossy venter; black, with black to
rufopiceus venter of abdomen, genae with metallic reflections in some specimens, a hint of metallic
reflection from dorsum and venter.

Body setae. Fine sparse setae on pleuron and sterna 1-5 (female) and 1-6 (male) of abdomen.

Other external features. Labrum unidentate, with 6 almost marginal setae (Fig. 120). Pronotum slightly
wider anteriorly, sutures shallow (Fig. 127). Coupling sulcus of mesepistemum of female a distinct
dorso-ventral groove. Apex of front trochanters each with one seta, middle trochanters glabrous. Elyra with

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apices slightly recurved or not, apical spine tiny or obsolete; maculations depressed with effaced or obsolete
middle band and apical lunule; punctures shallow, green; row of large markedly depressed punctures near
median suture and shoulder (Fig. 134).

Female genitalia. Sternum 8 with wide and moderately deep apical emargination; apices each with three
short stout setae (Fig. 145a). Second gonocoxa with grooves in medial margin (Fig. 145a). Second
gonapophyses as in Figure 145a. Syntergum 9 and 10 rectangular (Fig. 145b), bursa with larged wrinkled
and folded membranous sac with darkened area (Figs. 145c-f). Ventral sclerite wide, with a posterior
emargination and lateral projections; right dorsal apical extension of ventral sclerite setose (Figs. 145c-f).
Oviduct sclerite shield-like (Fig. 145c), spermatheca and duct lost in dissection.

Male genitalia. Median lobe and internal sac C. obsoletesignata type (Figs. 153a-e); flagellum 1.5 loops
at base (Figs. 153d, e). Stylet prominent (Figs. 153d, e). Arciform piece, right bar, small stiffening rib, and
central plate present (Figs. 153d, e).

Geographical Variation. — An adequate number of specimens was not available
for the determination of patterns of geographic variation.

Relationships.— Cicindela obsoletesignata and C. piligera are sister species.

Habitat and Period of Activity. — Unknown.

Geographical Distribution, Localities, Examined Specimens. — Figure 157.

Argentina. Chaco: no locality, IF, IPZE.

Brazil. Santa Catarina: no locality, 1M, 3F (including type), IPZE.

Cicindela (Cylindera) friedenreichi Dokhtouroff
Figs. 121, 128, 135, 146, 154, 157

Cicindela friedenreichi Dokhtouroff, 1887: 154 (TYPE LOCALITY, Sta. Cathari - Horn, 1904: 87;

1915: 407; 1926a: 309; 1938: 52. -Blackwelder, 1944: 18.- Rivalier, 1954: 263.- 1 andl, 1963: 578.

Recognition. — The character state combination of the unidentate labrum with
seven or eight setae close to the anterior margin (Figs. 121a, b), dark body, and
elytra with a humeral spot, complete and broad apical lunule, elongate and
depressed middle band (Figs. 135a, b), and a row of very large depressed punctures
along the median suture distinguishes adults of C. friedenreichi.

Synonyms and Types. — The name is based upon comparison of the original
description with specimens on loan from R.R. Murray.

Description-

Body length. 7.5 mm M, 7. 5-8.0 mm F.

Body colour. Dark, almost black with coppery and green reflections, dorsum slightly glossy, venter
glossy.

Body setae. Appressed setae sparse to moderate in anterior 0.5 and lateral portions of pronotum, pleuron
(mesepistemum glabrous mainly), lateral portions of metastemum and metacoxa, and lateral portions of
abdominal sterna 1-6 (sternum 6 of female with a few setae).

Other external features. Labrum convex, unidentate with seven or eight setae very close to margin (Figs.
121a, b). Pronotum widest in anterior 0.5, sutures deep in front and hind angles (Figs. 128a, b). Coupling
sulcus of mesepistemum of female a long sinuate groove. Front trochanters with one sensory seta. Elytra
with apical end tapered, slightly recurved in females, to small apical spine, maculations variable as in
Figures 135a, b, with humeral spot and complete apical lunule, apical 0.5 of middle band elongate and
depressed, distinct row of large depressed blue-green punctures near median suture and shoulder, punctation
shallow and green, microsculpture isodiametric bead-like.

Female genitalia. Sternum 8 with deep V-shaped apical emargination, apices each with three stout setae
(Fig. 146a). Second gonocoxa with grooves on median margin, second gonapophyses as in Figure 146a.
Syntergum 9 and 10 as in Figure 146b. Ventral sclerite with a pair of setose anterior lobes, anvil-shaped
(Figs. 146a-e). Spermatheca and duct lost in dissection.

Brazilian Species of Cicindela

325

Table 1. Classification of characters used in the construction of a genealogy of the
Brazilian species of Cicindela

No. Character

Character State

Plesiomorphous Apomorphous

1 Body size

2 Head and pronotum:

colour

3 Head and pronotum:

rugosity

4 Rugosity between eyes

5 Proepistemum

punctuation

6 Elytra surface

7 Elytra: ground colour

8 Frons: setae

9 Clypeus: setae

10 Genae: setae

1 1 Proepistemum: setae

12 Mesepistemum: setae

Average (7.0-8.0 mm)

Dark brown to black
with some metallic
reflections

Shallow

Shallow, no swirls

Shallow, random

Dull

Dark brown to black
with faint metallic
reflections

Absent

Absent

Absent

Sparse

Sparse

Large (>8.0 mm) A
Small (<7.0 mm) A1
Bright coppery (with
green in some
specimens) B
Metallic green or blue B 1
Deep C

Two shallow pits
(swirls of rugosity) D
Deep D1

Deep, few, evenly
distributed E
Matte, velvety F
Slightly glossy F1
Glossy F2
Metallic reflections
(coppery and green),
brighter G
Faintly purple G1
Present H

Dense appressed H1
Present I

Dense appressed I1
Present J

Dense appressed J 1
Moderate K
Dense K1

Dense appressed K2
Absent K3
Moderate L
Dense L1

Dense appressed L2
Glabrous L3

(continued on next page)

Quaest. Ent., 1989, 25 (3)

326

Freitag and Bames

Table 1 (continued)

No. Character

Character State

Plesiomorphous Apomorphous

13 Middle of abdominal Absent

sterna: setae

14 Setose lateral portions of 1-6

abdominal sterna

15 Labrum: width Average

16 Labrum: margin Unidentate

17 Labrum: setae position Submarginal

18 Labrum: setae 8 (average)

19 Mentum tooth: length Average

20 Antenna segments: colour Normal

21 Pronotum: width Average

22 Coupling sulcus: shape Groove (di act)

23 Middle trochanter: setae Absent

24 Abdominal sternum 5 Absent

of $: unpigmented
bell-shaped spot

Present M

Few additional erect M1

1- 5 N

2- 6 N1

3- 6 sparse N2
1-3 N3
Narrow O
Elongate centre O1
Elongate O2
Edentate P
Tridentate P1

5 -Dentate P2
7-Dentate P3
>7-Dentate P4
Nearer to margin
(some or all setae) Q
Marginal Q1
<8 (average) R
>8 (average) R1
ShortS
Long S1
5-6 pale T
5-11 pale T1
Narrow U
Broad IF

Groove with deeper
centre V

Groove with central pit V1
Pit V2

Almost absent V3
Present W
Present X

(continued on next page)

Brazilian Species of Cicindela

327

Table 1 (continued)

No. Character

Character State

Plesiomorphous Apomorphous

25 Elytra: pale markings

26 Elytra: foveae

27 Abdominal sternum 8 ?:
apical emargination

30 Ventral sclerite: shape

31 Ventral sclerite: setae

32 Oviduct sclerite: form

Narrow to broad,
complete or almost
complete

Obvious near shoulder
and median suture

Broad, average depth

Broad, tapered
posteriorly

Absent

Shield-shaped

Reduced Y
Reduced and partially
depressed Y1
Effaced Y2
Broad and fused Y3
Broad, fused, ragged,
spotted Y4
Ragged Y5
Narrow, fused, partly
ragged and spotted Y6
Not obvious Z
Few Z1
Absent Z2
Very large Z3
Broad, deep a
Broad, shallow a1
Very broad, deep a2
Very broad, shallow a3
Narrow, deep a4
Short b
Long b1
4c
5 c1
>5 c2

With median ridge d
Two elongate posterior
projections d1
Two anterior lobes
and two posterior
projections d2
Present e

Rectangular, not laterally
broadened f
Narrow f1
Membranous f2

28 Abdominal sternum 8 $: Average length

stout apical setae size

29 Abdominal sternum 8 ?: 2-3

stout apical setae number

(continued on next page)

Quaestv Ent., 1989, 25 (3)

328

Freitag and Bames

Table 1 (continued)

No. Character

Character State

Plesiomorphous

Apomorphous

33

Spermatheca and duct:

1.0-1. 5 mm

1.5-2. 5 mm g

length

>2.5 mm g1

34

Bursa copulatrix: setae

Absent

Present h

35

Bursa copulatrix: shape

Normal

One ventral right out-
pocketing i

Two anterior lateral out-

pocketings i1

36

Median lobe: apex

Short

Short, ventral notch j
Hooked ventrally j 1
Elongate j2
Bent dorsally j3

37

Flagellum

Present

Absent k

Fewer than 1 1/2 coils

(loops) at base k1

38

Tooth: condition

Sclerotized

Membranous 1

39

Tooth: shape

Elongate

Elongate with spatulate
apex m
Short m1

Elongate arising from two

long thin roots of equal
length m2

40

Internal sac: two finger-

Absent

Present n

like darkened fields,
one spatulate-like

41

Arciform piece: shape

Narrow

Broad o

Very broad tapered

apically o1

42

Arciform piece: position

Oblique

Longitudinal p

43

Central plate

Not joined to shield

Joined to shield q
Absent q1

44

Shield

Present

Absent r

(continued on next page)

Brazilian Species of Cicindela

329

Table 1 (continued)

No. Character Character State

Plesiomorphous Apomorphous

45 Shield: shape

46 Small stiffening rib

47 Stylet

48 Stylet: form

49 Sagittal sclerotized plates

50 Cuna (triangular piece)

Single apex

Present

Present

Average length to long

Absent

Absent

Single apex tapered s
Single apex rounded s1
Single apex rounded,
bent s2

Single apex tridentate s3
Two apices s4
Two apices short, blunt, s5
Heart-shaped two apical
lobes s6

Two apices elongate s7
Two apices long
filamentous s8
Two apices, one a
prominent round lobe s9
Two apices, round lobe,
short spine s10
Three apical lobes s11
Folded laminate s12
Folded laminate
leaf-like s13
Reduced s14
Absent t
Absent u
Short v
Bifid v1
Broad v2
One present w
Two present w1
Short x
Long x1

Quaest. Ent., 1989, 25 (3)

Table 2. Selected characters of South American, Central American and Caribbean species of the Subgenera Brasiella , Gaymara, Plectographa and Cylinde

330

Freitag and Bames

(continued on next page)

Table 2 (continued)

Brazilian Species of Cicindela

331

Quaest. Ent., 1989, 25 (3)

332

Freitag and Bames

Male genitalia. Median lobe with elongate and tapered apex (Figs. 154a-c). Internal sac with flagellum
1.5 loops at base. Tooth and stylet well developed. Arciform piece, right bar, small stiffening rib and central
plate well developed, additional stout sclerite between small stiffening rib and right bar (Figs. 154d, e).

Geographical Variation. — Insufficient material was available to determine
patterns of geographical variation; variable characters appear to be mainly body
colour ranging from greenish to black through coppery, and the maculations of the
elytra.

Relationships. — Cicindela friedenreichi is sister to the lineage that gave rise to
sister species C. piligera and C. obsoletesignata.

Habitat and Period of Activity.- — Unknown.

Geographical Distribution, Localities, Examined Specimens. —

Southern Brazil, Rio Grande do Sul, Santa Catarina (Fig. 157).

Brazil. Rio Grande do Sul. Torres, 1M, 6F, RRMC.

HISTORY OF SOUTH AMERICAN SPECIES OF SUBGENERA BRASIELLA,
GAYMARA, PLECTOGRAPHA AND CYLINDERA

Introduction

Organisms are wards of their history. Each taxon embodies an evolutionary
heritage based on a special kind of ancestry. In time this legacx is altered with
acquired genetic and biogeographical attributes which aie carrier o descendants.
Natural classifications reflect these processes as part of the lain course of
organismic history. They present branching lineages and herita ; changes in a
dichotomous phylogenetic framework.

In the phylogeny section that follows reconstructed phylogenies are based on
methods and philosophy of Hennig (1966). Phylogenetic relationships are inferred
from determinations of apomorphous (derived) and plesiomorphous (ancestral)
characters of adult beetles from which the reconstructed genealogies are expressed.
Biogeographic considerations, based on the reconstructed phylogenies, attempt to
deduce distributions of species and species groups.

Brazilian and related South American taxa of Cicindela were included, which
together appeared to form monophyletic assemblages largely within South America.
We examined adult specimens of 29 Brazilian and 16 non-Brazilian species. In
addition the genitalia of males and females were dissected and their characteristics
documented for each of 29 and 26 Brazilian species and 10 and 11 non-Brazilian
species respectively. Data for 16 non-Brazilian species not examined by us were
obtained from original descriptions, writings and figures in Horn (1915, 1938),
Rivalier (1954, 1955), Mandl (1963, 1973), Vidal Sarmiento (1966a, b, 1967), Pena
(1969), Pena and Barria (1973), and Sumlin (1979).

Phylogeny

The purposes of this section are to postulate phylogenetic relationships of sister
groups and to classify them. Detailed accounts and viewpoints of the method of
phylogenetic reconstructions can be obtained from Hennig (1966), Ross (1974),

Brazilian Species of Cicindela

333

Eldredge and Cracraft (1980), Wiley (1981), Charig (1982), Maynard Smith (1982),
and Patterson (1982).

Phylogenies are deduced in two steps. First character states are determined to be
apomorphous or plesiomorphous. Second, sister groups are established from
synapomorphous (shared apomorphous) character states. It is not always easy to
decide the direction of a character state trend in a group ( in-group comparison) or
whether a character state is apomorphous or plesiomorphous. By comparing
character states of a group with those of related groups ( out-group comparison)
usually one can decide reasonably which is apomorphous and which
plesiomorphous.

Plesiomorphous and apomorphous character states of Brazilian species of
Cicindela are postulated in Tables 1, 2 and 3, and used as the basis for the
construction of phylogenetic diagrams (trees, cladograms) given in Figures 158-163.
In Table 1 each character is described and numbered. Apomorphous states of
non-genitalic characters are assigned a capital letter and those of genitalic characters
a lower case letter. Superscript numbers placed beside the letters indicate one of two
or more apomorphous states of a character, and a series of numbers indicates a
morphocline. In Table 2, points and letters or numbered letters indicate
plesiomorphous and apomorphous character states respectively.

Decisions about plesiomorphy and apomorphy given below are based on
generalized out-group comparisons of character states among North and South
American species groups or subgenera of the genus Cicindela. As an operational
rule for out-group comparisons we followed that of Watrous and Wheeler (1981)
which states that for a given character with two or more states within a group, the
state occurring in related groups is assumed to be the plesiomorphous state. Thus the
word common as it is used here refers to the high frequency of a plesiomorphous
character state among species groups or subgenera.

Body size (1). Average size plesiotypic though arbitrarily viewed, is found in
most species groups.

Head and pronotum: colour (2) and rugosity (3). Dark brown or black with
shallow rugosity is very common and deemed plesiomorphous.

Rugosity between eyes (4). Two deep rugose pits are found only in four species:
apomorphous.

Proepisternum: punctuation (5). Deep widely spaced punctures present in C.
anulipes only: apomorphous.

Elytra: surface (6) and ground colour (7). A dull and dark brown or black
combination is plesiomorphous being the most common condition among groups.

Setae onfrons (8) clypeus (9) genae (10). Absence of setae is the condition of the
majority of species groups: plesiomorphous.

Setae on proepisternum (11) and mesepisternum (12). Density of setae tends to
be a gradient condition, though the distinctly sparse state appears to be widespread:
plesiomorphous.

Quaest. Ent., 1989, 25 (3)

334

Freitag and Bames

Setae on middle of abdomen (13). Absent is the plesiomorphous state.

Setae on lateral portions of abdominal sterna (14). Most species bear setae on
sterna 1-6: plesiomorphous.

Labrum: width (15) and margin (16). The average width and unidentate states
are widespread among species groups: plesiomorphous.

Labrum setae: position (17) and number (18). Most species have submarginal
setae (plesiomorphous); an average number of eight setae is deemed plesiomorphic
among species groups though fewer than 8 is common in Brazilian species.

Mentum tooth: length (19). Average length is the widespread state:
plesiomorphous.

Antenna segments: colour (20). Lack of metallic reflections (pale) is uncommon:
apomorphous.

Pronotum: width (21). Average width widespread though not uniformly so
within species groups: plesiomorphous.

Coupling sulcus: shape (22). A distinct groove straight or slightly sinuate is
found in the majority of species groups: plesiomorphous.

Middle trochanter: seta (23). Absent from most species: plesiomorphous.

Abdominal sternum 5 of female: unpigmented bell-shaped spot (24). Present in a
few species, apomorphous.

Elytra: pale markings (25). Highly varied in detail though general features
somewhat more stable; narrow to broad, complete or almost complete more
prevalent than other combined conditions: plesiomorphous.

Elytra: foveae (26). Obvious though not large near suture and shoulder is
plesiomorphous.

Abdominal sternum 8 of female: apical emargination (27). A somewhat varied
character but stable within some species groups. Broad, average depth:
plesiomorphous.

Abdominal sternum 8 of female: stout apical setae size (28) and number (29).
Average length: plesiomorphous; 2-3 characteristic of most species:

plesiomorphous.

Ventral sclerite: shape (30). Anterio id posterior projections and a median
ridge are uncommon in most species groups of South America: apotypic; and a well
defined median ridge is uncommon: apomorphous.

Ventral sclerite: setae (31). The absent state is widespread: plesiomorphous.

Oviduct sclerite: form (32). A shield shape is the common state: plesiomorphous.

Spermatheca and duct: length (33). A short (1.0- 1.5 mm) spermatheca and duct
is common: plesiomorphous.

Bursa copulatrix: setae (34) and shape (35). The presence of setae and
outpocketings are states uncommon in South Ameri in species: apomorphous.

Median lobe: apex (36). A short unhooked apex is a widespread state and by
out-group comparison considered to be plesiomorphous.

Brazilian Species of Cicindela

335

Flagellum (37). Absence of a flagellum is uncommon within Cicindela
world-wide: apomorphous.

Tooth: condition (38) and shape (39). A sclerotized elongate tooth is common:
plesiomorphous.

Internal sac: two finger-like darkened fields , one spatulate-like (40). These are
apomorphous if present; found in a few species.

Arciform piece: shape (41) and position (42). The narrow and oblique states are
common and therefore plesiomorphous.

Central plate (43). A central plate not joined to the shield appears to be common:
plesiomorphous.

Shield (44) and (45). A distinct shield, and single apex both plesiomorphous.

Small stiffening rib (46). The present rib state is slightly more common than the
absent one and thus deemed to be plesiomorphous.

Stylet (47) and form (48). An average to long stylet is common: plesiomorphous.

Two sagittal sclerotized plates (49). Rarely present: apomorphous.

Cuha (50). A small triangular sclerite near the dorsal side of the flagellum of the
male, shown by Vidal Sarmiento (1966b) to occur in species of Plectographa and
Cylindera: apomorphous.

Because of the large number of diverse characters used in the reconstructed
phylogeny a strict hierarchical system of character weighting was diffk ilt to
establish. As a general rule we judged characters in demonstrating relationships,
from good, if stable and corroborating with other stable characters inter-specifically,
to poor, if unstable intra-specifically. Characters of the male and female genitalia,
morphometric features, loss of a good character, distribution of body setae, colour
pattern of the elytra, and body colour, were generally but not consistently applied in
descending order of value. For accounts of theory and methods of character
weighting we refer the reader to Funk and Wheeler (1986), Neff (1986), and
Wheeler (1986), and, in particular to how they relate to carabid beetles, to Ball and
Nimmo (1983).

In the formation of a genealogy the largest groups of species that consistently
share apomorphous character states are treated as evolutionary units. In our scheme
they are a monophyletic complex of subgenera of Cicindela found primarily in
South America. Four subgenera are recognized, the species of which are listed in
Table 2.

The reconstructed phylogeny of the subgenus Brasiella (Fig. 158) is based on
characters of 29 species listed in Table 2. Genitalic characters of males and females
dissected by us were observed and documented for 1 1 species, and those of males
only were determined from publications for 23 species. Genitalic characters were
not determined for both sexes of four species, and for females of 14 species. Eight
non-genitalic characters were not determined among 21 species.

Seven characters account for 12 evolutionary reversals and 44 characters for 122
parallelisms. Although 452 characters were not determined, one third of the full

Quaest. Ent., 1989, 25 (3)

336

Freitag and Bames

potential character number 1450, we are confident that our reconstructed phylogeny
of this subgenus will stand up to rigorous testing, as the system is based mainly on
complex characters that frequently corroborate each other in various lineages.

The presence of setae (pubescence) on the middle of the abdominal sterna, short
apical setae on abdominal sternum 8 and membranous oviduct sclerite of the female,
and hooked apex of the aedeagus, absent flagellum and central plate of the male,
characterize the original ancestral lineage of Brasiella.

The relative positions of species in the viridicollis group may require changing
later on, because undocumented good characters are absent from this species group
genealogy. However for now we treat C. acuniai and C. viridicollis as sister species
on the basis of the bicolourous adult body, bright blue or green head and pronotum
with brown to coppery elytra though we recognize that different relationships may
be derived among species by differential weighting of B1, F1, R, Z1 or Z3, and when
the genitalia of both sexes for the four species become known.

The aureola species group consists of C. rivalieri, C. amaenula and the sister
species C. aureola/C. horioni. We have not seen adults of C. horioni and have
placed this species mainly on the basis of the maculations of the elytra as drawn by
Mandl (1956: 388). A bell-shaped unpigmented area of the hind margin of sternum
5 of females is present in the latter three species, and also in species of the argentata
species group. This is an unusual character found in no other Brasiella species.
Therefore, we are not comfortable with this separation of the aureola and argentata
groups but it seems the most parsimonious at the moment.

In the misella group, the positions of C. dolosulaffinis and C. tippmanni are
uncertain. We have put them together as sister species on the basis of general
habitus. We have not seen specimens and many adult and genitalic characters have
not been documented (Table 2).

The reconstructed phylogeny of subgenus Gaymara is based on five species
(Table 2, Fig. 159). Two characters account for two evolutionary reversals, nine
characters account for nine parallelisms.

The ancestral lineage of subgenus Gaymara is distinguished by the characters
elongate and tridentate labrum with fewer than eight setae, short apical setae on
abdominal sternum eight of the female, ventral sclerite of the bursa copulatrix with
two elongate posterior projections, both flagellum and small stiffening rib absent
from the internal sac of male, membranous tooth of the internal sac, and central
plate joined to shield.

The reconstructed phylogeny of the subgenus Plectographa (Fig. 160) is based
on apomorphous characters of 18 extant South American species listed in Table 2.
Two characters account for 34 evolutionary reversals and 16 characters for 91
parallelisms. Broad, fused, ragged and spotted maculations of the elytra, widespread
among extant species, appear to have evolved independently at least four times. A
large body size, long spermathecal duct, elongate tooth with a spatulate apex and
folded laminate shield in the internal sac of the male distinguish the ancestral

Brazilian Species of Cicindela

337

lineage of this subgenus. The eugenil apiata lineage stands apart from the others of
this subgenus mainly in its primitiveness. The most difficult lineage to place is the
monobasic species group halophila. Adults of C. halophila superficially resemble
those of the suturalis species group, but male genitalia are radically different. The
median lobe is equipped with a short, uncoiled flagellum somewhat similar to that
found in North American males of subgenus Cicindela and the tooth is short and
rounded unlike the type found in Plectographa males. A long cuna is present and
the shield appears stylet-like. On the other hand female genitalia are similar to those
of the suturalis species group, particularly in the elongate shape of the ventral
sclerite of the bursa copulatrix and presence of approximately eight setae on both
apices of sternum 8. Thus the female genitalia were given special weight in selecting
an uncertain phylogenetic position for C. halophila.

The species C. siccalacicola, C. sinuosa and C. suturalis are probably more
closely related than indicated in Figure 160 but we were unable to work out a
simpler scheme. On the basis of dense appressed setae on the prostemum and
absence of large foveae on the elytra, C. sinuosa and C. suturalis are established
sister species, and, as a consequence, glossy elytra and few erect setae in the middle
of abdominal sterna become parallel characters of C. sinuosa and C. siccalacicola.
If weighting of the two former and two latter characters were reversed C.
siccalacicola and C. suturalis would necessarily be treated as sister species.

The species C. nigrovittata is tentatively presented as a monobasic species group
with ancient links to Plectographa lineages. An unusual combination of
plesiomorphous and apomorphous characters makes it uniquely different from other
species of Plectographa. The male genitalia are distinctly Plectographa- like in
detail. A deep rugosity forming two large swirls or pits on the head between the eyes
of the adults indicates an extraordinary convergence with adults of several species in
subgenus Gaymara.

The reconstructed phylogeny of South American lineages of subgenus Cylindera
is based on nine species (Fig. 161, Table 2). North American species were not
included as a considerable geographical gap exists between them and South
American species. We assumed that species in South America are more closely
related to each other than to any in North America. The relationships of C.
granulipennis and C. malaris are tentative as their male and female genitalia are not
known. An additional 21 and six characters were not determined for these two
species and four others respectively. Seven characters account for eight evolutionary
reversals and 12 characters for 28 parallelisms.

The ancestral lineage is characterized by elytra slightly glossy with a few foveae,
moderately setose proepistemum, average to elongate labrum width, some or all
setae of labrum very near front margin, fewer than eight labrum setae, ventral
sclerite partly covered with setae and with two anterior lobes and two posterior
projections, and shield absent from the bursa copulatrix.

Quaest. Ent., 1989, 25 (3)

Table 3. Apomorphous characters shared by Brazilian subgenera of Cicindela*.

338

Freitag and Barnes

CO

1 — 1

t— 1

, i

«d*

> >

> >

> >

CO

'd*

+->

4->

LD

'd-

CO

CO

<d-

•d-

s-

S_

co

, 1

'd-

cr

c r

'd-

o

o

o

CO

CO

r-

CO

CD

CM

CM

CM

CO

•r-J

•fj

LD

CO

%r~

*«“

CO

I — 1

CO

cn

cn cn

cn cn

CM

CO

CO

4-

4-

s-

<L)

i — 1

+->

CO

cu

<U

O

n3

o

i — t

\ — i

i — i

s_

CO

"O

•o

"O

to

JCZ

r-~

t — i

r— t

i— 1 CM CO

CM CO

cd

CM

fd td

fd

a

a

a

a

a

a

a

CM

i — i

CM

>

>

> >

pH

* — I

r— 1

r— f

CM

ZD

ZD

ZD

CD

c — t

i — 1

i— 1

oo

OO

r"-

i — i

ax

ay

ax

CD

i — l CM

* — t

,—1

CM

i — l

Q_ Cl.

o_

Q.

O-

■d-

* — i

z

Z

z

cn

*— i

-

1— 1

CO

or

zc

re

r-'*

CD

CD

CD

,

CO

Lo-

Li-

Lo_ LO-

Lo_ U_

•d-

Ci

CO

co

C_>

CD

CD

td

fd

s_

sz

CU

Cl

c

rd

fd

fd

<D

r—

s-

S-

cn

r —

td

cn

cu

JO

a>

s-

O

-O

3

•i —

fd

-M

sz

CO

CO

. E

o

•r—

id

>>

cu

i — ■

s_

fd

r—

>>

CO

CD

O-

CD

s-

CU

O)

-O

+->

o

4->

c

CO

CD

c

cd

OJ

r—

CO

CO

CD

CM

- — - CO

CD

"d-

H

CM *

•> — •*

«o

CD CO

-d-

r— 1

*»

<U CM

cn

SZ «•

•»

O -d-

CO

SZ I— I

*» CD

*

r-- co

1 CO

. .

i r^

fd i

fd

< — i

S-

s-

rd

cu fd

cu

SZ 1

"D S-

e

D

sz cu

<u

fd fd

•r- -O

cn

s- s-

1— sz

-D

cr cu

>>•«-

3

o -a

CD . —

CO

+-> sz

' — . >i

U -r-

fd cd

cu

CU r—

sz \

cu

I— >s o. fd

s-

CL. CD

fd sz

sz

w

S- Q-

+->

fd fd

cn fd

a a

O s-

fd rd

+-> cn

JD

E E

a o

>> >

O) +-»

-a

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Brazilian Species of Cicindela

339

Quaest. Ent., 1989, 25 (3)

Fig. 158. Phylogenetic diagram for South American species of Brasiella. An asterisk indicates Brazilian species. Cicindela mandli Brouerius van Nidek has a median lobe with a rounded
hook and C. insularis Brouerius van Nidek, a median lobe with an abrupt hook (★).

chlorostiota* Kollar

340

Freitag and Bames

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species.

anulipes* Horn

halophila Sumlin

Brazilian Species of Cicindela

341

C r—

tO r—

Fig. 160. Phylogenetic diagram for South American species of Plectographa. An asterisk indicates Brazilian
species.

Quaest. Ent., 1989, 25 (3)

kollari* Gist!

342

Freitag and Bames

Fig. 161. Phylogenetic diagram for South American species of Cylindera. An asterisk indicates Brazilian
species.

obsoletesignata* Horn

Brazilian Species of Cicindela

343

ABC DA D B CACB D

Fig. 162. Fifteen possible dichtomous cladograms for the four subgenera of Brazilian Cicindela , and
characters in Table 3 for which apomorphous states are shared by members of each subgenus. Letters
represent subgenera as follows: A - Brasiella, B - Gaymara, C - Plectographa, D - Cylindera. Numbers in
brackets indicate gradient characters.

Quaest. Ent., 1989, 25 (3)

344

Freitag and Bames

<3

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Fig. 163. Phylogenetic tree of the Brazilian subgenera of Cicindela.

Inter-group relationships are summarized in Table 3 and Figures 162 and 163. In
the construction of Table 3, apomorphous states shared by subgenera, and those
which occur in one or more species of a subgenus, were considered as having equal
weight. Ideally apomorphous character states indicative of strong relationships are
shared by all of most species of related subgenera. These apomorphous character
states complement the less frequent apomorphous states. A broad pronotum and
ventral sclerite with two elongate posterior projections are shared by most species of
Gaymara, Plectographa and Cylindera. The absence of a flagellum and small
stiffening rib characterize most species of Brasiella and Gaymara. All species of
Brasiella and three of Cylindera lack a central plate, and most species of Gaymara
and Cylindera have setae on the ventral sclerite and a membranous tooth.

As shown in Figure 162, of the 15 possible dichotomous cladograms for the four
subgenera, either a or c probably indicates the true phylogeny because they each
incorporate the largest number of synapomorphous character states, 1 1 in each. We
favour cladogram a as it places the Gaymara lineage in an intermediate position
between the Brasiella lineage and the Plectographa/Cylindera lineage, with which
the former shares apomorphous character states of the genitalia approximately
equally (Figs. 162a, m).

The subgenus Brasiella is a Neotropical autochthon that occupies mainly the
northern half of South America and Central America where its origins and evolution
undoubtedly took place. It is sister of the Gaymara/ Cylindera lineage (Figs. 162a,
163). Brasiella seems to have retained a largely plesiomorphous habitus with several
apomorphous acquisitions. The absence of a flagellum in the internal sac of the male
is assumed to be an apomorphous state in Brasiella. It implies that earlier lineages

Brazilian Species of Cicindela

345

had a flagellum which was lost twice, once in Brasiella and again in Gaymara. If it
is assumed that ancestors lacked a flagellum, which was then acquired by the
Plectographa/Cylindera lineage parallel to and independent of other groups of tiger
beetles, it is difficult to account for the complex form of this new flagellum. In more
primitive subgenera of Cicindela the flagellum is almost straight, but in South
American species it has 1 .5 loops at the base. Thus we recognize the absence of the
flagellum in Brasiella as being a loss of a somewhat specialized flagellum. Similarly
the membranous oviduct prevalent in Brasiella is considered to be a loss of a well
developed sclerotized one which is widespread in Cicindela.

The subgenus Gaymara is endemic to central-eastern South America. It is sister
to the Plectographa/Cylindera lineage (Figs. 162a, 163). A narrow field-like strip or
a membranous “tooth” of the internal sac of the male is present in most species of
Gaymara , which is remarkably similar to the tooth with a spatulate apex found in
species of Plectographa. Similarities of this degree are normally given a great deal
of taxonomic weight, however, for these taxa it is difficult to conclude that these
structures are homologues. If these structures are indeed synapomorphous the true
genealogy is probably indicated by Figure 162c rather than Figure 162a.

The subgenus Plectographa is sister to subgenus Cylinder a (Figs. 162a, 163).
Most of its species are specialized in a number of characteristics including a broad
and ragged pattern of the elytral maculations, and structure of male and female
genitalia. This subgenus is endemic to the southern half of South America on both
sides of the Andes. Cicindela suturalis which ranges from Brazil to the West Indies
is the northernmost member of Plectographa.

The subgenus Cylindera is the only cosmopolitan South American subgenus of
Cicindela. Most South American species of Cylindera live in central-eastern parts of
that continent.

The relative age and origins of the subgenera are as follows: Brasiella, probable
ancient Neotropical endemic, sister group of Gaymara/Cylindera lineage, originated
in the northern half of South America and Middle America; Gaymara, South
American endemic, sister group of the Plectographa/Cylindera lineage, with
Brazilian origins south of the Amazon basin; Plectographa, sister group of
Cylindera, South American endemic, origins in Argentina; Cylindera, highly diverse
cosmopolitan complex, recent South American lineages probably originated in
regions of southeastern Brazil.

Quaest. Ent., 1989, 25 (3)

346

Freitag and Bames

BIOGEOGRAPHY

Introduction

Here we offer explanations of geographical distributions of extant taxa, their
ancestral origins, and pathways taken to where they now live. This procedure
combines current knowledge of geological and climatic history of the Neotropics,
geographical distributions of the studied species, and their phylogenetic
relationships.

Assumptions

Our integration of these data is based on three assumptions. The first is that
origins of South American subgenera of Cicindela preceded the Cenozoic Era, no
later than early Cretaceous. In addition, early lineages and species groups diversified
throughout the Tertiary Period, while extant species evolved during the late Pliocene
and Pleistocene, beginning about 2 or 3 million years ago. With the exception of
subgeneric origins, these time/lineage associations are in keeping with contemporary
ground beetle studies (Ball, 1985; Noonan, 1985; Ball and Shpeley, 1986).

The second assumption is that there has been a general tendency in Cicindela for
independent lineage adaptations to open country habitats in temperate and tropical
regions (Cazier, 1954; Willis, 1967, 1972; Freitag, 1979; and articles in the journal
Cicindela since 1969). Although riparian and woodland intrusion are plentiful in
various groups, they appear to indicate the primitive and intermedu adaptive states
respectively. This supports the “taxon cycle” concept demonstrated by carabids as
reviewed by Erwin and Adis (1982), in which wetland generalists radiate into a
biotic zone away form the waterside substratum and become evolutionary
specialists. Specializations in savanna and prairie biomes have been major trends in
a broad spectrum of lineages of Cicindela. As will be shown, the main distribution
patterns of South American lineages of Cicindela are closely allied to savanna and
open country conditions.

The third assumption is that speciation in Cicindela follows the allopatric mode
as defined by Mayr (1963), that is geographic speciation. Important to this process
are physical or climatic barriers, which isolate conspecific populations from one
another, disrupt gene flow among them, and ultimately cause the formation of
genetically incompatible separate species.

Hypotheses

Current hypotheses that explain range distributions and geographical histories of
South American organisms synthesize past geophysical, climatic, and biotic
processes which appear to have had a general influence on their biogeography. (For
reviews see Simpson and Haffer, 1978; Webb, 1978; and Haffer, 1981).

One such hypothesis is the Refuge Theory, which postulates that the biotic
richness of tropical forests is created by cha* ;ng vegetations due to climatic
fluxuations (Prance, 1982; Mayr and O’Hara, 86). A special kind of allopatric

Brazilian Species of Cicindela

347

speciation is therefore proposed in this theory which discards geophysical factors as
part of the diversification process.

There is a growing body of evidence, however, that implies forest disturbance
due to modem and past river dynamics is partially responsible for the high
biological diversity in the upper Amazon basin (Salo et al., 1986).

As stated by Haffer (1982) the Refuge Theory holds that forest and non-forest
biomes changed continuously in distribution during the geological past, breaking up
into isolated blocks and again expanding and coalescing under the varying humid to
arid climatic conditions of certain geological time intervals, especially during the
Quaternary.

Also important is the complex Tertiary geology of Central America. Unresolved
are the time of the complete isthmian connection of North and South America and
the extent of dry land within the sea gap before the continents were joined. The
region appears to have served as island repositories for some evolving groups, and
periodic crossings undoubtedly took place throughout the Tertiary, either by island
hopping or across wide stretches of ocean. (Howden and Young, 1981, and Ball and
Shpeley, 1986, discuss these problems in the light of beetle biogeography and
provide references.)

Two hypotheses focus on biogeographic affinities of the Greater Antilles to other
parts of the New World. One view, the vicariance model of Caribbean
biogeography, is that tectonic forces, during the early or middle Tertiary, displaced
portions of the Proto-Greater Antilles (land between North and South America)
north-eastward to form the Greater Antilles, and that pieces of the original fauna
were taken along as insular inhabitants (Rosen, 1975; Guyer and Savage, 1986).

The older dispersal model of Caribbean biogeography followed here proposes
that dispersals from mainland coasts bordering the Caribbean region account for
most of the fauna on the Greater Antilles. Records of offshore flights, up to
distances of 100 miles by C. trifasciata (Erwin, 1979; Graves, 1981) and its wide
geographical distribution in the West Indies (Leng and Mutchler, 1916; Elliott and
Salbert, 1978); the presence of C. marginata in coastal eastern United States,
Bahamas, and Cuba (Leng and Mutchler, 1916; Vaurie, 1952); and recent dispersals
of C. carthagena from Central America or Colombia to Jamaica (Brouerius van
Nidek, 1980; Freitag, 1985) and C. olivacea from Cuba to Florida (Woodruff and
Graves, 1963) are evidence for the dispersal model and examples of the highly
volant nature of adult Cicindela.

Past Climatic, Floristic and Geophysical Processes

Factors which very likely have had a bearing on the diversification of
Neotropical taxa of Cicindela are: (1) formation of South America and its westward
drift across the Pacific Ocean, following breakup of Gondwanaland in the late
Cretaceous (Hallam, 1981); (2) proximal insular connection and eventual joining of
North and South America through the Central American land bridge in the late

Quaest. Ent., 1989, 25 (3)

348

Freitag and Bames

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Table 4 (continued)

Brazilian Species of Cicindela

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Quaest. Ent., 1989, 25 (3)

350

Freitag and Bames

Tertiary; (3) presence of the Guayana and Brazilian shields, which have been above
sea level since Paleozoic time (Haffer, 1981); (4) marine ingressions in the Amazon
basin from the Atlantic and Pacific Oceans during the early Tertiary (Haffer, 1981);
(5) progressive Andean orogeny since the end of the Cretaceous, but principally that
of the Pliocene and Quaternary that led to the emergence of lowlands immediately
surrounding the Andes (Haffer, 1981); (6) origins of seasonal woodlands and
savanna in Patagonia and their gradual spread in South America during the Tertiary
and Quaternary (Webb, 1978); and (7) periodic expansions of non-forested areas
into the Amazon basin caused by glacial episodes in the Pleistocene (Prance, 1982).

Cenozoic vegetational changes were related directly to cooler and drier climates.
The Andean uplift and circulation patterns around 30° S latitude created a rain
shadow in temperate South America, and ultimately contributed to the development
of desert conditions inland and the driest east coast in the southern hemisphere
(Furley and Newey, 1983). Central America and the Amazon basin were not greatly
affected by cooling trends in the Tertiary, and a tropical rain forest dominated the
land.

Expected Geographical Patterns

Given the above, certain geographical patterns may be expected, as follows: (1) a
general fit of vicariance patterns, of any subgenus to the Cretaceous breakup of
Gondwana, and of any species group to the coming together of North and South
America and to the Guayana and Brazilian shields in the Tertiary; (2) grasslands and
open forests to hold the largest number of species as demonstrated by Willis (1972)
for North American Cicindela ; (3) the Amazon basin to be a major region of
diversification for Tertiary and Pleistocene lineages; (4) post-Pleistocene refugia or
major species centers to be found outside of the Amazon basin.

Distribution and Historical Interpretation - Subgenera

Subgenera Brasiella, Gaymara and Plectographa are Neotropical endemics, and
Cylinder a is Cosmopolitan (Tables 4-7).

Of the first three, Brasiella occupies ti orthem half of South America, Middle
America, and the West Indies, Gaymara is ^ nfined to eastern South America, and
Plectographa ranges across South America from central Chile and western
Patagonia to the West Indies. Subgenus Cylindera in South America is confined
mainly to Brazil south of the Amazon basin. This subgenus is part of a complex the
members of which Rivalier (1950, 1954, 1957, 1961, 1963) treated as subgenera
within the genus Cylindera. Subgenus Cylindera s. str. ranges in both the Old and
New Worlds and other members are found in temperate and tropical regions of the
Old World (Fig. 164; Table 7).

The reconstructed phylogeny (Fig. 163) shows three dichotomies (divergences):
Brasiella and lineage Gaymara - Plectographa - Cylindera ; Gaymara and lineage
Plectographa - Cylindera ; and Plectographa and Cylindera.

Brazilian Species of Cicindela

351

The collective distributional pattern of the four subgenera suggests a Gondwanan
ancestry. Divergence-spatial relationships among ancestral lineages are obscure as
the four subgenera are more or less sympatric in South America. The geographical
ranges of Plectographa and Cylindera might indicate a south-north division of their
immediate ancestor in western Gondwanaland.

Concerning the cosmopolitan geography of Cylindera , we propose that the
break-up of Gondwanaland in the Cretaceous was the initial cause of its disjunctions
in the southern hemisphere, that is between South America and Africa, followed by
late Cretaceous dispersal from South America into North America, and from Africa
into Eurasia during the early and middle Tertiary. (See Hallam, 1981, for a review
of plate movement, eustasy, climate, and organic response to them since the early
Mesozoic, as background information for the above hypothesis.)

Distribution and Historial Interpretation - Species Groups and Species

Ecological Notes.- — Species of Cicindela in the Neotropics and elsewhere
occupy similar habitats. Ecological requirements are specific and diverse among
taxa, though water availablility and soil types are principal factors in governing
geographical distributions of species. As a group, members of Cicindela are ground
residents living in open places in most biomes. Common habitat sites are roadsides,
paths in forests, banks of rivers and streams, edges of lakes, ponds, sea beach s and
salt marshes, and fields such as grasslands, pastures and salt playas. A few species
inhabit mountains, deserts, and low wetlands.

Species of Cicindela found in the Amazon River basin are largely ecological
generalists. The majority of these species are found in other areas as well. Tiger
beetles typical of tropical rain forests such as those of Odontocheila and
Ctenostoma, are arboreal, but species of Cicindela are obligate terrestrial forms
unsuited to dense forests and long and frequent periods of flooding, characteristic of
the Amazon lowlands. Nonetheless the fact that widespread and relict species of
Cicindela exist in the Amazon River basin indicates a history of establishments
there.

Data available for 17 (29%) of 61 species studied indicate spatial patterns of
habitat type or habitat factor preference.

Adults of insular and mainland species of the C. viridicollis species group
(. Brasiella ) are found on patches of bare ground and in dry grasslands. Adults of C.
argentata are found in moist grassy places and on river beaches.

Adults of sister species C. melaleuca/C. patagonica ( Plectographa ) are found on
sea coasts. The former species lives near marshes and dry places inland as well.
Three additional species of Plectographa are seashore residents and adults of one
other have been collected on salt playas. Adults of C. suturalis are riparian. The
habitats of these species, one third of a total of 18 species, in this subgenus, suggest
seaside origins and early development for salt tolerance in Plectographa.

Quaest. Ent., 1989, 25 (3)

352

Freitag and Barnes

Table 5. Distribution of species of the subgenus Gaymara according to geographic

area.*

Species Area

S.E. Brazil**

Braz. Hghld.

Amazon
R. Basin

Area No.

4

5

7

chlorosticta group

chlorosticta

V

V

V

staudingeria

V

V

V

nigroreticulata

V

paranigroreticulata
anulipes group

V

anulipes

V

V

TOTAL 5

5(2)

3

2

*Endemic species are shown in brackets.

**Includes NE. Argentina, E. Paraguay, and Uruguay.

Three species of Cylindera are riparian which may be characteristic of the
habitats for other species of this subgenus.

The majority of Neotropical taxa of Cicindela live in places between sea level
and 1500 meters, and a few others live at higher elevations up to 3500 meters.

Taxa of most South American species of Cicindela are active in summer in the
southern hemisphere, appearing anytime from October to April. Taxa within
southern parts of the Amazon River basin are also active in “summer”. Taxa north
of the basin appear to be active during summer of the northern hemisphere, while
those that transcend the basin are active all year emerging in warmer seasons of both
hemispheres. We have not overlooked the importance of these findings to
investigations of speciation and evolution of Cicindela in the tropics. An analysis is
beyond the scope of this study however, and for now we refer to the phenomenon as
“Hemispheric Control of Phenological Activity in Tropical Regions”.

Areas. — Fifteen geographical areas are designated as a basis for analysis of
distribution patterns. They are delimited by assemblages of taxa of Cicindela or
where geographical limits of taxa of Cicindela are congruent. Brief accounts of the
areas follow (taken from Bates, 1961; Garrett, 1981; and Furley and Newey, 1983).
Area numbers are listed with their representative taxa in Tables 4-8.

Table 6. Distribution of species of the subgenus Plectographa according to geographic area.

Brazilian Species of Cicindela

353

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356

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Fig. 164. Distribution of subgenus and species Cylindera s. str.

Brazilian Species of Cicindela

357

1 . Central Chile: Bordered by the west coast and high Andes this is a relatively
wet area, south of the Atacama Desert, and consists of coastal forests, eastern upland
taiga or tundra, and dry low scrubland in the north. Ball and Shpeley (1986) use the
term ‘Southern Trans-Andean South America’ for this area.

2. Southern Argentina: This area, temperate Patagonia north to approximately
Rio Negro, is bordered by the Andes in the west and the east coast. The area is
marked by dry western uplands and admixtures of desert, semi-desert and grasslands
in central and eastern parts.

3. Northern Argentina: This is the Pampas area. It is bordered by the Andes to
the west, the Rio Negro to the south and the east coast. The northern boundary
includes Chaco Central and Entre Rios. This is predominantly a grassland region.
Semideciduous forests are present in the north.

4. Southern Brazil: This area includes the Brazilian states south of Sao Paulo,
Uruguay, northern Argentina, and eastern Paraguay. The east coast, Entre Rios and
Rio Paraguay form the main boundaries. There is no distinct northern boundary.
Northern uplands gradually merge with those in the south end of area 5. The
vegetational zones are primarily coastal rain forests, prairies to the south, and
semideciduous forests to the north.

5. Eastern Brazil (Brazilian Highlands, Brazilian Shield): This vast region to the
south of the Amazon River basin, includes the east coast, eastern highlands, and
central uplands of Brazil. Western boundaries are Rio Araguaia, the pantanal
complex and Rio Paraguay. Semideciduous forests and savanna scrub cover much of
this area. Amazonian gallery forests along river valleys extend from the south end of
the Amazon basin into northern parts of this area, and extensive rain forests are
found on the east coast.

6. Western Paraguay and Eastern Bolivia: This area comprises the Chaco region
of both countries and the eastern Bolivian uplands, and is bordered by the Andes to
the west, Rio Paraguay to the east, Rio Guapore to the north, and Chaco central to
the south. The region is relatively dry dominated by savanna scrub, and
semideciduous forests in central Bolivia, and large river valleys.

7. Amazon River Basin: The area consists of the large upper and main drainage
systems of the Amazon River in northern Brazil. A more or less continuous rain
forest covers the area. Savanna scrub elements intrude southern and northwestern
regions of the basin from areas 5, 6 and 8.

8. Northern South America (Guay ana Shield): This area includes the Guay ana
Highlands, Orinoco River basin, northeastern Colombia, and the north coast of
South America. The Andes proper and Cordillera Oriental just west of the
Maracaibo basin form the designated western border of this area. Savanna scrub
covers western portions. Rain forests and grasslands cover central and eastern parts.

9. Northwestern South America: This area includes Colombia and Ecuador in
and west of the Andes. No endemic species of Cicindela are found there but
geographical ranges of a few species terminate in it. Coastal rain forests and upland

Quaest. Ent., 1989, 25 (3)

358

Freitag and Bames

semideciduous forests are widespread, and smaller zones of savanna scrub are
present near the Caribbean coast of northern Colombia and Pacific coast of southern
Ecuador. Ball and Shpeley (1986) refer to this area as ‘Northern Trans-Andean
South America’.

10. Lower Central America: This area includes Costa Rica and Panama. A series
of islands may have persisted in this area from the mid-Tertiary up until the late
Pleistocene. The topography is mixed and consists of central mountains and
lowlands. Tropical rain forest is widespread. West of the mountains on the Pacific
slope, rainfall and floral richness are reduced.

11. Nuclear Central America: Most of this area, between the Isthmus of
Tehuantepec and southern Nicaragua, has been dry land throughout the Tertiary
(Rosen, 1978), but periodically separated from Mexico (area 12) by high sea levels
(Ball and Shpeley, 1986).

12. Mexico north of the Isthmus of Tehuantepec.

13. Northwestern Mexico and Southwestern United States.

14. West Indies: Greater Antilles.

15. West Indies: Lesser Antilles.

Undesignated areas which are devoid of species are the dry west coast from
southern Ecuador to northern Chile, high Andes, and the southern end of Argentina
and Chile.

The Distributional Pattern. — Five of the seven Brasiella species groups
designated, reside chiefly within the northern half of South America (Table 4). Of
these, species groups aureola and stamatovi are endemic to the continent (areas 3-8,
and 3 respectively), species groups minarum and misella, extend into Middle
America (areas 3-11), and species group argentata is also found in southern islands
of the Lesser Antilles (areas 3-9, 15). Of the two remaining Brasiella species
groups, hemichrysea group is primarily Middle American (areas 9-12), and
viridicollis group is broadly disjunct being present both in Cuba (area 14), and
northwestern Mexico and southwestern United States (areas 12, 13). Thus in terms
of Brasiella species groups the greates iversity is in the northern half of South
America.

The 29 species of Brasiella collects y range from northern Argentina to
southwestern United States and the West Indies (Table 4). Of the 22 species known
from South America, 12 are found south of the Amazon River basin (areas 3-6), four
are within or on the edge of the basin (area 7), and six inhabit northern and
northwestern South America (areas 8-9). Among the remaining seven species, three
are confined to Central America and southern Mexico (areas 10-12), one inhabits
northwestern Mexico and southwestern United States (area 13), and three species
are found in Cuba (area 14).

The pattern of diversity comprises chiefly two species aggregates in South
America. One, south of the Amazon River basin (a 'eas 4-6) diminishes in species
number abruptly southward and gradually northward and eastward. A smaller

Brazilian Species of Cicindela

359

species aggregate north of the basin is confined largely by the Guayana Highlands
and the Andes (area 8).

In closer scrutiny, of the 12 species south of the Amazon River basin four have
substantial geographical ranges. The species C. amaenula and aureola are each
widely disjunct with isolated populations in the basin (Fig. 49), C. obscurella ranges
into southern Uruguay and northern Argentina (Fig. 48), and C. minarum is found
from near the coast to the Vacaria River area in Matto Grosso (Fig. 51). The
remaining eight species have restricted or disjunct ranges. They are C. horioni
(Bolivia), C. balzani (Bolivia, Ecuador), both on the margins of the Amazon basin,
C. dolosulaffinis and C. tippmanni (Paraguay), C. stamatovi (northwestern
Argentina), and C. banghaasi, C. brevipalpis and C. hamulipenis (southeastern
Brazil) (Fig. 51). The species C. dolosula and C. misella are found south, north and
northwest of the Amazon basin and occur in parts of Central America (Fig. 50).

The Amazon River basin is occupied by one widespread species and one local
species. The species C. argentata is the only species of Cicindela found in the arid
northeastern parts of area 5, and it extends into the Caribbean region (Fig. 47). The
species C. pretiosa is found only in the Manaus area (Fig. 48).

Among the six species that inhabit the region north and northwest of the Amazon
River basin those with comparatively larger geographical ranges, C. mendicula
(Nicaragua to western Venezuela) and C. nebulosa (Nicaragua to Colomb i and
Ecuador), are found in Central America and C. venustula occupies the region north
of the Guayana Highlands. The species C. rivalieri and C. venezuelensis are
confined to small areas in Venezuela, and C. insularis is found in Trinidad and on
the adjacent mainland.

Two species have local distributions in Central America and southern Mexico, C.
sphaerodera in the Quiche Mountains of Guatemala and C. mandli in Chiapas,
Mexico. C. hemichrysea ranges more widely from western Mexico to Panama. In
the north C. wickhami inhabits northwestern Mexico and southwestern United
States, and C. acuniai, C. cubana and C. viridicollis occupy Cuba.

Nine species are endemic in the southern end of the Brasiella range, eight may
be geographical relicts (Table 4). Four endemic species are found north of the
Amazon River basin. Three of them are probably geographical relicts. At the
northern end of the range there is one mainland endemic species, and three species,
very likely autochthonous, on Cuba. Only one endemic species is present in Central
America.

These findings indicate that the region south and southeast of the Amazon River
basin (areas 4-6) has been the chief center of diversification for Brasiella. A smaller
region north of the basin (area 8) has been a minor center.

The only two species groups of subgenus Gaymara, chlorosticta and anulipes,
are found in eastern South America. The species have a composite range mainly in
Brazil south of the Amazon River basin (areas 4-5) (Table 5). The area of greatest
diversity is Rio Grande do Sul (area 4) in which at least three and probably all five

Quaest. Ent., 1989, 25 (3)

360

Freitag and Barnes

species exist (Fig. 77). Three species, C. chlorosticta, C. staudingeria and C.
anulipes, are comparatively widespread and enter the southern margins of the
Amazon River basin (area 7). Two species, C. nigroreticulata and C.
paranigroreticulata, appear to have local ranges.

It is concluded from these data that the major center of diversity for Gaymara
has been southeastern Brazil (area 4).

Of subgenus Plectographa, species groups suturalis, melaleuca and apiata,
range largely within northern Argentina (area 3). Species, group halophila is
endemic there, and species group nigrovittata, is endemic to southeastern Brazil
(area 5). Species group diversity in Plectographa is therefore greatest in northern
Argentina (Table 6).

The composite geographical range for the 18 Plectographa species covers most
of South America from temperate Chile and Patagonia to the north coast and West
Indies (Table 6). Thirteen species are resident in northern Argentina (area 3); three
species are known from Chile (area 1); one species is found in eastern Brazil (area
5); and one species is widespread in the northern half of South America and the
West Indies (areas 4-8, 15).

The pattern of diversity consists principally of a large species aggregate in
northern Argentina. Species decline in number from this region, abruptly northward
and southward, but more gradually eastward.

Of the 13 species in northern Argentina three, C. drakei, C. ritesmai and C.
mixtula are found in the west end of this region and the latter species also inhabits
Bolivia and possibly western Paraguay. Three species C. siccalacicola, C.
hirsutifrons and C. eugeni appear to have restricted ranges, mainly in Cordoba.
Intrusions into the eastern portions of northern Argentina, or Paraguay, Uruguay,
and southeastern Brazil have been made by C. halophila , C. sinuosa, C. ramosa, C.
melaleuca (Fig. 113), and C. apiata (Fig. 114). The species C. patagonica seems to
be confined to Rio Grande do Sul, Buenos Aires and southern Uruguay (Fig. 113).
The coastal species C. nivea ranges from Patagonia to Brazil (Fig. 1 12).

Of the three Chilean species, C. nahuelbutae is found locally in Arauco
Province, and C. chiliensis , and C. gormazi extend into southwestern Argentina.

The species C. nigrovittata inhabits a small area in eastern Brazil (Fig. 1 14), and
C. suturalis is widespread north of Argentina (Fig. 1 12).

These data show that all or part of the geographical ranges of 13 species are
located in the northern half of Argentina (area 3), and two others in southwestern
Argentina (area 2). Only three species live entirely outside of northern Argentina. In
terms of endemism, six species are located in northern Argentina (area 3), in which
three are geographical relicts. One species is endemic to Chile (area 1) and another
to eastern Brazil (area 5), both are geographical relicts.

This leads us to conclude that northern Argentina has been the major center of
diversity for Plectographa.

Brazilian Species of Cicindela

361

South American species groups of subgenus Cylinder a, morio and friedenreichi,
are endemic to the northern half of South America, and occupy mainly southeastern
and eastern Brazil (areas 4, 5), Amazon basin (area 7), and marginally, northern
Argentina (area 3).

As a group the nine species of South American Cylindera range across the
middle area of the continent south of the Amazon River from southeastern Brazil to
Ecuador and Peru (Table 7). Six species are found in southeastern Brazil (areas 4,
5). One species is found both in southeastern Brazil and Ecuador (areas 5, 7). One
species is found in Ecuador (area 7), and one other in Peru (area 7).

The pattern of species diversity consists of a broad aggregate in southeastern
Brazil and three isolated taxa in western South America.

Of the seven species found in southeastern Brazil, C. kollari and C. morio enter
southern parts of the Amazon River basin (Figs. 155, 156), C. confluentesignata
ranges near the coast into Uruguay (Fig. 155), C. marquardti is found in Matto
Grosso (Fig. 156), C . friedenreichi exists as a small isolated population at the south
end of this region in Rio Grande do Sul (Fig. 157), C. piligera is widely disjunct,
found only in Minas Gerais and Ecuador (Fig. 157), and C. obsoletesignata ranges
into southern Paraguay and northern Argentina (Fig. 157).

The non-Brazilian species C. granulipennis and C. malaris found in Ecuador and
Peru respectively, along with the Ecuadoran C. piligera population, represent
western area 7 (Amazon basin) isolates of South American Cylindera. None of these
three taxa appears to exist west of the Andes.

Based on these data we conclude that the major center of diversity for taxa of
South American Cylindera has been southeastern Brazil (areas 4, 5).

Geographical distributions of species per designated area and endemism of
species for the four subgenera collectively indicate a consistent pattern (Table 8,
Fig. 165). Northern Argentina (area 3), southeastern Brazil (area 4) and eastern
Brazil (area 5) are the major centers of taxa concentration. They contain the largest
number of species, about 20 in each area.

Together these areas also contain the largest number of endemic species
determined, 17 (54%) of a maximum number of 31 found in all designated areas.
Endemism based on species found only in each area is remarkably high, 37% for
area 3, 24% for area 4, and 23% for area 5. Fourteen species are found in the
Amazon River basin (area 7), and three species (21%) are endemic. Next in
importance is northern South America (area 8), in which four (57%) of seven
species are endemic. Northwestern South America (area 9) and lower Central
America (area 10), each containing six species, are noted for their lack of endemic
species. Nuclear Central America (area 11) contains six species including one
endemic (17%). Three species are endemic to Cuba, (area 14). One endemic species
(10%) of 10 is found in area 6. Other areas (1,2,12,13,15) with three or fewer
species have one or no endemic species.

Quaest. Ent., 1989, 25 (3)

362

Freitag and Bames

Table 8. Number of species and endemic species of subgenera Brasiella, Gaymara,
Plectographa, and Cylindera for designated geographical areas.

Geographical Area

Taxa Total

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15

Species 3 3 19 21 22 10 14 7 6 6 6 1 1 4 2

Endemic 1-755134--1-13 - 31

Species

Phylogenetic - Geographical Relationships and Areas of Diversification

Allopatric patterns of sister lineages, derived in the phylogenetic section, are
important to the identification of geographical origins and regions of diversification.
Figure 166 and Tables 9-12 show comparative geographical positions for sister
lineages of the studied subgenera. Geographical areas 1 to 15 e the basis for
allopatric reference. Allopatric states of sister taxa between and vithin the same
geographical area are included.

Beginning with subgenus Brasiella (Table 9), two pairs of early sister lineages
have allopatric members. One pair is divided between areas 7 and 9 with a small
overlap in area 8. Another pair is allopatric in area 12.

Within species groups there are allopatric members in 13 sister lineages. Based
on nearest areas of these disjunct sister lineages, the area combinations are, three
7/8, two 5/5, two 11/11, and one each of 5/7, 5/9, 6/7, 5/6 or 7/9, 8/11, and 13/14.

Concerning individual areas alone, nine are associated with allopatric sister
lineages in Brasiella. The areas with their respective number of allopatric
associations shown in brackets are as follows: area 7 (7), area 5(5), area 8(4), area
11(3), aiea 9(3), area 6(2), area 12(1), area 13(1), and area 14(1).

For Brasiella lineages then, the main areas in and among which diversification
has occurred are the Amazon River basin (area 7), eastern Brazil (area 5), northern
Brazil (area 8), and the junction of Central and South America (area 9 to 11) (Fig.
116).

With the exception of C. nigroreticulata/C. paranigroreticulata , all members of
sister lineages in Gaymara are sympatric in areas 4, 5 or 7 (Table 10). The former
sister species are allopatric in central parts of southwestern Brazil (area 4) which
probably has been the principal area of diversification for the subgenus as the other
three species of Gaymara exist there as well.

Brazilian Species of Cicindela

363

Members of three pairs of Plectographa sister lineages are allopatric (Table 11).
Their nearest area combinations are 3/1, 3/1 or 2, and 3/4. Area 3 is associated with
all three allopatric combinations, and sympatric sister lineages are most frequently
found there. Thus we conclude that diversification of the majority of Plectographa
lineages has taken place in or adjacent to northern Argentina (area 3).

Four pairs of Cylindera sister species have allopatric members (Table 12), for
which nearest area combinations are 3 or 4/5, 7, 4/4, 4 or 5/7, and 7/7. Based on
these data diversification of Cylindera lineages has been prominent in or among the
Amazon River basin (area 7), southwestern Brazil (area 4), and eastern Brazil (area
5).

As expected, among the principal centers of species diversification the Amazon
basin appears to have played a major role, particularly during the Pleistocene.
Additional evidence of late Pleistocene effects are the small refugia on the eastern
slopes of the Andes along the edges of the Amazon basin, in Bolivia, Peru, and
Ecuador. A few species appear to be wedged between the high mountain and
rainforest barriers (i.e., C. balzani, C. granulipennis, C. horioni, C. malaris, and
western populations of C. piligera).

There is at least one small refugium within the basin near Manaus, that of C.
pretiosa, and isolated populations of C. amaenula and C. aureola.

Sister groups residing on opposite sides of the Amazon basin are evidence of
Amazonian disruptions of widespread ancestral lineages, as for example C.
minaum/nebulosa-mandli-insularis.

Geographical History

Historical aspects of geography for taxa of the four subgenera are pieced together
from the above reconstructed phylogeny, assumptions, hypotheses, evidence for past
climates and geophysical processes of the Neotropics, and distribution patterns of
related and unrelated taxa.

We propose the following events for Brasiella. (1) Early Tertiary vicariances of
early lineages caused by marine ingressions in the Amazon basin, and founder
dispersals over tropical forests in developing open forest and savanna, resulting in
diversification of ancestral stocks into cubana-pretiosa lineage, aureola group,
misella group, stamatovi group, and minarum group. (2) Early Tertiary vicariance of
cubana-pretiosa lineage and its diversification into ancestors of northern South
American (Guayana Shield, area 8) cubana-hemichrysea lineage and eastern Brazil
(Brazilian Shield, area 5) argentata group. (3) Diversification of
cubana-hemichrysea lineage in northern South America, and a middle Tertiary
ocean crossing to Central America resulting in North American viridicollis group
and South American stock of hemichrysea group. (4) Dispersal into northern
Mexico and diversification of viridicollis group, a late Tertiary ocean crossing to
Cuba from eastern Mexico or the United States by cubana-viridicollis lineage and
its Pleistocene diversification on Cuba, and extinctions of southern and eastern

Quaest. Ent., 1989, 25 (3)

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dolosulaf finis- tippmanni/ venezuelensis-dolosula-misella 4/3 to 10 sympatric

(continued on next page)

Sister Lineages Geographical Areas Distributional Relationships

of Lineages

Brazilian Species of Cicindela

365

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366

Freitag and Barnes

Table 10. Phylogenetic - distributional relationships of Gaymara sister lineages.

Sister Lineages

Geographical

Distributional

Species Groups
C. chlorosticta group

Areas

Relationships
of Lineages

C. chlorosticta/C. staudingeria

4-5_7/4-5-7

sympatric

C. nigroreticulata/

C. paranigroreticulata

4/4

allopatric

chlorosticta-staudingeria/
nigroreticulata-paranigroreticulata
C. anulipes group

4-5— 7/4

sympatric

C. anulipes
Early Lineages

4-5

chlorosticta-paranigroreticulata/
C. anulipes

4— 5— 7/4-5

sympatric

mainland populations of viridicollis group. (5) Post middle Tertiary dispersal of
hemichrysea group stock across ocean into Central America and its diversification
there, extinctions of elements on the South American mainland, and a Pleistocene
reinvasion of mendicula-sphaerodera lineage into western Colombia. (6)
Vicariance, dispersal and diversification from middle Tertiary through the
Pleistocene by the argentata group, aureola group, misella group, and minarum
group, within and among the three main centers of taxa concentration, northern
South America (area 8, Guayana Shield), eastern Brazil (area 5, Brazilian Shield),
and the Amazon basin (area 7), mainl) relation to expanding open forests and
savanna and their intermittent intrusic into the Neotropical rainforests. (7)
Pliocene vicariance of ancestral lineage ot nebulosa/mandli-insularis by Andean
orogeny in northwestern South America resulting in C. nebulosa west of the Andes
and ancestor of mandli-insularis east of the Andes, and a Pleistocene founder
dispersal by the latter across the Andes into middle America resulting in C. mandli
and C. insular is.

Extensive sympatry among sister lineages obscures the geographical history of
Gaymara. It is likely that southeastern Brazil was the region in which its
diversification took place, given that all extant taxa exist there.

Distribution patterns of the taxa and phylogenetic-distributional relationships
imply a general history for Plectographa in northern Argentina. Late Tertiary
dispersals westward across the Andes by the ancestral stocks of

Table 11. Phylogenetic - distributional relationships of Plectographa sister lineages.

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Brazilian Species of Cicindela

369

Table 12. Phylogenetic - distributional relationships of Cylindera sister lineages.

Sister Lineages

Geographical

Areas

Distributional
Relationships and
Lineages

Species Groups
C. morio group

C. kollari/C. malaris

5-7/7

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C. confluentesignata/
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Early Lineages

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melaleuca-patagonica/gormazi-chiliensis and of C. ramosa/C. nahuelbutae resulted
in the Chilean species C. gormazi, and the stock of C. chiliensis/C. nahuelbutae
which diversified west of the Andes, and resulted in the latter sister species.

Sympatric relationships of sister lineages of relative early and intermediate age
obscure historical events of Cylindera. Allopatric relationships of recent lineages
however indicate the following events: (1) Pleistocene vicariance and diversification
of stock of C. kollari/C. malaris caused by expansions of rainforest during an
interglacial phase resulting in Amazon basin species C. kollari (east) and C. malaris
(west). (2) Events as in (1) for stock of C. confluentesignata/C . granulipennis
resulting in C. confluentesignata (east Brazil) and C. granulipennis (Ecuador). (3)
Vicariance and diversification of ancestral stock of C. friedenreichi/C. piligera-C.
obsoletesignata in southeastern Brazil during Pliocene or early Pleistocene, possibly
as a result of habitat decrease caused by increased aridity in eastern Brazil, resulting
in C. friedenreichi and stock of C. piligera/C. obsoletesignata , and vicariance of the
latter resulting in C. piligera and C. obsoletesignata. (4) Dispersal of C. piligera

Quaest. Ent., 1989, 25 (3)

370

Freitag and Bames

Fig. 165. Distribution of species of Brasiella, Gaymar< lectographa, and Cylindera according to
geographical area. Symbols: the number beside a circle d' es geographical area; squares and circles are

major and minor centres of species concentration respec ly; letters denote subgenus; first and second

numbers beside letters denote number of species and endemic species respectively.

Fig. 166. Number of geographical relationships of allopatric sister lineages of Brasiella, Gaymara,
Plectographa and Cylindera. Symbols: the number beside a circle denotes the geographical area; squares
and circles are major and minor centres of species diversification respectively in designated geographical
areas; letters denote subgenus; number beside letters denotes number of allopatric sister lineages, within
(inside circles) or between (outside circle) geographical areas.

Quaest. Ent., 1989, 25 (3)

372

Freitag and Bames

throughout the southern part of the Amazon basin during the last glacial episode,
and its vicariance pattern recently caused by the spread of rainforest in the basin
resulting in two allopatric populations, one in Ecuador and another in eastern Brazil.

Summary

Origins of the four studied subgenera were in western Gondwana prior to the
formation of South America, and subsequent Neotropical diversification within
subgenera took place mainly in the northern half of South America.

Major centers of species concentration, or refugia, of Brasiella, Gaymara and
Cylindera are in eastern Brazil south of the Amazon River basin and that of
Plectographa in northern Argentina. These centers have been long standing, perhaps
for the entire Cenozoic Era, given their mix of related and unrelated taxa of different
phylogenetic age. They were the chief sources of dispersal during hospitable
climatic and geophysical episodes, and into which taxa retreated during the hostile
episodes.

Major barriers over or through which taxa periodically passed, became
geographically isolated and diversified, include the Amazon basin, Andes in Chile
and Colombia, and Tertiary Central America. At the present time members of many
taxa pairs are found on opposite sides of any one of these barriers Arid regions in
central parts of northern Argentina, and Pleistocene marim ingressions in
northeastern Argentina (Noonan, 1985) may have been barrier;, effective in the
diversification process of Plectographa lineages.

Climatic reversals resulting in vegetational changes in the Amazon basin during
the last half of the Tertiary and Pleistocene were a fundamental cause of taxa
formation. Open country conditions developing in glacial periods allowed general
expansions of ranges into the basin. Division, isolation, and diversification of
populations occurred with the return of humid tropical forests during interglacial
periods.

PROBLEMS AND PREDICTIONS

1 . Definitive taxonomic positions of C. stamatovi, C. halophila, C. mixtula, C.
nigrovittata, C. malaris and C. granulipennis will be partly or wholly resolved by
careful examinations of external and genitalic characters, and the reassessment of
character weights.

2. Subgenus Brasiella species for which genitalic structures are not yet known,
will have the flagellum absent from the male genitalia and the oviduct sclerite will
be replaced by a membrane in females. Similarly, Plectographa species will have an
elongate tooth with or without spatulate apex in males and ventral sclerite with two
elongate posterior projections in females. Subgenus Cylindera species will have the
central plate absent from males and ventral sclerite with two anterior lobes and two
posterior projections in females.

Brazilian Species of Cicindela

373

3. The relationships analysis of Cylindera lineages is incomplete in the context
of their New World history. Some South and North American taxa appear more
closely related than we earlier suspected. This may be so for other groups not
currently recognized as part of the Cylindera complex. One example is North
American subgenus Dromochorus Guerin, the adults of which have a general
habitus remarkably similar to those of the morio group.

4. This biogeography will change with changes in the reconstructed phylogeny or
by geographical range extensions. The latter is likely to occur as new collections of
specimens are made for taxa in areas peripheral to and within western parts of the
Amazon basin. Until now that region has not been well collected, and what appear
to be disjunct distributions may turn out to be more or less continuous geographical
ranges. Candidates for range changes are C. misella, C. dolosula and C. piligera,
among others.

5. Additional species of subgenus Brasiella that may be discovered will be in the
New World, and mainly from eastern Brazil.

6. Additional species of subgenus Gaymara will be South American from
southeastern Brazil.

7. Additional species of subgenus Plectographa will be from northern Argentina.

8. Additional South American species of Cylindera will be from eastern Brazil
and the Amazon River basin.

9. It is evident that most species are closely associated with unforested regions.
Many related taxa are divided by humid forests, particularly those of Amazonia.
Dispersal routes followed by ancestors of these disjunct taxa have not been
identified in our study. We do know that humid forests periodically gave way to
grasslands and open country, but we do not know where they developed, and if there
existed major and minor dispersal routes. In Webb’s (1978) review of South
American history of savanna vertebrates two major north-south routes are discussed,
the Andes, “high road”, and the Amazon basin “low road”. Both routes were
followed by non-forest vertebrate species. Taxa of Brasiella and Cylindera appear to
form a composite geographical distribution pattern that coincides with the “low
road”. It extends from Colombia to southeastern Brazil through the western central
and southern areas of the Amazon basin. Collections in western Brazil, Bolivia,
Peru, and Ecuador will add distributional evidence in support of the Amazonian
route for South American taxa of Cicindela.

ACKNOWLEDGEMENTS

This study was made possible with the generous help of the following people
who sent specimens on loan from their own collections or collections in their
charge: M.E. Bacchus and G. Kibby (BMNH, London); G. Ekis and R. Davidson
(ICCM, Pittsburg); T.L. Erwin and D.R. Davis (USNM, Washington D.C.); H.F.
Howden (Carleton University, Ottawa); M.M. Kaulbars (Carleton University,

Quaest. Ent., 1989, 25 (3)

374

Freitag and Barnes

Ottawa); D.H. Kavanaugh (CASC, San Francisco); H. Morge (IPZE, Germany);
R.R. Murray (Fort Worth, Texas); G. Scherer (ZSMS, Germany); A. Smetana and
J.E.H. Martin (BRI, Ottawa).

We are particularly grateful to those persons who permitted the senior author and
his colleague W.M. Graham to visit their institutions, and to others who kindly
assisted in the examination of specimens in their care: P.E. Vanzolini, N. Papavero,
C. Costa, M.E. Jorge-Silva (MZSP, Sao Paulo); M.A. Monne (MNRJ, Rio de
Janiero); N.E. Penny (INPA, Manaus); H. Perrin, J. Menier (MNHP, Paris).

G.E. Ball, Edmonton, Alberta, W.M. Graham, Thunder Bay, Ontario, who also
joined the senior author in a visit to institutions in Brazil, and R.L. Huber Prairie
Village, Kansas, read all or part of the manuscript and made many helpful
suggestions that greatly improved the final version.

The study was financed by the National Research Council of Canada and Natural
Sciences and Engineering Research Council of Canada through their respective
bilateral exchange programs with France and Brazil, and NSERC Operating Grant
A-4888 held by the senior author.

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Rivalier, E. 1955. Les Brasiella du groupe de argentata F. Ibid 22(2): 79-100.

Rivalier, E. 1957. Demembrement du genre Cicindela L. III. Faune Africo-
Malagache. Ibid 24: 312-342.

Rivalier, E. 1961. Demembrement du genre Cicindela L. (suite). IV. Ibid., 28(3):
121-149.

Rivalier, E. 1963. Demembrement du genre Cicindela L. (fin). V. Faune
Australienne. Ibid 30(1): 3(M18.

Rivalier, E. 1970. Cicindelidae (Coleoptera) recoltes en Guyane Fran£aise pa la
Mission du Museum National d’Histoire Naturelle. Annales de la Societe
Entomologique de France 6 (new series): 857-859.

Rosen, D.E. 1975. A vicariance model of Caribbean biogeography. Systematic
Zoology 24: 431-464.

Rosen, D.E. 1978. Vicariant patterns and historical explanation in biogeography.
Ibid 21: 159-188.

Ross, H.H. 1974. Biological Systematics. Addison-Wesley Publishing Co., Inc.,
Reading, Massachusetts. 345 pp.

Salo, J., R. Kalliola, I. Hakkinen, Y. Makinen, P. Niemela, M. Puhakka and P.
Coley. 1986. River dynamics and the diversity of Amazon lowland forest. Nature
322: 254-258.

Schilder, F.A. 1953. Studien zur evolution von Cicindela. Wissenschaftliche
Zeitschrift der Martin-Luther Universitat Halle-Willenberg. Mathematisch-

Brazilian Species of Cicindela

381

Natursissenschaftliche Reich 3(2): 539-576.

Simpson, B.B. and J. Haffer. 1978. Speciation patterns in the Amazon forest biota.
Annual Review of Ecology and Systematics 9: 497-518.

Steinheil, E. 1875. Beschreibung neuer Arten aus Colombia. Coleopterologische
Hefte 13: 95-103.

Sumlin, W.D., III. 1979. A brief review of the genus Cicindela of Argentina
(Coleoptera: Cicindelidae). Journal of the New York Entomological Society
87(2): 98-117.

Thomson, J. 1857. Description de quatorze especes nouvelles. Archives
Entomologiques 1: 129-136.

Varas Arangua, E. 1925. II. Contribution - Cicindelidae recogidos por F. Schade en
Paraguay. Revista Chilena de Historia Natural Pura y Applicada 29: 36^40.

Varas Arangua, E. 1930. Contributions al estudio de los Cicindelidae. VII. Notas
zoogeograficas sobre algunos Cicindelidae Americanos. Revista Chilena de
Historia Natural Pura y Applicada 34: 152-154.

Vaurie, P. 1952. Insect collecting in the Bimini island group, Bahama Islands.
American Museum Novitates 1565: 1-24.

Vidal Sarmiento, J.A. 1966a. Determinacion generica en base a estructuras genitales
de la hembra de Cicindelidos (Coleoptera). Physis 26: 253-261.

Vidal Sarmiento, J.A. 1966b. Las especies Argentinas de los generos Cicindelidia
Riv., Brasiella Riv., y Cylindera West. Revista del Museo do la Plata, section
Zoologia 9(68): 25^46.

Vidal Sarmiento, J.A. 1967. Determinacion a nivel especifico en hembras de
Cylindera Westwood (Coleoptera: Cicindelidae). Acta Zoologica Lilloana,
Argentina 23: 207-216.

Watrous, L.E. and Q.D. Wheeler. 1981. The out-group comparison method of
character analysis. Systematic Zoology 30: 1-11.

Webb, S.D. 1978. A history of savanna vertebrates in the new world. Part II: South
America and the great interchange. Annual Review of Ecology and Systematics
9: 393^126.

Westwood, J.A. 1831. Memoire pour servir a l’histoire naturelle de la famille des
Cicindeletes (1). Annales des Sciences Naturelles 22: 299-317.

Wheeler, Q.D. 1986. Character weighting and cladistic analysis. Systematic
Zoology 35(1): 102-109.

Wiley, E.O. 1981. Phylogenetics: The theory and practice of phylogenetic
systematics. John Wiley and Sons, New York. 439 pp.

Willis, H.L. 1967. Bionomics and zoogeography of tiger beetles of saline habitats in
the central United States (Coleoptera: Cicindelidae). University of Kansas
Science Bulletin 47(5): 145-313.

Willis, H.L. 1968. Artificial key to the species of Cicindela of North America north
of Mexico. Journal of the Kansas Entomological Society 41(3): 303-317.

Willis, H.L. 1972. Species density of North American Cicindela Cicindela 4(2):

Quaest. Ent., 1989, 25 (3)

382

Freitag and Bames

29-34.

Woodruff, R.E. and R.C. Graves. 1963. Cicindela olivacea Chaudoir, an endemic
Cuban tiger beetle, established in the Florida Keys (Coleoptera: Cicindelidae).
The Coleopterists’ Bulletin 17: 79-83.

Brazilian Species of Cicindela

383

INDEX TO NAMES OF TAXA
(Synonyms in italics)

jVIILY GROUP TAXA
indelina, 248
indelini, 248
nostomatini, 248
*acephalini, 248
mtocheilina, 249
thymina, 249

SfERA AND SUBGENERA
siella Rivalier, 246, 249, 254,
278, 332, 335-336, 339,
343-345,350-351,358-359,
362-363, 370-373
siliella Mandl, 254
indosa Motschulsky, 246, 317
indera Westwood, 246, 249,
254, 284,317, 332, 337,
|342-345,350-352, 356,
361-363, 369-373
mochorus Guerin, 373
unara n. subg., 249, 254, 278,
284, 332, 336-337, 344-345,
1350, 359-360, 362, 366,
370-373

btographa Rivalier, 249, 254,
284, 289, 298-299, 305,317,
332, 336-337, 341, 343-345,
1350-351,353,360, 362-363,
366, 370-373

iCIES AND SUBSPECIES
mpsa Chaudoir, Cicindela , 320
jniai Mutchler, Cicindela, 336,
359

oguttata Audouin and Brulle,

Cicindela , 27 1

tenula Chaudoir, Cicindela,

253, 255-256, 258, 262,
270-271,336, 359, 363
anulipes Horn, Cicindela, 252,
278-283, 288-289, 360
apiata apiata Dejean, Cicindela,
305, 307

apiata claussenii Putzeys,
Cicindela, 305, 307
apiata Dejean, Cicindela, 253,
290-291, 297, 305-307, 360
argentata argentata Fabricius,
Cicindela, 265

argentata ecuadorensis Mandl,
Cicindela, 265

argentata Fabricius, Cicindela,
253-257, 260, 265-268, 270,
273-274, 351,359
argentata macella Rivalier,
Cicindela, 265-266
argentata pallipes Fleutiaux and
Salle, Cicindela, 265-266
argentata semicircumscripta
Mandl, Cicindela, 265
argentata umbrogemmata Horn,
Cicindela, 265, 267
argyrosticta Gemminger and
Harold, Cicindela , 271, 359
aureola alvarengai Mandl,
Cicindela, 271-272
aureola alverengai Mandl,
Brasiliella, 27 1

aureola aureola Klug, Cicindela,
271

aureola jatahyana Rivalier,
Cicindela, 271-272
aureola Klug, Cicindela, 25 1 ,
255-256, 258, 262, 270-272,
336, 363

balzani Horn, Cicindela, 359, 363

lest. Ent., 1989, 25 (3) -

384

banghaasi Horn, Cicindela, 250,

254- 256, 259, 264, 275-277,
359

brevipalpis Horn, Cicindela, 25 1 ,

255- 256, 259, 264, 275-277,

359

celeripedestris Horn, Cicindela,
267-268

chiliensis Audouin and Bruelle,
Cicindela, 360, 369
chlorocephala Mannerheim,
Cicindela , 298

chlorosticta Kollar, Cicindela, 252,
279-286, 289, 360
chlorosticta smaragdina Horn,
Cicindela , 284

chrysocollis Mandl, Brasiella , 242,
244, 267-268

confluens Horn, Cicindela, 319
confluentesignata Horn, Cicindela,
250, 275,308-310,314,317,
319-320, 361,369
conspersa Dejean, Cicindela,
300-301

cubana Leng and Mutchler,
Cicindela, 359

cyanitarsis Kollar, Cicindela, 271
denticulata Klug, Cicindela, 320
dimidiaticornis Lucas, Cicindela,
317-318

dolosula Rivalier, Brasiella, 273
dolosula Rivalier, Cicindela, 253,
255-256, 263, 266, 272-274,
359, 373

dolosulaffinis Mandl, Cicindela,
336, 359

drakei Horn, Cicindela, 360
egaensis Thomson, Cicindela, 265
egena Chaudoir, Cicindela, 320
eugeni Castelnau, Cicindela, 307,

360

friedenreichi Dokhtouroff,

Freitag and Barnes

Cicindela, 254, 308-309,
312-313,316, 322, 324, 332,
361,369

germanica Linnaeus, Cicindela,
317

gormazi Reed, Cicindela, 289,

298, 360, 369

granulipennis Bates, Cicindela,
319, 337,361,363,369, 372
guerin Gory, Cicindela, 265
halophila Sumlin, Cicindela, 337,
360, 372

hamulipenis Horn, Cicindela, 253,
255-256, 259, 264, 275-277,
359

hebraea guadeloupensis Fleutiaux
and Salle, Cicindela, 298
hebraea Klug, Cicindela, 298-299
hemichrysea Chevrolat, Cicindela,
359

hieroglyphica Klug, Cicindela,

298

hirsutifrons Sumlin, Cicindela,
301,360

hirticollis Dejean, Cicindela, 317
horioni Mandl, Cicindela,
271-272, 336, 359, 363
inaequalis Motschulsky,
Cicindosa, 320

insidiosa Gistl, Cicindela, 298
insularis Brouerius van Nidek,
Cicindela, 359, 366
intricata Brulle, Cicindela, 278,
300, 339

kollari Gistl, Cicindela, 253^
308-310,313-314,317-318,
361,369

lucorum Gistl, Cicindela, 265
malaris Horn, Cicindela, 318, 337,
361,363,369, 372
mandli Brouerius van Nidek,
Cicindela, 278, 339, 359, 366

Brazilian Species of Cicindela

385

marquardti Horn, Cicindela, 25 1 ,
308-309,311,313,315,317,
321-322, 361

melaleuca Dejean, Cicindela, 250,
259, 290-291,293,296,
301-302, 304, 351,360
mendicula Rivalier, Cicindela, 359
minarum Putzeys, Cicindela,

254- 256, 259, 264, 275, 277,
359

misella Chaudoir, Cicindela, 252,

255- 256, 263, 266, 272-274,
359, 373

misella misella Chaudoir,
Cicindela, 274

misella transversalis Rivalier,
Cicindela, 274

mixtula Horn, Cicindela, 318, 360,
372

morio Klug, Cicindela, 252, 290,
308-311,313,315,317,
320-322, 361

nahuelbutea Pena, Cicindela, 360,
369

nebulosa Bates, Cicindela, 278,
359, 366

nigroreticulata Horn, Cicindela,
252, 279-284, 286-289, 360,
362

nigrovittata Horn, Cicindela, 250,
290-291, 293-294, 297,
304-305, 337, 360, 372
nivea Kirby, Cicindela, 250,
290-292, 298, 301, 360
nivea nivea Kirby, Cicindela, 295,
300-301

nivea orbignyi Guerin-Meneville,
Cicindela, 300-301
nocturna Steinheil, Cicindela ,
298-299

obliquealba Motschulsky,
Cicindosa, 317, 320

obscurella constricta Rivalier,
Cicindela , 267-268
obscurella Klug, Cicindela, 251,
255-257, 261, 265-269, 359
obsoletesignata Horn, Cicindela,
253,308-309,312-313,
322-324, 332, 361, 369
ocskayi Gistl, Cicindela , 320
pallidipes Mandl, Brasiliella, 268
paranigroreticulata n.sp.,

Cicindela, 251, 279-284,
287-289, 360, 362
patagonica bergiana Horn,
Cicindela, 303

patagonica Brulle, Cicindela, 250,
290-291,296, 301,303-304,
351,360

patagonica cherubim Chevrolat,
Cicindela, 303

patagonica patagonica Brulle,
Cicindela, 303-304
piligera Horn, Cicindela, 251,
308-309, 31 1-313, 322-324,
332, 361,363,369, 373
pretiosa Dokhtouroff, Cicindela,
251,255-256, 258, 261,
265-266, 268-269, 359, 363
pseudoaragentata Mandl,
Brasiliella , 265, 268
ramosa Brulle, Cicindela, 360, 369
ritsemai Horn, Cicindela, 360
scutellaris Gistl, Cicindela , 317
siccalacicola Sumlin, Cicindela,
337, 360

sinuosa Brulle, Cicindela, 299,
337, 360

stamatovi Sumlin, Cicindela, 359,
372

staudingeri Horn, Cicindela, 285,
360

staudingeria Horn, Cicindela, 252,
279-286, 289

Quaest. Ent., 1989, 25 (3)

386

Freitag and Bames

suturalis Fabricius, Cicindela, 254,
290-292, 294-295, 298-299,
337, 345,351,360
suturalis helvaea Horn, Cicindela ,
298

tippmanni Mandl, Cicindela, 336,
359

trifasciata boliviana Mandl,
Cicindela , 298

trifasciata Dejean, Cicindela , 298
trifasciata Fabricius, Cicindela,

347

tripunctata Dejean, Cicindela , 267
venezuelensis Mandl, Cicindela,
359

venustula Gory, Cicindela , 266,
359

viridicollis Dejean, Cicindela, 336,
351,359

wickhami Horn, Cicindela, 359

Book Review

387

BOOK REVIEW

SCHWALM, F. E. 1988. Insect Morphogenesis. S. Karger AG, Basel, xix + 356 pp.,
94 figs., 14 tables, subject index. $ 261.00 CAN!

Among the most active research areas in Biology today is that dedicated to
understanding how genes control embryonic development in animals. Although
progress towards achieving this goal has been phenomenal this past decade,
principally because of the imaginative use of genetic engineering techniques, it has
resulted from intensive study of less than a dozen, easy-to-maintain, “lab animals”:
the nematode, Caenorhabditis elegans, a few sea urchin and annelid species, the
amphibian Xenopus laevis, Drosophila melanogaster, and the house mouse, Mus
musculus. Not the least among the dangers of such narrow focus is production of a
generation of biologists expert in the burgeoning techniques of molecular biology
but unable to teach development from a comparative viewpoint and only at the
lowest levels of biological organization. These levels are remote from the personal
experience of most students and inaccessible to the senses but for ‘laboratories’ of
high tech equipment. Also, it can be argued (e.g. Bartholomew, 1986. Bioscience 36:
324-329) that to ask incisive questions about any aspect of its life, we must know the
phylogenetic position of the animal being studied and have knowledge of higher
levels in its organization particularly those concerning the whole animal and how it
lives. Thus, is there still a place for a ‘comparative embryology’ such as this volume
by Fritz Schwalm.

The book contains a preface by Friedrich Seidel (the “father” of experimental
insect embryology), an introduction (3 pp.) and eight chapters concerning,
respectively: systematics (4 pp.), imaginal morphology (17 pp.), gametogenesis and
sex determination (63 pp.), insect culture and egg collecting techniques (13 pp.),
oviposition and parental care (7 pp.), mating, egg structure and fertilization (19 pp.),
tables of embryonic development (12 pp.) and a concluding chapter (at 147 pp., the
“meat” of the book) on the structural aspects of insect embryogenesis.

In his preface, Seidel summarizes some high points in the early experimental
study of insect embryos, emphasizing particularly results from his laboratory in the
1920’s that demonstrated the existence of three physiological centres whose activity
was required for normal blastoderm (the cleavage centre) and germ anlage
formation (the activation centre) and for subsequent differentiation (the
differentiation centre) of damselfly ( Platycnemis pennipes ) embryos. In addition, he
comments at some length on the negative impact that evolutionary thinking has had
on progress in experimental embryology and lauds Schwalm for writing “a
monograph on ‘pure’ morphology” devoid of evolutionary speculation. Chief among
the ‘destructive’ contributions he mentions is Ernst Haeckel’s [1866] ‘Biogenetic
Law’ (i. e. “Ontogeny recapitulates phylogeny”), pervasive in the late 19th and early
20th centuries, which suggested that since phylogeny was the “cause” of ontogeny,
its experimental analysis was unnecessary. Because of his stature at that time in the

Quaest. Ent., 1989, 25 (3)

388

Book Review

European scientific community, Haeckel was successful for some years in
preventing such analysis from beginning. (Haeckel also postulated the existence of
an hypothetical ancestral taxon, the “Gastraea” whose adult stage was never found
despite numerous attempts to do so and which was thought to be recapitulated in the
ontogeny of many modem animals as the gastrula). Seidel considers the primary
contribution of developmental morphology to be provision of a structural basis for
experimental study and implies that use of developmental information to establish
homology of structure and to reconstruct phylogeny is of minor importance. The
many biologists to-day ( e.g Alberch, Bonner, Gould, Nelson, Smith, etc.) who are
attempting to re-introduce consideration of ontogeny into evolutionary thinking,
would disagree.

Seidel also discusses the concept of the “Korpergrundgestalt” or basic body
pattern : an embryonic stage in the development of an animal that clearly identifies
the higher level taxon to which that animal belongs. Members of a monophyletic
lineage not only express their common phylogenetic origin in an easy to see way for
the first time at this stage but diversify from it with subsequent development. For
most hexapods, this stage is the “polypod” embryo: a fully segmented form usually
bearing paired appendages on most of its body segments. Species as distantly related
as bristle tails, mayflies, cockroaches and beetles pass through b on their way to
hatching. (One should not confuse this concept with the “( ound Plan” of
phylogenetic systematists which constitutes the collective, ancestn L character states
of a taxon).

In his introduction, Schwalm refers briefly to some of the recent, illuminating,
experimental work on regulatory genes now being carried out in Drosophila
embryos: specifically the ‘segmentation' genes whose normal expression results in
determination of blastoderm cells to organise into a repeating series of homologous,
segment primordia and the ‘homoeotic’ genes which specify segment identity and
position. He recalls too E. B. Lewis’ idea that the mesothorax is the ‘ basic body
segment' or “segment grundgestalt” upon which products of the homoeotic genes
work to generate the diverse insect segments we now see.

Schwalm’s objectives in writing the book were to: “...present embryonic
morphogenesis in a sequence of ‘typical’ events which interdigitate with earlier and
later events as the embryo forms...” and to identify differences in development
among insects that are worthy of further analysis. He is reasonably successful in
achieving these goals.

His chapters on systematics and adult morphology were obviously written by one
who has little knowledge of or interest in either since they are poorly done and are
not referred to again. Chapter 1 includes two cladograms revised from the works of
Hennig, Kastner and Kristensen that can be used by the reader to organise the
observations presented elsewhere in the book within an evolutionary framework. He
considers the entognathous apterygotes (Collembola, Protura, Diplura) to be
monophyletic and to constitute the sister group of the remaining insects; not to be

Book Review

389

separate classes as many recent authors (e.g. Jamieson, Manton) have suggested.
Chapter 2 contains some appropriate, though superficial, information on male and
female reproductive systems and genitalia, but its brief summaries of distribution
and behavioural adaptations seem totally out of place in a book on embryogenesis.
He offers nothing about the role of sexual selection in shaping insect genitalia (see
Eberhard, 1985. Sexual Selection and Animal Genitalia. Harvard).

In Chapter 3 are brief but reasonably up-to-date summaries of sex determination,
hermaphroditism, parthenogenesis, heterogony and viviparity, oogenesis and egg
membranes, spermatogenesis, sex ratios, life cycles and metamorphosis, neoteny,
paedogenesis, caste formation in social insects, and quiescence and diapause.
Coverage of comparative aspects is encyclopaedic and is presented in tables that
include appropriate literature citations: Table I (6 pp.): oogenesis; II (2 pp.):
synthetic activities of oocyte nuclei; III (2 pp.): types of parthenogenesis; IV (1 p.):
sex ratios in Hymenoptera; and V (4 pp.): fecundity and duration of developmental
stages.

Rearing and egg collecting methods are described in Chapter 4 for 69 species in
16 orders (Archaeognatha-2 spp., Zygentoma-2, Ephemeroptera-3, Odonata-10,
Plecoptera-1, Phasmatodea- 1 , Orthoptera-6, Dictyoptera-2, Isoptera-2, Hemiptera-5,
Coleoptera- 1 1 , Hymenoptera-6, Trichoptera-1, Lepidoptera-6, Mecoptera-1, and
Diptera-11) and should be useful to investigators seeking for research beasts other
than Drosophila upon which to work.

A straight, descriptive account of oviposition and egg masses in Chapter 5
informs us that Collembola “...deposit about 250 eggs in batches...” and that ”...
Psocoptera lay individual eggs or form cocoons with egg batches...”. Schwalm
seems not to realise that individual females, not taxa, deposit eggs. Generalizations
such as these are of little value since they are based on detailed knowledge of a few
species whose identities are only sporadically revealed.

Mating behaviour, and the size (Table VI: 3 pp.), shape, maturation, internal
structure and fertilization of eggs are discussed in Chapter 6 while tables of
embryogenic development comprise Chapter 7: Table VII (6 pp.): the literature of
descriptive embryogenesis arranged by order and VIII (4 pp.): timetables for major
morphogenetic events in the embryogenesis of 52 species in 18 orders
(Collembola- 1 sp., Archaeognatha-2, Zygentoma-1, Ephemeroptera- 1 , Odonata-1,
Plecoptera-1, Phasmida-1, Orthoptera-5, Dictyoptera-3, Isoptera-1, Hemiptera-4,
Coleoptera-6, Megaloptera- 1 , Neuroptera-1, Hymenoptera-6, Trichoptera-1,
Lepidoptera-10 and Diptera-4) . Both tables are filled with information that would
require much effort to extract from the primary literature.

The final chapter is a detailed, well illustrated summary of embryonic
development beginning with cleavage and ending with hatching. Coverage is
extensive and up-to-date and on insects selected from throughout the class. Only
very recent discoveries are not mentioned (e.g., Ball and Goodman’s [1983, 1985]
ingenious experimental work on muscle pioneers in locust embryos [Nature 301:

Quaest. Ent., 1989, 25 (3)

390

Book Review

66-69; Dev. Biol. Ill: 383-418], Campos-Ortega and Hartenstein’s [1985] superb
book on the embryogenesis of Drosophila and recent papers on the embryogenesis
of Thysanoptera [Heming, 1979. J. Morph. 160: 323-344; 1980. Ibid. 164: 235-263;
Haga, 1985. Rec. Adv. Ins. Embryol. Jap. 1: 45-106; and Moritz, 1988. Zool. Jb.
Anat. 117: 1-64]). The book contains 1255 references of which 156 were published
after 1980.

Much comparative information is, again, presented in tables: Table IX (4 pp.):
duration of nuclear cycles during cleavage; X (1 p.): vitellophage fate; XI (1 p.):
pole cell formation; XII (1 p.): intercalary appendages; XIII (1 p.): head
segmentation; and XIV (1 p.): embryonic abdominal appendages and their fate.

The book is nicely printed on good quality paper and seems to be strongly bound
but is exorbitantly priced and has numerous typographical errors and a substantial
number of errors in interpretation. In his preface, Seidel hopes “...that the volume
will find wide distribution and that it will generate new developmental concepts for
experimental analysis” (p. xvi). At $ 261.00 (CAN) this is hardly possible since onlv
the most pecunious of libraries and individuals will be able to afford it. Also, much
of its content is better presented in other works that are readily available in most
libraries ( e.g ., Anderson, 1973. Embryology and Phylogeny in Annelids and
Arthropods, Pergamon Press; Counce and Waddington [eds.], 1972, 1973.
Developmental Systems: Insects, Vols. 1 & 2, Academic Press; Haget, 1977. Traite
de Zoologie, Vol. 8, part 5Ba: 1-262; 279-387. Masson et Cie; and Sander, Gutzeit
and Jackie, 1985. pp. 319-385 In Kerkut and Gilbert [eds.] Comparative Insect
Physiology, Biochemistry and Pharmacology. Vol. 1. Pergamon Press).

Below I note some of the more obvious errors in interpretation:

p. viii: insects do not constitute a phylum;

p. 14: the Devonian formations he refers to are 375^100 million years old not
350;

p. 20: the Monarch Butterfly, Danaus plexippus, is a new world species;

p. 46: Thysanoptera are not known to have nonparthenogenetic paedogenesis.
Rather males of the thripid, Limothr i£ lenticornis develop more rapidly than

females, are preferentially attracted to an ate with female pupae and lodge fully
differentiated spermatozoa in their still urn li shed spermathecae (Boumier, 1956.
Arch. Zool. exp. gen. 93: 219-317);

Tables I, III, V, X: numerous misplacements of genera or the laboratory species
of various authors into families and higher taxa;

p. 73: adult aphid macropterae usually develop in crowded conditions not the
opposite;

p. 230: the pleuropodia of the phasmid, Carausius develop from modifications in
the embryonic appendages of abdominal segment 1 ^ectodermal), as in many other
insects, not from secondary coelomic sacs (mesodermal) (Louvet. 1976. Int. J. Ins.
Embryol. Morphol. 5: 35-49);

Book Review

391

p. 238: Some neuroblasts rather than degenerating towards the end of
embryogenesis in holometabolous embryos, persist and divide teloblastically again
during larval development, eventually to generate large numbers of imaginal
neurons which differentiate during metamorphosis (Heming [1982] J. Morphol. 172:
23-43; Booker and Truman [1987] J. Comp. Neurol. 255: 547-559; Truman and
Bate [1988] Dev. Biol. 125: 145-157.);

p. 254: the description of mouthpart morphogenesis in thrips is derived from
Risler’s (1956) account of adult mouthparts in Thrips physapus. A full description
of their embryonic development in Haplothrips verbasci is presented by Heming
(1980. J. Morphol. 164: 235-263).

p. 255: labels for labial palpus and paraglossa are incorrect in Fig. 85c;

p. 257: salivary gland invaginations in Drosophila are here said to originate
between the labial and prothoracic segments; in Fig. 86c, they are shown as being
located between the maxillary and labial segments.

Finally, there are numerous inconsistencies in the use of subtitles and some of his
paragraphs go on for pages. Considering the above, one is better advised to purchase
one of the general references cited above.

B. S. Heming
Department of Entomology
University of Alberta

Quaest. Ent., 1989, 25 (3)

Quaest

lones

Entomolog

icae

A periodical record of entomological investigations,
published at the Department of Entomology,
University of Alberta, Edmonton, Canada.

VOLUME 25

NUMBER 4

FALL 1989

Publication of Quaestiones Entomologicae was started in 1965 as part of a
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Issued December 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 1989

CONTENTS

O'Hara — Systematics of the Genus Group Taxa of the Siphonini (Diptera:

Tachinidae) 1

Commentary 231

Book Review — Ives, W. G. H. and H. R. Wong. 1988. Tree and Shrub Insects

of the Prairie Provinces 239

Freitag and Barnes Classification of Brazilian Species of Cicindela and
Phylogeny and Biogeography of Subgenera Brasiella, Gaymara new
subgenus, Plectographa and South American Species of Cylindera

(Coleoptera: Cicindelidae) 241

Book Review — Schwalm, F.E. 1988. Insect Morphogenesis 387

Ward — Systematic studies on pseudomyrmecine ants: revision of the

Pseudomyrmex oculatus and P. subtilissimus species groups, with

taxonomic comments on other species 393

Book Review — Campbell, I. 1988. Zoological Catalogue of Australia. Vol. 6.

Ephemeroptera, Megaloptera, Odonata, Plecoptera, Trichoptera 469

Book Notice — Nichols, S. W. (Compiler), and Randall T. Schuh. 1989. The

Torre-Bueno Glossary of Entomology 471

Editor's Acknowledgements and Comments 473

i

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 4 1989

CONTENTS

Ward — Systematic studies on pseudomyrmecine ants: revision of the

Pseudomyrmex oculatus and P. subtilissimus species groups, with

taxonomic comments on other species 393

Book Review — Campbell, 1. 1988. Zoological Catalogue of Australia. Vol. 6.

Ephemeroptera, Megaloptera, Odonata, Plecoptera, Trichoptera 469

Book Notice — Nichols, S. W. (Compiler), and Randall T. Schuh. 1989. The

Torre-Bueno Glossary of Entomology 471

Editor's Acknowledgements and Comments 473

g

i'ehi-

1

SYSTEMATIC STUDIES ON PSEUDOMYRMECINE ANTS: REVISION OF
THE PSEUDOMYRMEX OCULATUS AND P. SUBTILISSIMUS SPECIES
GROUPS, WITH TAXONOMIC COMMENTS ON OTHER SPECIES

Philip S. Ward
Department of Entomology
University of California
Davis, California 95616
U. S. A.

Quaestiones Entomologicae
25:393-468 1989

ABSTRACT

The first part of this paper contains a synopsis of the major species groups and
revisions of two such groups (P. oculatus group, P. subtilissimus group) in the large
Neotropical ant genus Pseudomyrmex. Eleven species are recognized in the P.
oculatus group, of which three are new: P. alustratus Ward (from Peru), P. cretus
Ward ( Costa Rica), and P. pisinnus Ward (Brazil). P. caeciliae (Forel) is raised to
species ( stat. novj, and the following new synonymy is proposed: P. elongatus
(Mayr) = P. alliodorae (Wheeler); P. oculatus (F. Smith) = P. altinodus (Mann) = P.
tuberculatus ( Enzmann ) = P. wessoni (Enzmann); P. urbanus (F. Smith) = P. chodati
(Forel) = P. ogloblini (Santschi). Four species are recognized in the P. subtilissimus
group of which two are new: P. spiculus Ward (Costa Rica), and P. villosus Ward
(Brazil). P. culmicola (Forel) is proposed as a junior synonym of P. tenuissimus
(Emery). A final section of the paper contains taxonomic comments, including new
synonymy and lectotype designations, on 22 other species of Pseudomyrmex, in
various species groups, with emphasis on taxa which are associated with ant-plants.
No new species are described but three taxa [P. pazosi (Santschi), P. rufiventris
(Forel), and P. tachigaliae (Forel)] are raised to species (stat. novJ; two species [P.
dendroicus (Forel), P. osurus (Forel)] are removed from synonymy (stat. revalj; a
replacement name (P. laevifrons Ward, nom. novj is provided for P. laeviceps (F.
Smith), 1877 (nec F. Smith, 1859); and 32 new synonymies are proposed. Identities
of some of the Pseudomyrmex species inhabiting Acacia, Tachigali, and Triplaris
are clarified.

RESUME

La premiere partie de cet travail contient un sommaire des groupes principaux d’ especes et revisions de
deux tels groupes (groupe P. oculatus, groupe P. subtilissimus) dans le grand genre Neotropicaux,
Pseudomyrmex. Onze especes sont reconnues dans le groupe P. oculatus, desquelles trois sont nouvelles: P.
alustratus Ward ( de Peru), P. cretus Ward (Costa Rica), et P. pisinnus Ward (Brazil). P. caeciliae (Forel) est
eleve a I’etage d’espece (stat. novj et le nouveau synonymie suivant est propose: P. elongatus (Mayr) = P.
elongatus (Wheeler); P. oculatus (F. Smith ) = P. altinodus (Mann) = P. tuberculatus (Enzmann) = P.
wessoni (Enzmann); P. urbanus (F. Smith ) = P. chodati (Forel) = P. ogloblini (Santschi). Quatre especes

394

Ward

sont reconnues dans le groupe P. subtilissimus, desquelles deux sont nouvelles: P. spiculus Ward (Costa
Rica), et P. villosus Ward (Brazil). P. culmicola (Forel) est propose comme un nouveau synonyme de P.
tenuissimus (Emery). Une section finale contient notes taxonomiques, comprenant nouveau synonymie et
designation des lectotypes, sur 22 autres especes de Pseudomyrmex, dans quelques groupes d’ especes, avec
accent sur taxa qui sont associes avec des plantes myrmecophiles. On ne decrit pas nouvelles especes, mais
trois taxa [P. pazosi (Santschi), P. rufiventris (Forel), et P. tachigaliae (Forel)] sont eleves au niveau
d’espece fstat. novj; deux especes [P. dendroicus (Forel), P. osurus (Forel)] sont enleves de synonymie
fstat. revalj; un nom de remplacement (P. laevifrons Ward, nom. novj est fourni pour P. laeviceps (F.
Smith), 1877 (nec F. Smith, 1859); et 32 nouveaux synonymes sont proposes. Les identites de quelques
especes de Pseudomyrmex qui vivent dans Acacia, Tachigali et Triplaris sont clarifiees.

TABLE OF CONTENTS

Introduction 394

Materials and Methods 396

Collections 396

Metric Measurements and Indices 397

Other Conventions 401

Synopsis of Major Species Groups in Pseudomyrmex 402

Taxonomic Treatment of Pseudomyrmex oculatus and subtilissimus Groups 405

Pseudomyrmex oculatus Group 405

Introduction 405

Synonymic Lists of Species 406

Key to Species 407

Species Accounts 409

Pseudomyrmex subtilissimus Group 429

Introduction 429

Synonymic Lists of Species 429

Key to Species 430

Species Accounts 430

Taxonomic Comments on Other Pseudomyrmex Species 435

Introduction 435

Species Accounts 435

Summary of Taxonomic Changes 446

Concluding Remarks 447

Acknowledgements 448

Literature Cited 449

Figures 454

Index 463

INTRODUCTION

The New World ant genus Pseudomyrmex contains a large number of species
and is a conspicuous element of the Neotropical arboreal ant fauna. It is known from

Pseudomyrmecine Ants

395

southern United States (with northward extensions to about 40°N latitude on the east
and west coasts) south to approximately 36°S in Chile and Argentina. A closely
related Old World genus, Tetraponera, is distributed from Africa to southeastern
Asia and Australia. Much taxonomic work remains to be done on these ants, both in
terms of species diagnosis and phylogenetic evaluation. Within the genus
Pseudomyrmex there are more than 250 available species-level names, of which
about 70% are valid presently. Most of these names accumulated in a piece-meal
fashion in the late nineteenth and early twentieth centuries, without the benefit of
any synthetic, revisionary studies. As a first attempt to deal with the taxonomy of
this genus, Kempf (1958, 1960, 1961, 1967) delineated five species groups,
containing about half of the described taxa, and clarified the identities of some
species within these groups. The Nearctic species of Pseudomyrmex were reviewed
by Ward (1985), who recognized two additional species groups (P. elongatus group,
P. pallidus group).

The present paper contains a reevaluation and synopsis of the major species
groups, and revisions of two such groups: (i) the P. oculatus group, which is
expanded to include the “ elongatus group” of Ward (1985), and (ii) the P.
subtilissimus group, here defined and diagnosed for the first time. A fourth section
contains taxonomic comments about some of the remaining species of
Pseudomyrmex. It is based on a study of type material in European and North
American collections, and helps to provide a foundation for future revisionary work
on the other major species groups. Such work, currently in progress, will include
keys to species groups and to isolated species of uncertain placement, as well as
phylogenetic assessments of the relationships among groups. A preliminary analysis
of these relationships is presented in Ward (1990a). A generic revision of the
subfamily Pseudomyrmecinae and a higher-level phylogenetic analysis appears in
Ward (1990b).

The genus Pseudomyrmex is richly endowed with sympatric sibling species and
taxonomically confusing “species complexes”, a situation which is apparent even in
the depauperate Nearctic fauna (Ward, 1985). The resolution of such taxonomic
imbroglios requires careful analysis of character variation in large samples of
diverse geographic origins. Pseudomyrmex workers do not exhibit striking
interspecific variation in sculpture or spinescence; differences between closely
related species often reside in subtle contrasts of size and shape. I have attempted to
quantify these differences wherever practical and to avoid subjective qualitative
statements. But quantitative assessments of size and shape often reveal an
impressive amount of apparent intraspecific variation, making the search for
diagnostic differences challenging. Needless to say, precise metric measurements
are necessary for the accurate determination of many Pseudomyrmex species, and
the taxa dealt with in this paper are no exception.

Quaest. Ent., 1989, 25 (4)

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Ward

MATERIALS AND METHODS

Collections

Material was examined in the following collections (abbreviations follow those
of Arnett & Samuelson (1986) as closely as possible):

BMNH British Museum (Natural History), London, U.K.

CASC California Academy of Sciences, San Francisco, CA, USA
CDAE California Dept, of Food and Agriculture, Sacramento, CA, USA
CELM Coleccion Entomologfa “Luis Maria Murillo”, Instituto Colombiano
Agropecuario, Bogota, Colombia

CISC California Insect Survey, University of California at Berkeley, CA, USA
CKIC C. Kugler Collection, Radford University, VA, USA
CNCC Canadian National Collection of Insects, Biosystematics Research
Institute, Ottawa, Canada

CUIC Cornell University Insect Collection, Ithaca, NY, USA
EBCC Estacion de Biologia Chamela, Jalisco, Mexico
FFIC Fernando Fernandez Collection, Bogota, Colombia
GBFM Graham B. Fairchild Museo de Invertebrados, Universidad de Panama,
Panama

GCWC G. C. & J. Wheeler Collection, Silver Springs, FL, USA
GHPC G. H. Perrault Collection, Paris, France
ICCM Carnegie Museum of Natural History, Pittsburg, PA, USA
IEGG Istituto di Entomologia “Guido Grandi”, Universita de Bologna, Italy
INHS Illinois Natural History Survey Insect Collection, Champaign, IL, USA
INPA Instituto Nacional de Pesquisas da Amazonia, Manaus, Brazil
IZAV Instituto de Zoologia Agricola Universidad Central de Venezuela,
Maracay, Venezuela.

JTLC J. T. Longino Collection, University of California at Santa Barbara, CA,
USA

KSUC Kansas State University Insect Collection, Manhattan, KS, USA
KWJC K. W. Jaffe Collection, Universidad Simon Bolivar, Caracas, Venezuela
LACM Los Angeles County Museum of Natural History, Los Angeles, CA, USA
MCSN Museo Civico di Storia Naturale, Genoa, Italy

MCZC Museum of Comparative Zoology, Harvard University, Cambridge, MA,
USA

MHNG Museum d’Historie Naturelle, Geneva, Switzerland
MJPL Museo de Historia Natural “Javier Prado”, Lima, Peru
MNCR Museo Nacional de Costa Rica, San Jose, Costa Rica
MNHN Museum National d’Histoire Naturelle, Paris, France
MZSP Museo de Zoologia da Universidade de Sao Paulo, Brazil
NHMB Naturhistorisches Museum, Basel, Switzerland
NHMV Naturhistorisches Museum, Vienna, Austria

Pseudomyrmecine Ants

397

NHRS Naturhistoriska Riksmuseet, Stockholm, Sweden

OSUO Department of Entomology Collection, Oregon State University,
Corvallis, OR, USA

PSWC P. S. Ward Collection, University of California at Davis, CA, USA

SEMC Snow Entomological Museum, University of Kansas, Lawrence, KS,
USA

TAMU Department of Entomology Insect Collection, Texas A&M University,
College Station, TX, USA

UCDC Bohart Museum of Entomology, University of California at Davis, CA,
USA

UCRC UCR Entomological Collection, University of California at Riverside,
CA, USA

UNCB Museo de Historia Natural, Instituto de Ciencias Naturales, Universidad
Nacional de Colombia, Bogota, Colombia

USNM National Museum of Natural History, Washington, DC, USA

UWEM Entomology Museum, University of Wisconsin, Madison, USA

WPMC W. P. MacKay Collection, College Station, TX, USA

WWBC W. W. Benson Collection, Campinas, SP, Brazil

ZMHB Zoologisches Museum, Museum fur Naturkunde der Humboldt -
Universitat, Berlin, East Germany

ZMUC Zoologisk Museum, University of Copenhagen, Dennmark

ZMUH Zoologisches Institut und Zoologisches Museum, Universitat Hamburg,
West Germany

ZSMC Zoologische Sammlung des Bayerischen Staates, Munich, West Germany

Metric Measurements and Indices

Metric measurements were made at 50X power on a Wild microscope, with a
dual-axis (X-Y) pair of Nikon micrometers wired to an Autometronics digital
readout. All measurements were taken to the nearest thousandth of a millimeter.
Most have been rounded to two decimal places for presentation here.

Some of the measurements described below differ from those used in taxonomic
studies of other ants. These modifications are partly dictated by peculiarities of the
Pseudomyrmecinae. For example, following Kempf (1960), the compound eyes are
included in the measurement of head width in workers and queens (as well as
males), because the maximum width of the head capsule frequently occurs behind
the eyes and is blocked from view. In addition I use DPL (diagonal length of the
propodeum; see below) as a proportional measure of mesosoma (=alitrunk) length
rather than WL (Weber’s length of the entire mesosoma) because the free
articulation of the pronotum with the rest of the thorax renders the measurement of
WL imprecise.

A number of important measurements are taken with the head in a full-face,
frontal (i.e., dorsal) view. To make such measurements comparable among all

Quaest. Ent., 1989, 25 (4)

398

Ward

pseudomyrmecine ants, the head is positioned so that its anterolateral margins
(above the mandibular insertions) are in the same focal plane as the occipital margin
(see Fig. 6). The head length (HL) which is measured in such a view includes the
median clypeal lobe, but the axis of measurement is not unduly altered by a strong
anterodorsal protrusion of the fronto-clypeal complex as occurs, for example, in
some Tetraponera. As a result, HL is slightly less than the “maximum critical” head
length (Brown, 1953), but it and other measurements which are taken in a full-face
view of the head are more appropriate for comparisons across the entire subfamily.

For Pseudomyrmex workers and queens a full-face, frontal view of the head is
roughly equivalent to positioning the median ocellus and the frontal carinae in the
same plane of view (while turning the head so that the maximum possible head
width is achieved). Such a positioning rule does not usually work in Tetraponera ,
due to discrepancies in head shape and the frequent lack of ocelli in workers.

In discussions below, the antennal sclerite is the sclerite which encircles the
antennal fossa. In most ants, the inner margin of the antennal sclerite is developed as
a median lobe (Fig. 3) which is raised dorsolaterally and often covers part of the
basal condyle of the antenna. In pseudomyrmecine ants these san e median lobes
typically fuse with the frontal carinae anteromedially (Fig. 3); in many other ants the
frontal carinae are themselves expanded laterally to form frontal lobes which cover
the median lobes of the antennal sclerites.

The following is a list of measurements used in this and other continuing studies
of pseudomyrmecine ants. Explanatory illustrations of some of these measurements
are found in Ward (1985); see also Figs. 1-5 of this paper.

HW

VW

HL

EL

OD

OOD

Head width: maximum width of head, including the eyes, measured in
full-face dorsal view.

Vertex width: width of the posterior portion of the head (vertex),
measured along a line drawn through the lateral ocelli, with the head in
full-face, dorsal view (see Fig. 1 in Ward [1985]).

Head length: midline length of head proper, measured in full-face, dorsal
view, from the anterior clypeal margin to the midpoint of a line drawn
across the occipital margin.

Eye length: length of compound eye, measured with the head in full-face,
dorsal view.

Ocellar distance: distance from the middle of the median ocellus to the
midpoint of a line drawn between the lateral ocelli, measured with the
head in full-face dorsal view (see Fig. 1 in Ward [1985]).

Oculo-ocellar distance: distance from the middle of the median ocellus to
the midpoint of a line drawn across the posterior margins of the
compound eyes, measured with the head in full-face, dorsal view (see
Fig. 1 in Ward [1985]) (this distance is negative in value if the posterior
margin of the compound eye exceeds the median ocellus).

Pseudomyrmecine Ants

399

CD Clypeal distance: distance from the anterior clypeal margin to the
midpoint of a line drawn across the anterior margins of the antennal
sclerites.

MD1-

MD9 A series of mandibular measurements (see Figs. 1, 2) taken with the
mandibles removed from their insertions and mounted separately on a
point. For measurements of width and overall length (MD1 - MD3), the
mandible is positioned so that the dorsal abductor swelling and the apical
tooth are in the same focal plane (Fig. 2) and it is then rotated on this axis
until MD2, the width of the mandible at the juncture of the basal and
masticatory margins, is made maximum. Measurements of the lengths of
the basal and masticatory margins, and of the relative positioning of teeth
along those margins (MD4 - MD9), are taken with the extreme ends of
the respective margin in the same focal plane. If there is more than one
mesial basal tooth, MD4 is the distance to the outermost one.

MFC Minimum frontal carinal distance: minimum distance between the frontal
carinae, posterior to their fusion with, or approximation to, the antennal
sclerites (Fig. 4).

ASD Antennal sclerite distance: maximum distance between the lateral
margins of the median lobes of the antennal sclerites, measured in
full-face, dorsal view of the head (Fig. 4).

ASO Antennal sclerite distance, outer margins: maximum distance between the
outer, lateral margins of the antennal sclerites (Fig. 4).

EW Eye width: maximum width of compound eye, measured along its short
axis, in an oblique dorso-lateral view of the head.

EL2 Eye length: maximum length of compound eye, measured along its long
axis in the same plane of view as EW.

SL Scape length: length of the first antennal segment, excluding the radicle.

LF1 Length of first funicular segment: maximum measurable length of the
first funicular segment (pedicel), including its basal articulation in
workers and queens but excluding the basal articulation in males (where
it is usually hidden).

LF2 Length of second funicular segment: maximum measurable length of the
second funicular segment.

LF3 Length of third funicular segment: maximum measurable length of the
third funicular segment.

WF2 Width of second funicular segment.

FL Fore femur length: length of the fore femur, measured along its long axis
in posterior view (see Fig. 3 in Ward [1985]).

FW Fore femur width: maximum measurable width of the fore femur,
measured from the same view as FL, at right angles to the line of
measurement of FL.

Quaest. Ent., 1989, 25 (4)

400 Ward

DPL Diagonal length of the propodeum: length of the propodeum, measured in
lateral view along a diagonal line drawn from the “metapleural” lobe to
the metanotal groove (see Fig. 2 in Ward (1985)).

BF Length of the basal (=dorsal) face of the propodeum, measured in lateral
view from the metanotal groove to the point on the surface of the
propodeum which is maximally distant from the diagonal propodeal line.

DF Length of the declivitous face of the propodeum, measured in lateral view
from the metapleural lobe to the point on the surface of the propodeum
which is maximally distant from the diagonal propodeal line.

MP Depth of metanotal groove (“mesopropodeal impression”), measured in
lateral view from the bottom of the metanotal groove to a line drawn
across the dorsal surface of the mesonotum and propodeum.

PL Petiole length: length of the petiole, measured in lateral view from the
lateral flanges of the anterior peduncle to the posterior margin of the
petiole (see Fig. 4 in Ward [1985]).

PND Petiolar node distance: distance from the lateral flanges of the anterior
petiolar peduncle to the maximum height of the node, measured from the
same view as PL and along the same line of measurement (see Fig. 4 in
Ward [1985]).

PH Petiole height: maximum height of the petiole, measured in lateral view
at right angles to PL, but excluding the anteroventral process.

PPL Postpetiole length: length of the postpetiole, measured in lateral view,
from the anterior peduncle (of the postpetiole) to the point of contact with
the fourth abdominal tergite, excluding the pretergite (see Fig. 4 in Ward
[1985]).

DPW Dorsal petiolar width: maximum width of the petiole, measured in dorsal
view.

MPW Minimum petiolar width: minimum width of the petiole, measured in
dorsal view, anterior to DPW.

PPW Dorsal postpetiolar width: maximum width of the ppstpetiole, measured
in dorsal view.

LHT Length of hind tibia: maximum measurable length of hind tibia,

excluding the proximal part of the articulation which is received into the
distal end of the hind femur (Fig. 5).

LHS Length of hind basitarsus: maximum measurable length of hind

basitarsus.

Indices calculated from the preceding measurements include the following ratios
(the orthodox but redundant and potentially misleading procedure of multiplying
ratios by 100 has not been adopted):

Pseudomyrmecine Ants

401

Cl Cephalic index: HW/HL

OI Ocular index: EW/EL

012 Ocular index, using EL2: EW/EL2

REL Relative eye length: EL/HL

REL2 Relative eye length, using HW: EL/HW

OOI Oculo-ocellar index: OOD/OD

VI Vertex width index: VW/HW

FCI Frontal carinal index: MFC/HW

FCI2 Frontal carinal index, using ASD: MFC/ASD

ASI Antennal sclerite index: ASD/ASO

SI Scape index: SL/HW

SI2 Scape index, using EL: SL/EL

FLI Funicular length index: (LF2 + LF3)/WF2

FI Fore femur index: FW/FL

PDI Propodeal index: BF/DF

MPI Metanotal index: MP/HW

NI Petiole node index: PND/PL

PLI Petiole length index: PH/PL

PLI2 Petiole length index, using PPL: PPL/PL

PWI Petiole width index: DPW/PL

PWI2 Petiole width index, using PPW: DPW/PPW

PWI3 Petiole width index, using MPW: MPW/DPW

PPWI Postpetiole width index: PPW/PPL

PPWI2 Postpetiole width index, using HW: PPW/HW

Other Conventions

Palp formula refers to the number of maxillary palp segments, followed by the
number of labial palp segments. In counting the numbers of teeth on the basal and
masticatory margins of the mandibles, the apico-basal tooth (see Fig. 1) is
considered to be part of the dentition of the masticatory margin. The following setal
counts are employed:

MSC Mesosoma setal count: number of standing hairs, i.e., those forming an
angle of 45° or more with the cuticular surface (Wilson, 1955), visible in
outline on the mesosoma dorsum in lateral view.

HTC Hind tibial setal count: number of standing hairs visible in outline on the
outer (extensor) surface of the hind tibia, with the line of view orthogonal
to the plane of tibial flexion.

MTC Mid tibial setal count: equivalent count for mid tibia.

Terms for surface sculpture follow Harris (1979). When material was examined
under a light microscope, an opaque (Mylar) filter was placed between the source of

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402

Ward

illumination and the specimen, at a distance of about 4 cm from the latter. The
consequent reduction of glare renders the details of sculpture more visible, and
permits greater discrimination between different degrees of reflectance of the
integument. The terms opaque, subopaque, sublucid, and lucid are used to make
distinctions along this continuum of reflectance, opaque indicating a lack of
reflectance, on the one hand, and lucid referring to a strongly shining integument, on
the other hand, under the conditions of soft lighting described above.

Scanning electron micrographs were taken with an ETEC Autoscan, using
gold-palladium coated specimens.

The term stat. nov. is used for a change in status of a valid name (in all instances
here: elevation from infraspecific to specific status), while stat. reval. refers to the
revalidation of a name previously considered invalid (in all instances here:
resurrection of a former synonym). The distinction between these two kinds of name
changes was not made in Ward (1985).

In the lists of material examined, records are arranged alphabetically by country
and by principal administrative region within larger countries. The abbreviations for
Brazilian states follow Kempf (1972), with the following additions: MS = Mato
Grosso do Sul; RR = Roraima.

The abbreviation “c.u.” signifies collector unknown.

SYNOPSIS OF MAJOR SPECIES GROUPS IN PSEUDOMYRMEX

Among ants whose workers possess a distinct postpetiole and well-developed
sting, those of Pseudomyrmex are recognized by their elongate compound eyes (OI
0.48-0.66, REL 0.36-0.68), closely set frontal carinae (FCI 0.01-0.13), and short
scapes (SI 0.40-0.52). The median lobes of the antennal sclerites are also closely
adjacent (ASI 0.41-0.73) and visible in a full-face view of the head. The antennae
are 12-segmented. The mandibles are relatively small, with well differentiated basal
and masticatory margins bearing 2-3 and 5-10 teeth, respectively.

Within the genus Pseudomyrmex I recognize currently nine major species
groups, containing about 85% of the described taxa. The most salient worker
characteristics of these groups are summarized in Table 1.

The outline of species groups is tentative, details of which undoubtedly will
undergo refinement. The pallens group, in particular, appears to be rather
heterogeneous and is probably paraphyletic. Additional undescribed species are in
most of these groups; and there is a residue of taxonomically isolated species, and
taxa of uncertain identity ( incertae sedis ). Nevertheless Table 1 should permit the
assignment of most worker-based material of Pseudomyrmex to a species group.

The valid described taxa in each group are given below. All names are implicitly
listed as binomina (since I expect all infraspecific names eventually either to enter
synonymy or be raised to species), and the nomenclatural changes of this paper have
been incorporated.

Species Palp No. teeth on Lateral comers of median Pilosity on mesosoma Tibial pilosity: common Eye size: relatively Other

group formula(e) masticatory clypeal lobe: rounded dorsum: common ([HTC + MTC] >10) short (REL 0.36-0.46)

margin of (or bluntly angled) (MSC>10) versus versus sparse versus elongate

mandibles3 versus sharply angulate sparse (MSC<8) ([HTC + MTC]<8) (REL 0.48-0.68)

Pseudomyrmecine Ants

403

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Quaest. Ent., 1989, 25 (4)

404

Ward

P. ferrugineus group

alfari (Forel), atrox (Forel), bicinctus (Santschi), ferrugineus (F. Smith),
flavicornis (F. Smith), gaigei (Forel), infernalis (Wheeler), nigrocinctus (Emery),
peltatus (Menozzi), peperi (Forel), scelerosus (Wheeler), spinicola (Emery),
veneficus (Wheeler).

P. gracilis group

alternans (Santschi), alvarengai Kempf, argentinus (Santschi), atrinodus
(Santschi), atripes (F. Smith), bicolor (Guerin), browni Kempf, canescens (F.
Smith), cladoicus (F. Smith), dimidiatus (Roger), excisus (Mayr), faber (F. Smith),
glabriventris (Santschi), godmani (Forel), gracilis (Fabricius), laevigatus (F. Smith),
maculatus (F. Smith), major (Forel), mexicanus (Roger), niger (Donisthorpe),
nigropilosus (Emery), osurus (Forel), pilosulus (F. Smith), pupa (Forel), sericatus
(F. Smith), simulans Kempf, squamiferus (Emery), unicolor (F. Smith), velifer
(Stitz), venustus (F. Smith), volatilis (F. Smith).

P. oculatus group

alustratus Ward, caeciliae (Forel), cretus Ward, cubaensis (Forel), curacaensis
(Forel), eduardi (Forel), elongatus (Mayr), oculatus (F. Smith), pisinnus Ward,
schuppi (Forel), urbanus (F. Smith).

P. pallens group

adustus (Borgmeier), apache Creighton, championi (Forel), elongatulus (Dalle
Torre), gibbinotus (Forel), incurrens (Forel), landolti (Forel), longus (Forel),
lynceus (Spinola), monochrous (Dalle Torre), pallens (Mayr), phyllophilus (F.
Smith), salvini (Forel), thoracicus (Norton).

P. pallidas group

acanthobius (Emery), belgranoi (Santschi), brunneus (F. Smith), cocae
(Santschi), colei (Enzmann), distinctus (F. Smith), ejectus (F. Smith), ethicus
(Forel), euryblemma (Forel), fiebrigi (Forel), flavidulus (F. Smith), fuscatus
(Emery), holmgreni (Wheeler), leptosus Ward, levivertex (Forel), lizeri (Santschi),
pallidus (F. Smith), pazosi (Santschi), peruvianus (Wheeler), rochai (Forel),
rufiventris (Forel), seminole Ward, simplex (F. Smith), solisi (Santschi), terminalis
(F. Smith), virgo (Santschi), vistanus (Enzmann).

P. sericeus group

acaciarum (Wheeler), beccarii (Menozzi), castus (Wheeler), cordiae (Forel),
fords (Forel), heterogynus (Wheeler & Mann), huberi (Santschi), humboldi
(Enzmann), ita (Forel), lisus (Enzmann), longior (Forel), pictus (Stitz), rubiginosus
(Stitz), sericeus (Mayr), vinneni (Forel).

Pseudomyrmecine Ants

405

P. subtilissimus group

spiculus Ward, subtilissimus (Emery), tenuissimus (Emery), villosus Ward.

P. tenuis group

boopis (Roger), denticollis (Emery), tenuis (Fabricius), termitarius (F. Smith).

P. viduus group

concolor (F. Smith), cordobensis (Forel), coronatus (Wheeler), dendroicus
(Forel), endophytus (Forel), kuenckeli (Emery), malignus (Wheeler), nigrescens
(Forel), opacior (Forel), rurrenabaquensis (Wheeler & Mann), symbioticus (Forel),
tachigaliae (Forel), triplaridis (Forel), triplarinus (Weddell), viduus (F. Smith).

Incertae sedis

antiguanus (Enzmann), brunnipes (Enzmann), costaricensis (Enzmann),
depressus (Forel), duckei (Forel), fervidus (F. Smith), filiformis (Fabricius), gebelli
(Forel), goeldii (Forel), haytianus (Forel), laevifrons Ward, mandibularis (Spinola),
oki (Forel), perboscii (Guerin), rufomedius (F. Smith), santschii (Enzmann), subater
(Wheeler & Mann), voytowskii (Enzmann), weberi (Enzmann), wheeleri (Enzmann).

TAXONOMIC TREATMENT OF PSEUDOMYRMEX OCULATUS AND
SUBTILISSIMUS GROUPS

PSEUDOMYRMEX OCULATUS GROUP

Introduction

Diagnosis (worker). — Small to medium-size species (HW 0.47-0.98), head
longer than broad, often markedly so (Cl 0.61-0.88), with medium to large eyes
(REL 0.48-0.61); mandibles with external and basal margins parallel or diverging
slightly (MD1/MD2 0.87-1.00); basal margin of mandibles with a single distal
tooth, masticatory margin with five or (less commonly) six teeth; frontal carinae
subcontiguous (FCI 0.017-0.047), fusing anterolaterally with the antennal sclerites,
so that in lateral view the frontal carinae do not form a continuous curve with the
clypeus (Fig. 17) (in contrast to the P. subtilissimus group, see below and Fig. 19);
median lobe of clypeus tectiform, notably produced anteriorly, the anterior margin
sharp-edged and weakly flared (Figs. 14-16); in dorsal view anterior margin of
median clypeal lobe convex and laterally rounded; petiolar node relatively short and
high (PLI 0.67-1.06), anterior peduncle not conspicuously developed. Head and
mesosoma at least partly punctate, and usually with substantial areas of the
integument opaque or subopaque. Erect pilosity conspicuous on most parts of body,
including the antennae, legs, head, and dorsum of mesosoma. Body covered with
light to moderate density of fine pubescence. Palpal formula: 6,3 (reduced to 5,3 in
P. eduardi, P. pisinnus, and in some individuals of P. caeciliae and P. urbanus ;
these species are among the smallest in the group).

Quaest. Ent., 1989, 25 (4)

406

Ward

Comments. — The present concept of the P. oculatus group is a much expanded
one from that given in Kempf (1961), who treated only three species. Within the P.
oculatus group, I have included also the cluster of species allied to P. elongatus
(Mayr) (the “ elongatus group” of Ward [1985]). As presently defined, the P.
oculatus group is minimally diagnosed by the following combination of worker
features: tectiform clypeal lobe, with weakly flared, sharp-edged, anterior margin
(Figs. 14-16); relatively short, high, apendunculate petiole ( e.g ., Figs. 7, 8, 9b, 10b);
and conspicuous standing pilosity. Relationships with the P. subtilissimus group are
discussed under the latter.

Within the P. oculatus group are several reasonably distinctive species, but a
bewildering residue of forms remains centering around P. caeciliae (Forel), P.
cubaensis (Forel) and P. urbanus (F. Smith). Species relationships within what may
be called the urbanus complex have not been resolved fully in this study, and will
require more extensive material and further analysis (particularly larger samples of
worker-associated males and queens).

Synonymic List of Species

P. alustratus Ward, sp. nov. Peru
P. caeciliae (Forel, 1913), stat. nov. Texas to Panama
P. cretus Ward, sp. nov. Guatemala, Costa Rica
P. cubaensis (Forel, 1901b). Caribbean; Mexico to Bolivia
P. curacaensis (Forel, 1912). Panama to Bolivia, Brazil
P . eduardi (Forel, 1912). Jamaica; Costa Rica to Peru, Brazil
P. elongatus (Mayr, 1870). Florida; Mexico to Bolivia, Brazil
= P. alliodorae (Wheeler, 1942), syn. nov.

= P. allidorus (Enzmann, 1945).

= P. tandem (Forel, 1906).

P. oculatus (F. Smith, 1855). Mexico to Bolivia, Brazil
= P. advena (F. Smith, 1855).

= P. dolichopsis (Forel, 1899).

= P. implicatus (Forel, 1911).

= P. altinodus (Mann, 1916), syn. nov.

= P. tuberculatus (Enzmann, 1945), syn. nov.

= P. wessoni (Enzmann, 1945), syn. nov.

P. pisinnus Ward, sp. nov. Brazil, Peru
P. schuppi (Forel, 1901a). Argentina, Paraguay, Brazil
= P. confusior (Forel, 1901a).

= P. geraensis (Forel, 1912).

P. urbanus (F. Smith, 1877). Panama, South America
= P. chodati (Forel, 1920), syn. nov.

= P. ogloblini (Santschi, 1936), syn. nov.

Pseudomyrmecine Ants

407

Key to Species (workers and queens)

Note. — The queen caste is included tentatively in this key. To underscore the
rather limited amount of material on which this is based, I have given sample sizes
(n = ...) for queen measurements. Sample sizes within single lugs of a couplet are all
identical, and are not repeated after the first metric measurement.

1 Petiolar node broadly rounded in lateral profile, without
strongly differentiated dorsal and posterior faces (e.g., Figs.

8,9b); summit of the node usually occurring in the middle third

of the petiole length (NI 0.51-0.69) 2

1' Petiolar node with a flat, posteriorly uplifted, dorsal face,

rounding suddenly into a vertical posterior face (Fig. 7);
summit of node displaced to the posterior quarter of the petiole
length (NI 0.72-0.85); medium-sized species (worker HW
0.73-0.91; queen HW 0.78-0.85, n=7) with broad head (worker
Cl 0.77-0.87; queen Cl 0.71-0.76) and long eyes (worker REL
0.55-0.62; queen REL 0.53-0.57) (Mexico to Bolivia and
Brazil) P. oculatus (F. Smith), p. 422

2 (1) Larger species, worker HW 0.74-0.98, usually greater than

0.79 (queen HW 0.78-0.95, n=15); if HW <0.79mm, then
occipital margin of head concave (in frontal view) and
dorsolateral margination of petiole relatively sharp so that in
dorsal view the maximum petiole width appears to coincide
with the lateral margination; funicular segments II and III
ranging from slightly broader than long to notably longer
than broad 3

2' Smaller species, HW <0.79; occipital margin of worker flat to

weakly concave (in frontal view); dorsolateral margination of
petiole rather gentle so that in dorsal view the maximum petiole
width occurs slightly below the margination; funicular
segments II and III broader than long, usually markedly so 5

3 (2) Petiole about as high as long in worker (Fig. 8) (worker PLI

0.94-1 .07) and appearing about as broad as long in dorsal view
(PWI 0.84-0.99) (in queens: PLI 0.81-0.85, PWI 0.78-0.85,
n=4); head densely punctate, opaque (Argentina, Paraguay, s.e.

Brazil) P. schuppi (Forel), p. 426

3' Petiole longer than high (Figs. 9b, 10b) (worker PLI 0.71-0.89;
queen PLI 0.57-0.76, n=ll) and notably longer than broad in
dorsal view (worker PWI 0.59-0.78; queen PWI 0.61-0.75);
head punctate and sublucid, with small shiny interspaces
between many of the punctures 4

4 (3') Larger species (worker HW 0.85-0.98; queen HW 0.90-0.95,

Quaest. Ent., 1989, 25 (4)

408

Ward

5 (2')
5'

6 (5)
6'

7 (5')
T

n=3) with longer scapes and shorter eyes (worker SI2
0.74-0.80; queen SI2 0.69-0.73; worker REL2 0.62-0.65;
queen REL2 0.68-0.71) (Fig. 9a); regressions of HL on HW
and EL on HW lying below those of curacaensis (Figs. 26, 27)

(Guatemala, Costa Rica) P. cretus, sp. nov., p. 412

Smaller species (worker HW 0.74-0.89; queen HW 0.78-0.88,
n=8) with shorter scapes and relatively longer eyes (worker SI2
0.60-0.74; queen SI2 0.58-0.66; worker REL2 0.64-0.77;
queen REL2 0.72-0.82) (Fig. 10a); regressions of HL and EL
on HW lying above those of cretus (Figs. 26,27) (Panama south

to Bolivia and Brazil) P. curacaensis (Forel), p. 416

Punctures on dorsum of worker head and pronotum contiguous
almost throughout, rendering the integument largely opaque; in
particular, punctures mostly contiguous and without extensive
shiny interspaces (i.e., separated by half their diameters or less)
posteromesad of compound eye, and immediately posterior to
compound eye (except for a thin shiny margin about
0.010-0.020 mm. wide), when viewed under a light microscope
(see also SEM micrographs, Figs. 20, 21); queens similar but
sometimes with slightly more extensive shiny interspaces

between ocelli and compound eye 6

Dorsum of head and pronotum more extensively shiny or
sublucid, especially in queens; at least some punctures
separated by half their diameters or more, especially between
the ocelli and compound eye, and immediately posterior to
compound eye, when viewed under a light microscope (see also

SEM micrographs, Figs. 22-25) 7

Smaller species (worker HW 0.56-0.68, usually <0.65; queen
HW 0.56-0.61, n=19; worker HL 0.73-0.93; queen HL

0.87-0.97) (Florida; Mexico south to Bolivia and Brazil)

P. elongatus (Mayr), p. 419

Larger species (worker HW 0.69-0.75; queen HW = 0.67, m=l;

worker HL 0.96-1.04; queen HL = 1.05) (Peru)

P. alustratus, sp. nov., p. 409

Very small species (worker HW 0.47-0.51, queen unknown);
worker with elongate head (Cl 0.61-0.68), short scapes (SI2
0.50-0.54), long basal face of propodeum (PDI 1.46-1.69), and
low petiole (PLI 0.67-0.77) (Fig. 11); light orange-brown

(Brazil, Peru) P. pisinnus, sp. nov., p. 425

Larger species (worker HW 0.50-0.78) with broader head
(worker Cl 0.67-0.79) and longer scapes (worker SI2
0.55-0.74); at lower end of worker size range (HW 0.50-0.60),

Pseudomyrmecine Ants

409

basal face of propodeum shorter (PDI 0.91-1.45) and petiole
higher (PLI 0.79-0.92); color variable, often medium to
dark brown 8

8 (7') Front of head smooth and shiny with scattered fine punctulae

(Fig. 25); small (worker HW 0.50-0.57; queen HW 0.51-0.55,
n=3), dark brown, shiny species (Jamaica; Costa Rica to Peru

and Brazil) P. eduardi (Forel), p. 418

8' Front of head shiny to subopaque, with coarser (0.010 to

0.020 mm in diameter) and denser punctures, separated by
0-4 diameters ( e.g Fig. 24); size and color variable
(i urbanus complex) 9

9 (8') Larger species (worker HW 0.65-0.78; queen HW 0.65-0.76,

n=17); at lower end of worker size range, fore femur is
relatively slender and eyes relatively short (see Figs. 28,29)

(Florida, West Indies, Mexico south to Brazil)

P. cubaensis (Forel) ( s.l .), p. 413

9' Smaller species (worker HW 0.51-0.66; queen HW 0.51-0.64,

n=9); at upper end of worker size range, fore femur tends to be

broader and eyes longer (Figs. 28,29) 10

10 (9') Larger species (worker HW 0.58-0.66; queen HW 0.58-0.64,
n=3); ocellar distance (OD) generally smaller (worker OD/HW
0.103-0.149) (see Fig. 30); usually light orange-brown in color,

sometimes darker (Panama, South America)

P. urbanus (F. Smith), p. 427

10' Smaller species (HW 0.51-0.61, usually < 0.59; queen HW

0.51-0.59, n=6), with a larger ocellar distance (worker OD/HW
0.127-0.169) (see Fig. 30); medium to dark brown in color
(Texas south to Panama) P. caeciliae (Forel), p. 410

Species Accounts

Pseudomyrmex alustratus sp. nov.

(Figures 13a, 13b)

Holotype worker. — PERU, Junfn: Chanchamayo, Anashirone River, 16.vii. 1963,
C.A. Portocarrero #A47 (Kempf No. 4060) (MZSP). HW 0.73, HL 1.03, EL 0.59,
PL 0.42, PH 0.35.

Paratypes. — Five workers, same data as holotype (BMNH, LACM, MCZC,
MZSP). Seven workers, one dealate queen, same locality as holotype, 22.viii.1963,
C.A. Portocarrero #73 (Kempf No. 4072) (MJPL, MZSP, PSWC).

Additional non-paratypic material is listed below.

Quaest. Ent., 1989, 25 (4)

410

Ward

Worker measurements (n=9). — HL 0.96-1.04, HW 0.69-0.75, MFC 0.018-0.024, Cl
0.70-0.76, 01 0.55-0.58, REL 0.55-0.58, REL2 0.76-0.80, OOI 0.13-0.53, VI 0.80-0.86, FCI 0.024-0.034,
SI 0.45-0.46, SI2 0.58-0.60, FI 0.47-0.53, PDI 1.32-1 58, MPI 0.040-0.055, NI 0.59-0.64, PLI 0.83-0.92,
PWI 0.63-0.7 1,PPWI 1.18-1.43.

Worker diagnosis. — Medium-sized species (HW 0.69-0.75), with elongate head and eyes (Fig.
13a; Cl 0.70-0.76, REL2 0.76-0.80); sides of head slightly convex, occipital margin weakly concave in
full-face view; second and third funicular segments broader than long (FLI 1.49-1.69, n=5). Fore femur
relatively broad; metanotal groove of moderate depth but not long; basal face of propodeum flat to slightly
convex, not notably raised above level of mesonotum, longer than the declivitous face into which it rounds
rather suddenly (the basal and declivitous faces of the propodeum form a distinct obtuse angle in some
individuals). Petiolar node relatively short and high (Fig. 13b; see metrics), with a conspicuous, rounded
subpetiolar process; postpetiole broader than long.

Mandibles striate with scattered punctures. Head and pronotum densely punctate and opaque (weakly
sublucid in the centre of the pronotum); remainder of mesosoma punctate or coriarious-imbricate, and
subopaque. Petiole, postpetiole, and gaster subopaque to weakly shining, with numerous fine piligerous
punctures. Fine erect pilosity and appressed pubescence common over most of the body, including the
mesosoma dorsum. Medium brown, apices of appendages (antennae, legs) lighter in color; mandibles pale
luteous.

Comments. — This species evidently is related closely to P. elongatus, from
which it differs primarily in size (HW >0.68). The head is even more densely
punctate and opaque than in most P. elongatus workers, the basal face of the
propodeum tends to be relatively longer (compare PDI values), and the subpetiolar
process is perhaps a little better developed than in P. elongatus. The punctate
opaque head, elongate eyes (REL2 >0.76), relatively broad fore femur, and high,
rounded petiole (PLI >0.82), distinguish P. alustratus from P. cubaensis, the only
species with which it overlaps substantially in size.

Biology. — Essentially nothing is known about the biology of P. alustratus. Two
workers from Sotileja, Peru were foraging on Ochroma.

Material Examined. — Type material listed above plus the following: PERU Junm:
Valle Chanchamayo, 800m (Weyrauch) (MZSP); Madre de Dios: Sotileja, jet. Rio Manu & R. Sotileja
(D.W. Davidson) (PSWC).

Pseudomyrmex caeciliae (Forel), stat. nov.

(Figures 23, 24, 28, 29, 30)

Pseudomyrma eduardi race caeciliae Forel, 1913: 214. Two syntype workers, Patulul, Guatemala (Peper)

(MHNG) [Examined]. One worker here designated LECTOTYPE.

Worker measurements (n=26). — HL 0.73-0.91, hw 0.51-0.61, MFC 0.012-0.023, ci
0.67-0.74, OI 0.52-0.60, REL 0.48-0.55, REL2 0.72-0.81, OOI 0.22-0.80, VI 0.73-0.91, FCI 0.021-0.041,
SI 0.42-0.49, SI2 0.55-0.63, FI 0.45-0.51, PDI 0.91-1.45, MPI 0.048-0.084, NI 0.55-0.63, PLI 0.79-0.91,
PWI 0.61-0.74, PPWI 1.09-1.39.

Worker diagnosis. — Small species (HW 0.51-0.61) with elongate eyes (REL2 0.72-0.81); ocelli
relatively well-separated, OD/HW >0.127; sides of head subparallel, rounding into the occipital margin, the
latter flat to weakly concave, in full-face view; second and third funicular segments broader than long,
usually notably so (FLI 1.23-1.72, n=5). Fore femur relatively broad; metanotal groove well marked; basal
face of propodeum flat to weakly convex, often raised perceptibly above the level of the mesonotum; basal
face rounding into declivitous face of propodeum, to which it is subequal or slightly greater in length (see
PDI values). Petiole relatively short and high (PLI >0.79), with an anteroventral process, variously
developed; postpetiole of typical dimensions, broader than long.

Pseudomyrmecine Ants

41

Mandibles striate (sometimes weakly so), with scattered punctures. Head densely punctate, the
punctures separated by shiny interspaces at least on parts of the upper third of the head, and especially in the
area immediately posterior to the compound eye, so that these parts of the head appear sublucid to shiny (in
contrast to the predominantly opaque head of P. elongatus)', head punctures (Figs. 23, 24) relatively coarse,
0.010 to 0.020 mm in diameter; mesosoma punctate to coriarious-imbricate, sublucid in the centre of the
pronotal and mesonotal discs, and on the side of the pronotum, more opaque elsewhere; petiole, postpetiole,
and gaster sublucid, with numerous fine piligerous punctures. Fine, erect pilosity (sometimes rather short)
and appressed pubescence common over most of the body, including the mesosoma dorsum. Body medium
to dark brown in color, the mandibles, antennae, and tarsi varying from concolorous to a paler luteous
brown.

Comments. — This previously overlooked species ranges from Texas to Panama.
It includes populations from Texas and parts of Mexico which I misidentified as P.
elongatus in Ward (1985). The characteristic features of P. caeciliae are its small
size, relatively dark color, and the sublucid punctate appearance of parts of the head
and mesosoma. It is distinguished from P. elongatus primarily on the basis of this
shiny appearance (all of their metric measurements overlap broadly, although P.
elongatus averages larger in size, with relatively longer eyes - compare HW, HL
and REL). Typically P. elongatus workers have a densely punctate and opaque
head, while in P. caeciliae the punctures on the head are less dense, with
(correspondingly) more extensive shiny interspaces between them, especially in the
area immediately posterior to the compound eye (see Figs. 20, 21, 23); but both
species vary considerably, and the extremes of sculptural variation come close to
overlapping.

Among material examined in the present study, an old series of workers from
Escuintla, Guatemala (Wheeler leg.) included both P. elongatus, P. caeciliae, and
several individuals seemingly intermediate between the two. In western Mexico
(Jalisco, Sinaloa), where P. caeciliae is apparently absent, some P. elongatus
workers approach P. caeciliae in head sculpture. On the other hand in Costa Rica,
where the two species are broadly sympatric, I have seen no intermediates among
material referable to the two species.

P. caeciliae differs from P. cubaensis by its smaller size (HW <0.62), relatively
longer eyes (REL2 0.72-0.81) (Fig. 28), broader fore femur (Fig. 29), and shorter
petiole. While Mexican and Central American populations of these two species are
rather distinct, Jamaican “ caeciliae ” partly bridge the gap between the two. These
Jamaican individuals may, in fact, represent small P. cubaensis.

The relationship of P. caeciliae to the South American species, P. urbanus, is
even more problematic. The existing differences between the two are slight (see key
to species) and not wholly diagnostic. More material, especially worker-associated
queens and males from northern South America, will be needed to resolve this issue.

Biology. — P. caeciliae has been collected nesting in dead twigs of Quercus and
Prosopis in Texas and northern Mexico. Costa Rican nest-site records include dead
twigs of Gliricidia sepium, Anacardium and Laguncularia racemosa. Habitat
records cover a broad range (tropical dry forest, mangrove, littoral vegetation,
riparian forest, edge of second-growth rain forest, rain forest) but suggest a

Quae st. Ent., 1989, 25 (4)

412

Ward

preference for open areas. In Costa Rica the species is notably less common than P.
elongatus.

Material Examined ( CDAE , CNCC, GCWC, JTLC, LACM, MCZC, MHNG,
MZSP, PSWC, USNM). —

COSTA RICA Guanacaste: 6 mi S, 6 mi W Canas, Taboga (D. H. Janzen); Hda. La Pacifica, nr. Canas,
50 m (P. S. Ward); Palo Verde (C. M. Herrera); Santa Rosa Natl. Pk. (D. H. Janzen); Santa Rosa Natl. Pk., <
5 m (P. S. Ward); Limon: Talia Farm, Estrella Valley (G. C. Wheeler); Puntarenas: 9.7 mi W Esparta (D. H.
Janzen); Boca Barranca (D. H. Janzen); Estrella, 300 m (J. T. Longino); Golfito, 5 m (P. S. Ward); Manuel
Antonio Natl. Pk., 10 m (P. S. Ward); Manuel Antonio Natl. Pk., < 5 m (P. S. Ward); Pita, 120 m (J. T.
Longino); Pita, 200 m (J. T. Longino); Sarmiento, 200-300 m (J. T. Longino); Sirena, Corcovado Natl. Pk.,
10 m (P. S. Ward); Sirena, Corcovado Natl. Pk., 50 m (J. T. Longino); San Jose : 2 mi. W Rio Oro (D. H.
Janzen); 40 km S San Isidro de General (D. H. Janzen); San Jose (H. Schmidt).

GUATEMALA Escuintla : Escuintla (W. M. Wheeler; N. L. H. Krauss); Izabal: Los Amates
(Kellerman); Suchitepequez : Patulul (Peper).

HONDURAS Atlantida : La Ceiba (F. S. Dyer); Cortes : Choloma (W. M. Mann).

JAMAICA Kingston (A. Forel); Ford 1 mi SE Stony Creek (E. A. Chapin).

MEXICO “Mexico”, intercepted at Brownsville (c.u.); Camp.: 29 mi E, 12 mi S Campeche (Ruinas
Edzna) (D. H. Janzen); N.L.: El Pastor, Montemorelos, 2200 ft. (W. S. Creighton); S.L.P.: 3 mi N Valles,
300 ft. (W. S. Creighton); Tamps.: 6.2 mi S San Fernando on Hwy 101 (D. H. Janzen); Canon del Abra,
1000 ft. (W. S. Creighton); Ver.: Camaron (E. Skwarra).

NICARAGUA Estelt : 7.5 mi NW San Isidro, 550 m (D. H. Janzen).

PANAMA Chiriqui : “Prov. Chiriqui” (F. M. Gaige); Panama: Trinidad Rio (A. Busck).

UNITED STATES Texas: Cameron Co.: 10 mi W Boca Chica (R. R. Snelling); 5 mi W Boca Chica (G.
C. & J. Wheeler); Brownsville (H. S. Barber; c.u.); Harlingen (W. F. Buren); Hidalgo Co.: Mission (W. F.
Buren); Monte Alto, 60 ft. (W. S. Creighton); Kenedy Co.: 26 mi N Raymondsville (W. S. Creighton).

Pseudomyrmex cretus, sp. nov.
(Figures 9a, 9b, 26, 27)

Holotype worker. — COSTA RICA, Prov. Guanacaste : Santa Rosa Natl. Park,
300m, 10o51'N, 85°37'W, 14.xii. 1983, ex dead twig of Schoepfia, tropical dry forest,
P. S. Ward ace. no. 6434 (MCZC). HW 0.95, HL 1.18, EL 0.61, PL 0.52, PH 0.42.

Paratypes. — Same locality and date as holotype, P. S. Ward ace. nos. 6434 and
6427-2: two series of workers; COSTA RICA, Prov. Guanacaste : Santa Rosa Natl.
Park, 270m, 10°50'N, 85°38’W, 14.xii.1983, P. S. Ward ace. nos. 6443, 6444, 6446,
and 6448, workers and dealate queens. (BMNH, INPA, IZAV, LACM, MCZC,
MJPL, MNCR, MZSP, PSWC, UCDC, UNCB, USNM)

Type material restricted to the above series from Santa Rosa National Park,
Costa Rica. Additional material, believed to be conspecific, is listed below.

Worker Measurements (n=14). — hl 0.99-1.22, HW 0.85-0.98, MFC 0.027-0.035, Cl
0.78-0.86, OI 0.57-0.60, REL 0.49-0.53, REL2 0.62-0.65, OOI 0.07-0.35, VI 0.74-0.81, FCI 0.028-0.039,
SI 0.46-0.50, SI2 0.73-0.80, FI 0.40-0.46, PDI 1.21-1.43, MPI 0.047-0.068, NI 0.60-0.65, PLI 0.77-0.89,
PWI 0.69-0.78, PPWI 1.21-1.50.

Worker Diagnosis. — A relatively large species for the P. oculatus group (HW 0.85-0.98), with
broad head, long scapes, and relatively short eyes (Fig. 9a; SI2 0.73-0.80); sides of head broadly convex,
occipital margin weakly concave in frontal view; second and third funicular segments as long as, or longer
than, wide (FLI 1.94-2.40, n=5). Fore femur relatively slender; metanotal groove conspicuous but not
sharply excavated; basal face of propodeum flat to weakly convex, raised slightly above the level of the
mesonotum, longer than the declivitous face from which it is usually well differentiated; juncture of basal

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413

and declivitous faces of propodeum often marked laterally by a slight angle, which continues as weak
margination flanking the flat declivitous face. Petiole (Fig. 9b) apendunculate with a well-developed
anteroventral process, the node high and broadly rounded in lateral view, rather broad in dorsal view (see
petiolar indices); lateral margination of petiolar node more marked than in most other species of the oculatus
group; postpetiole notably broader than long.

Mandibles striato-punctate; head densely punctate, subopaque, upper third becoming sublucid due to
shiny interspaces between the punctures (which are separated by 0.25 to 1.0 their diameters); mesosoma
punctate to coriarious-imbricate, the mesonotum and centre and sides of pronotum sublucid, remainder
mostly opaque. Petiole, postpetiole, and gaster sublucid, the lustre subdued by numerous fine piligerous
punctures and associated pubescence. Fine erect pilosity and appressed pubescence common on most of the
body, including mesosoma dorsum; the pilosity a little longer and denser than is typical for the oculatus
group. Light to medium castaneous brown, the gaster and femora darker brown (variably so), with varying
degrees of modest infuscation of the mesosoma; mandibles paler, luteous brown.

Comments. — The combination of large size (HW > 0.84) and petiole shape
(node longer than high, broadly rounded in lateral view) separates P. cretus from all
other species except P. curacaensis. P. cretus and P. curacaensis appear to be sister
species; their males both possess unusually lengthened palpal segments and
shortened antennal segments, conditions not known to occur in other members of
the P. oculatus group

The worker of P. cretus averages larger in size than that of P. curacaensis , and in
the region of size overlap (HW 0.85-0.90) it possesses a broader head, shorter eyes,
and longer scapes (see Figs. 26, 27, and worker key). The two species have not been
collected sympatrically, P. cretus being known from Guatemala and Costa Rica, P.
curacaensis from Panama and most of South America. It is possible that
intermediate populations will be located in Central America, and the name P. cretus
would then become a junior subjective synonym of P. curacaensis , but the Panama
collections of P. curacaensis which I have examined do not approach P. cretus in
size or head shape.

Biology. — The available records suggest that P. cretus is an inhabitant of
tropical dry forest. Colonies have been collected in dead twigs of Gliricidia sepium
and Schoepfia sp. at the type locality (Santa Rosa National Park, Costa Rica). At the
same locality workers were observed foraging on low vegetation and visiting
extrafloral nectaries of Croton. Workers collected at Estrella, Costa Rica (J. Longino
leg.) appeared to be nesting in a dead fence post.

Material Examined. — Type material listed above, plus the following (JTLC,
LACM, MZSP, PSWC):

COSTA RICA Guanacaste : 5 km S. Liberia (D. H. Janzen); La Cruz, 250 m (J.T. Longino); Palo Verde
(D. E. Gill; C. M. Herrera); Santa Rosa Natl. Park (L. Gillespie; D. H. Janzen); Santa Rosa Natl. Park, 100 m
(J. T. Longino); Puntarenas : Estrella, 300m (J. T. Longino); Pita, 200m (J. T. Longino).

GUATEMALA Retalhuleu : N edge Champerico (D. H. Janzen); 2 mi. NE Champerico (D. H. Janzen).

Pseudomyrmex cubaensis (Forel)
(Figures 22, 28, 29)

Pseudomyrma elongata var. cubaensis Forel, 1901b: 342. Holotype (unique syntype) worker, Bahia Honda,
Cuba (M. J. Schmitt) (MHNG) [Examined].

Pseudomyrma elongata var. cubaensis Forel; Forel, 1913: 215. [Description of queen].

Quaest. Ent., 1989, 25 (4)

414

Ward

Pseudomyrmex elongatcr, Creighton (nec Mayr), 1955: 18. (partim) [Synonymy of P. cubaensis under P.

elongatus],

Pseudomyrmex cubaensis (Forel); Ward, 1985: 226. [Revalidated as a distinct species].

Worker measurements, Florida and Caribbean (cubaensis 5.5.) (n=24). — hl
0.89-1.02, HW 0.65-0.73, MFC 0.017-0.029, Cl 0.71-0.75, 01 0.52-0.58, REL 0.47-0.51, REL2
0.65-0.71, OOI 0.41-0.95, VI 0.74-0.84, FCI 0.024-0.044, SI 0.44-0.48, SI2 0.64-0.74, FI 0.42-0.48, PDI
1.09-1.44, MPI 0.038-0.084, NI 0.53-0.63, PLI 0.67-0.78, PWI 0.55-0.71, PPWI 1.01-1.28.

Worker measurements, mainland Latin America (n=31). — HL 0.82-1.06, HW
0.65-0.78, MFC 0.015-0.031, Cl 0.69-0.79, 01 0.53-0.60, REL 0.50-0.56, REL2 0.66-0.76, OOI
0.16-0.73, VI 0.75-0.86, FCI 0.022-0.044, SI 0.44-0.50, SI2 0.60-0.72, FI 0.41-0.50, PDI 1.12-1.51, MPI
0.038-0.066, NI 0.54-0.68, PLI 0.75-0.88, PWI 0.57-0.71, PPWI 1.13-1.36.

Worker diagnosis (cubaensis S.S.). — Medium-sized species (HW 0.65-0.73), with elongate
head and relatively short eyes (REL 0.47-0.51); sides of head shallowly convex, occipital margin flat to
slightly concave, in frontal view; funicular segments II and III broader than long. Fore femur only
moderately swollen; metanotal groove conspicuously impressed; basal face of propodeum more or less flat,
raised slightly or not at all above level of mesonotum, rounding gently into the declivitous face which it
exceeds in length. Petiole apenduculate, with a conspicuous anteroventral process (blunt or tooth-like), node
rather long relative to its height (see metrics), and with gentle dorsolateral margination; postpetiole slightly
longer than broad.

Mandibles striato-punctate; head (Fig. 22) densely punctate, subopaque to sublucid, with rather
conspicuous shiny interspaces between the punctures on the upper third of the head, especially between the
ocelli and the compound eye and immediately posterior to the eye. Mesosoma punctate to
coriarious-imbricate, subopaque, with sublucid areas on the side of the pronotum and centre of the pronotum
and mesonotum; petiolar node sublucid, especially its posterior face; postpetiole and gaster sublucid,
covered with numerous fine piligerous punctures. Fine erect pilosity and appressed pubescence covering
most of body, including mesosoma dorsum. Medium to dark brown, the antennae, mandibles, and tarsi
lighter in color.

Comments. — The above description applies to the populations of P. cubaensis
inhabiting Florida and the Caribbean. Elsewhere, in Mexico, Central America, and
South America is a variable array of cubaensis- like populations which I currently
treat as conspecific with the Caribbean form. Workers in these mainland populations
tend to have longer eyes and a shorter and higher petiole (compare REL, REL2, and
PLI in the lists of measurements given above). They also tend to be lighter in color,
especially in South America where some populations contain rather large
orange-brown workers, quite different in appearance from Caribbean P. cubaensis.
Taken together as a whole, however, these mainland populations overlap in size,
shape, and color with the more typical P. cubaensis, and I find no clear
discontinuities in the available (largely worker-based) material. Future work
involving the analysis of queens, males, or biochemical characters might well reveal
several cryptic species.

P. cubaensis ( s.l .) can be diagnosed minimally by the combination of medium
size (HW 0.65-0.78); a punctate-sublucid head; relatively short eyes (REL2
0.65-0.76); moderately broad fore femur (FI 0.41-0.50); and petiole with rounded
dorsolateral margination. The largest workers of P. cubaensis (s.l.) from South
America overlap in size with the smallest workers of P. curacaensis ; they are
distinguished from the latter by the possession of a broader fore femur, longer head,
flatter occipital margin, shorter funicular segments, and gentler dorsolateral
margination of the petiole (see discussion under P. curacaensis ). At the lower limits

Pseudomyrmecine Ants

415

of its size range, P. cubaensis ( s.l .) can be difficult to distinguish from P. urbanus.
Workers of the latter have broader fore femora and longer eyes for a given head
width, compared to P. cubaensis (s.l.) (see scatter plots in Figs. 28 and 29, but note
the variation).

Workers of P. cubaensis (s.l.) differ from those of P. elongatus by their larger
average size, shorter eyes and lower petiole height in the region of size overlap, and
sublucid head and mesosoma.

Biology. — In the Caribbean region, colonies of P. cubaensis occupy dead stems
of various woody and herbaceous plants (Ward, 1985), often in littoral
environments. Mainland populations come from a variety of habitats, including
tropical dry forest, savannah, roadside verge, second-growth lowland rain forest,
rain forest edge and montane rain forest edge. Nesting habits of these mainland
populations are probably also generalized, but unfortunately most records consist of
foraging workers rather than nest series.

Material examined (BMNH, CASC, CKIC, CNCC, CUIC, EBCC, INPA, IZAV,
JTLC, KWJC, LACM, MCSN, MCZC, MHNG, MNHN, MZSP, PSWC, UCDC,
USNM, WPMC). —

P. cubaensis (5.5.)

BAHAMAS Andros I. (W. M. Wheeler); Conception I., S. Bahamas (J. Greenway); Crooked Is., Gun
Point (B. Valentine & R. Hamilton); Fish Hawk Key, Andros I. (W. M. Wheeler); High Ridge Cay (W. M.
Mann); Mangrove Cay, Andros I. (B. Cole); Nassau (W. M. Wheeler); New Providence (B. Cole).

CAYMAN ISLANDS. Grand Cayman (M. E. C. Giglioli); Little Cayman (F. Banks).

CUBA “Cuba” (Cresson; c.u.); 7 km N Vinales (c.u.); Aguada de Pasajeros (W. M. Wheeler); Anafe,
Habana (G. Aguayo); Bahia Honda (Wickham; c.u.); Bolondron (W. M.Wheeler); Cabada (W. M. Mann);
Cabanas, Pinar del Rio (c.u.); Cayamas (Baker; E. A. Schwarz); Cienaga de Zapata (W. M. Wheeler);
Cienfuegos (W. M. Mann); Cristo (W. M. Mann); El Caney (W. M. Mann); Gomez Mena (San Nicholas)
(G. Salt); Guanajay, Pinar del Rio (E. O. Wilson); Guantanamo (W. M. Mann); Guavivo Cave, Soledad (Fal.
Smith); Hanabanilla Falls, Trinidad Mts. (G. Salt); Harvard Bot. Gardens, Soledad, Cienfugos (N. A.
Weber); Havana (A. Bierig; P. Serre); Hormiguero (H. J. MacGillavry); Jaruca, Habana (P. Bermudez);
Jiquari (Barbour & Shaw); Juragua, Las Villas Prov. (E. O. Wilson); La Milpa, nr. Cienfugos (G. Salt);
Limones (W. M. Mann); Limones Seboruco (N. A. Weber); Malas Aguas, Pinar del Rio (E. O. Wilson);
Mina Carlota, Trinidad Mts. (W. M. Mann); Pied, de las Tunas (W. M. Mann); Pinar del Rio (H. J.
MacGillavry); Pinares, Oriente (W. M. Mann); Rio Hanabana (c.u.); San Bias, Trinidad Mts., Santa Clara
(c.u.); Santa Clara, Las Villas Prov. (E. O. Wilson); Soledad (J. G. Myers; Fal. Smith); Soledad Gardens (F.
M. Carpenter); Soledad, Cienfuegos (C. T. Brues; C. T. & B. B. Brues; W. S. Creighton; Darlington);
Soledad, nr. Cienfuegos (N. Banks); Vinales, Pinar del Rio (A. F. Arden); nr. Casa Harvard, Soledad,
Cienfuegos (N. A. Weber).

HAITI “Haiti” (E. C. Leonard); Grande Riviere (W. M. Mann); Mts. N of Jacmel (W. M. Mann).
JAMAICA Balaclava (Wight); Bluefields Bay (E. A. Chapin); Kingston (N. L. H. Krauss; c.u.); St.
Eliz., Pedro Cross, 230 m (J. T. Longino); Troy (Wight); Westmor, Auchindown, 150 m (J. T. Longino).

UNITED STATES Florida: Charlotte Co.: 5 mi S Punta Gorda (E. S. Ross); Punta Gorda (c.u.); Collier
Co.: Collier-Seminole St. Pk. (P. S. Ward); Marco (W. T. Davis); Dade Co.: no specific locality (J. N.
Knull); Biscayne Bay (A. Slosson); Cards Point (W. M. Wheeler); Long Pine Key (W. M. Wheeler); Miami
(F. N. Young); Miami & vicinity (A. E. Wight); Miami Beach (W. L. Brown; A. C. Cole); Paradise Key (D.
Fairchild; A. Wetmore; W. M. Wheeler); Paradise Key, Everglades Nat. Pk. (T. H. & S. P. Hubbell);
Highlands Co.: no specific locality (F. J. Moore); Archbold Biol. Stn. (R. Silberglied); Highlands Hammock
St. Pk. (L. & C. W. Obrien); nr. Sebring (R. W. Klein); Hillsborough Co.: no specific locality (J. C. Bowyer;
R. F. Tinker); Tampa (M. H. Muma); Lake Co.: no specific locality (W. A. Hiers); Lee Co.: “Ft. Meyers”
(c.u.); Ft. Myers (J. C. Bradley); Monroe Co.: Key West (c.u.); Lower Matecumbe Key (W. M. Wheeler); N.
Key Largo (R. W. Klein); No Name Key (P. S. Ward); Osceola Co.: Lake Alfred (M. H. Muma); Palm

Quaest. Ent., 1989, 25 (4)

416

Ward

Beach Co:. Boynton Beach (Wood & Davidson); Sarasota Co.:30mi SE Sarasota, T38S, R22E, S.33 (J. T.
Longino); Long Branch Key, Sarasota (A. C. Cole); Sarasota (A. C. Cole); Seminole Co: Sanford (A. B.
Gahan).

P. cubaensis, mainland

ARGENTINA Misiones: El Dorado (N. Kusnezov); Esperanza (N. Kusnezov); Salta: Coronel Moldes
(Monros).

BOLIVIA Beni: 14 km SW San Borja, 210 m (P. S. Ward); Espiritu (F. Koya).

BRAZIL AM: Ig. Marianil, Rio Branco Rd., 24 km NE Manaus (W. L. Brown); Manaus to Itacoatiara
Hwy, km. 34 (W. L. & D. E. Brown); MT: Vila Vera (M. Alvarenga); PA: C. Araguaia (J. A. Rafael);
Tucurui (W. L. Overal); PE: Ilha Fernando Noronha (M. Alvarenga); RJ: Monumento Rodoviario (c.u.).

COLOMBIA Cauca: Popayan (W. & E. MacKay); Medellin, 1800 m (N. A. Weber); Magdalena:
Gairaca Bay, Parque Tayrona, 0-5 m (C. Kugler); Meta: Pto. Lopez (W. & E. MacKay).

COSTA RICA “Costa Rica” (F. Nevermann); Alajuela: 1 1 mi N Florencia (D. H. Janzen); Guanacaste:
Est. Biol. Palo Verde, 10 m (J. T. Longino); N side Lago de Arenal, 570 m (J. T. Longino); Palo Verde (C.
M. Herrera); Heredia: 3km S Pto. Viejo, 50 m (J. T. Longino); Limon: 5 mi. W Guapiles (D. H. Janzen);
Hamburg Farm, Santa Clara (F. Nevermann); Puntarenas: 1 km NE Tarcoles, 20 m (P. S. Ward);
Monteverde, 1 100 m (P. S. Ward); Ojo de Agua, 800 m (J. T. Longino); Sirena, Corcovado Natl. Pk., 0-100
m (J. T. Longino); Sirena, Corcovado Natl. Pk., 50 m (J. T. Longino); San Jose: San Isidro de General (C. L.
Hogue); San Jose (H. Schmidt).

GUATEMALA Escuintla: Escuintla (W. M. Wheeler); Guatemala: hills above Lake Amatitlan (D. H.
Janzen); Izabal: Lago Izabal, 1.5 km NE El Estor (D. H. Janzen).

GUYANA Mazaruni-Potaro: Kartabo, Bartica Dist. (W. M. Wheeler); Kartabo Point (W. Steiner et al.).
MEXICO Camp: 29 mi E, 12 mi S Campeche (Ruinas Edzna) (D. H. Janzen); Gro: 25.4 mi S
Chilpancingo (D. H. Janzen); Is. Tres Marias: Maria Madre I. (H. H. Keifer); Maria Magdalena I. (R. R.
Snelling); Jal: Est. Biologi'a Chamela, 100 m (P. S. Ward); Mich: 9 mi E Capirio, 800 ft. (E. Fisher & P.
Sullivan); Mor: 5 mi S Cuernavaca (W. S. Ross); Sin: 20 mi E Villa Union, 235 m (E. I. Schlinger); Tab: 3
mi W Cardenas (D. H. Janzen); 13 km W border Chiapas, Rt. 186 (W. P. MacKay); Ver: 9 km NNW
Sontecomapan, 20 m (P. S. Ward); 10 km NW Isla, Ruta 145, 10 m (W. P. MacKay); Est. Biol. “Los
Tuxtlas”, nr. San Andres Tuxtlas (G. Ibarra M.); Los Tuxtlas, 10 km NNW Sontecomapan, 200 m (P. S.
Ward); Mirador (E. Skwarra); Palma Sola (P. Reyes); Sontecomapan (D. H. Janzen).

PANAMA Canal Zone: vie. Pta de los Chivos, 3 km SW Gatun, 100 m (W. L. Brown et al.).

PERU Loreto: Yurac, 67 mi E Tingo Maria (E. I. Schlinger & E. S. Ross).

VENEZUELA Aragua: El Limon, 480 m (C. J. Rosales); Barinas: Pueblo Nuevo (H. Romero); 17 km
SSW Ciudad Bolivia, 240 m (P. S. Ward); Cojedes: 20 km W San Carlos (W. L. & D. E. Brown); Monagas:
Caripe (EMC, JL, CHFH); Uverito (C. J. Rosales & J. A. Gonzales).

Pseudomyrmex curacaensis (Forel)
(Figures 10a, 10b, 26, 27)

Pseudomyrma dolichopsis var. curacaensis Forel 1912: 29. Two syntype workers, Cura5ao (Bugnion)

(MHNG) [Examined], One syntype here designated LECTOTYPE.

Pseudomyrmex curagaoensis [sic] (Forel); Kempf, 1961: 379 [Raised to species].

Worker measurements (n=37). — hl 0.85-1.21, HW 0.74-0.89, ivifc 0.014-0.037, Cl
0.70-0.88, OI 0.53-0.59, REL 0.51-0.58, REL2 0.64-0.77, OOI (-0.04)-0.47, VI 0.74-0.86, FCI
0.018-0.044, SI 0.44-0.49, SI2 0.60-0.74, FI 0.42-0.49, PDI 1.09-1.60, MPI 0.030-0.059, NI 0.56-0.69,
PLI 0.71-0.89, PWI 0.59-0.78, PPWI 1.15-1.64.

Worker diagnosis. — Similar to P. cretus ( q.v.) ; averaging smaller in size (HW 0.74-0.89), with
longer head, shorter scapes, and longer eyes (Fig. 10a) (SI2 0.60-0.74; REL2 0.64-0.77) (see also Figs. 26
and 27); occipital margin varying from flat to (frequently) rather conspicuously concave, in full-face frontal
view; funicular segments II and III about as long as broad, or nearly so (FLI 1.55-2.04, n=6). Propodeum
and petiole (Fig. 10b) similar to that of P. cretus, except correspondingly more slender in smaller
individuals. Sculpture, pilosity, and pubescence more or less as in P. cretus. Color variable: typically
medium to dark brown, with lighter appendages, and a darker infuscated patch on the upper third of the

Pseudomyrmecine Ants

417

head, including the ocelli; some populations ( e.g ., northern Colombia, Curasao) have workers which are
lighter orange-brown in color, with contrasting dark brown hind femur, postpetiole, and gaster.

Comments. — It is curious that this species, one of the more common members of
the P. oculatus group, has been unrecognized since its original description. Kempf
(1961) raised P. curacaensis to species without being able to identify any material
referable to it. In collections I have often found specimens of P. curacaensis
misidentified as P. oculatus, to which they bear only a superficial resemblance,
having a very different petiole shape (compare Figs. 7 and 10b) and head sculpture.

The key characteristics of P. curacaensis are its relatively large size,
sublucid-punctate head, and relatively sharp lateral margination of the petiole (such
that, in dorsal view, the maximum width of the petiole occurs at, rather than below,
the dorsolateral margination). In lateral view, the petiolar node (Fig. 10b) presents a
rounded profile, typical of the P. oculatus group. P. curacaensis is very closely
related to the Central American species, P. cretus ; for distinguishing features see the
description above, the keys to workers and queens, and the discussion under P.
cretus. At the lower end of its size range (worker HW 0.74-0.79), P. curacaensis
may be confused with P. cubaensis ( s.l.) ; the most useful distinguishing characters
are the marginate petiole, more markedly concave occipital margin, and the more
elongate funicular segments, of P. curacaensis. The infuscated patch around the
ocelli also tends to be characteristic, but it is not invariably present. Finally, P.
curacaensis tends to have a less elongate head, and a more slender fore femur, than
P. cubaensis.

Biology. — I have encountered this species in a broad range of habitats, from
tropical thorn forest and roadside vegetation to primary and second-growth rain
forest. Nest-site records include dead twigs or branches of Prosopis juliflora,
Gliricidia sepium, Triplaris sp., Pilosocereus lanuginosus, and unidentified
bombacaceous and leguminous trees.

Material examined (BMHN, CASC, CDAE, GHPC, INPA, IZAV, KWJC, LACM,
MCSN, MCZC, MHNG, MNHN, MZSP, NHMB, PSWC, UCDC, USNM, UWEM,
WPMC, WWBC). —

BOLIVIA Beni: Blancaflor (W. M. Mann); Espiritu (F. Koya).

BRAZIL AM: “Amazonas” (Bates); Praia Grande, Rio Negro, Manaus (W. W. Benson); Res. Campina,
Manaus (W. W. Benson); Rio Taruma Mirim-Igapo (J. Adis); Taruma-Mirim (J. Adis); BA: Salvador (W.
W. Kempf); CE: Itapipoca (C. R. Gongalves); GO: Jatai (F. M. Oliveira); MA: Bacabal (W. W. Kempf);
MG: Arassuahy (Thieman); Campo Grande (Gilbert-& Muller); MS: Faz. Dr. Jose Mendes, Tres Lagoas
(Exp. Depto. Zool.); MT: Barao de Melgago (W. F. Buren); Pluva, 15 km N I. de Taiama (C. R. F. Brandao);
Sto. Antonio (c.u.); Utiariti, Rio Papagaio, 325 m (K. Lenko); PA: Belem (W. F. Buren; C. R. Gongalves);
Cach. Caju (Sampaio); Igarape-Agu (c.u.); Soure (c.u.); PE: Olinda (Reichensperger); RD: Porto Velho (C.
R. Gonsalves); RN: Ceara-Mirim (W. M. Mann); RR: Boa Vista (C. R. Gon?alves); SP: Faz. Itaquere, Boa
Esperanga do Sul (K. Lenko); Faz. Itaquere, Nova Europa (K. Lenko).

COLOMBIA Guajira: Maicao (W. L. Brown & C. Kugler); Serrania de Macuira, 6-8 km S Nazareth
(W. L. Brown & C. Kugler); Huila: 10 km. SE Hobo (W. & E. MacKay); 12 km. S Neiva (W. & E.
MacKay); 8 km. S Neiva (W. & E. MacKay); Colombia (W. & E. MacKay); Magdalena: 5 km SE Rio Frio,
100 m (P. S. Ward); Arroyo de Arena (F. M. Gaige); Gairaca, Parque Tayrona (C. Kugler); Rio Frio
(Darlington); Rio Frio, 20 m (P. S. Ward); Santa Marta, 40 m (P. S. Ward); Meta: Carimagua, 370 m (M.
Com); El Castillo (W. & E. MacKay); Tolima: Ibague (W. & E. MacKay).

Quaest. Ent., 1989, 25 (4)

418

Ward

CURASAO “Curacao” (Bugnion); Damacar (J. Maldonado C.).

ECUADOR prov. unknown: Limon Cocha & vie. (H. R. Hermann).

FRENCH GUIANA Cayenne (Pillault); Roura (G. H. Perrault).

GUADELOUPE no locality specified (A. Bonfils).

GUYANA Mazaruni-Potaro: Forest Settlement, R. Mazaruni (N. A. Weber); Kartabo, Bartica Dist. (W.
M. Wheeler); Rupununi : Dadanawa, 100 m (J. T. Longino); Karanambo, 100 m (J. T. Longino).

PANAMA Canal Zone : 5 km WNW Gatun Dam, 160 m (P. S. Ward); Frijoles (W. M. Wheeler); vie.
Pta de los Chivos, 3 km SW Gatun, 100 m (W. L. Brown et al.); Code : La Negrita, nr. Penonome (D.
Cordero); Colon: Colon (E. Adams); Panama: Las Cumbres (M. Daykin); Matfas Hernandez (N. L. H.
Krauss); Rio Piedras (D. H. Janzen).

PERU Huanuco: Monson Valley, Tingo Maria (E. I. Schlinger & E. S. Ross); Panguana, 220 m (M.
Verhaagh); Madre de Dios: 30 km SW Pto. Maldonado, 290 m (T. L. Erwin et al .); San Martin:: 24 km
NNE Tarapoto, 220 m (P. S. Ward); Convento, 26 km NNE Tarapoto, 220 m (P. S. Ward); La Perla, 21 km
NNE Tarapoto, 220 m (P. S. Ward); Tarapoto, 350 m (P. S. Ward).

ST. VINCENT Layou, St. Vincent (J. F. Lynch); Leeward side, St. Vincent (H. H. Smith); St. Vincent
(c.u.).

TRINIDAD “Trinidad” (A. Forel; N. A. Weber; c.u.); Curepe (F. D. Bennett); Imp. Coll. Trop. Agr. (R.
D. Shenefelt); Mayaro Bay (N. A. Weber); Nariva Swamp (N. A. Weber); St. Augustine (N. A. Weber).

VENEZUELA Anzoategui: La Leona (H. Romero); Aragua: El Limon, 450 m (C. J. Rosales); Maracay,
450 m (P. S. Ward); Barinas: 17 km SSW Ciudad Bolivia, 240 m (L. Lattke; P. S. Ward); Bolivar: Cuidad
Bolivar (E. A. Klages); Rio Cuyuni, 66 km SSE El Dorado, 250 m (P. S. Ward); Dto. Federal: Camuri
Grande, Pica “La Guardia” (A. Escalante); Caracas (N. A. Weber); Valle Camuri Grande, nr. La Guaira,70
m (I. Massin); Guarico: Altagracia de Orituco (C. R. F. Brandao & L.O.Campos; illegible); Est. Exp. Los
Llanos, 27 km S Calabozo (C. R. F. Brandao); Hato Masaguaral (Y. D. Lubin); Miranda: Higuerote (K.
Jaffe; G. Villegas); Monagas: Estacion Uverito (H. Romero); Uverito (C. J. Rosales & J. A. Gonzales);
Portuguesa: El Oso (H. Romero); Yaracuy: San Felipe, El Penon (C. Fuentes); Zulia: El Tucuco, Perija (R.
W. Brook et al.)', Maracaibo (E. Poirier; R. G. Wesson); state unknown: Akuriman (Anduze).

Pseudomyrmex eduardi (Forel)
(Figures 16, 17, 25)

Pseudomyrma eduardi Forel, 1912:29. Holotype (unique syntype) worker, Kingston, Jamaica (Forel)

(MHNG) [Examined],

Worker measurements (n = 11). — - HL 0.74-0.80, HW 0.50-0.57, MFC 0.009-0.018, Cl

O. 67-0.72, OI 0.53-0.59, REL 0.52-0.55, REL2 0.74-0.79, OOI 0.28-0.70, VI 0.79-0.86, FCI 0.017-0.032,
SI 0.43-0.47, SI2 0.55-0.62, FI 0.46-0.53, PDI 0.93-1.31, MPI 0.039-0.073, NI 0.52-0.64, PLI 0.79-0.92,
PWI 0.62-0.72, PPWI 1.19-1.50.

Worker diagnosis. — Similar to P. caedliae (q.v.) in size, shape, and color; differing in sculpture-
head largely smooth and shining, with scattered fine punctures (Fig. 25) which become coarser (but still
somewhat effaced) in the region posterior to the compound eye; the punctulate-coriarious sculpture of the
mesosoma is also somewhat effaced, producing rather strongly shining areas in the centre of the mesosoma
dorsum and on the side of the propodeum; petiole, postpetiole, and gaster with a shinier appearance than in

P. caedliae, although still covered with moderately dense pubescence. Dark brown, the mandibles and tarsi
lighter. Clypeus as in Figs. 16 and 17.

Comments. — This species is easily recognized by its combination of small size
(HW <0.58), dark color, and smooth shiny appearance (especially on the head and
mesosoma dorsum). The sculpture in the middle of the head between the compound
eyes is particularly characteristic, consisting of very fine punctures and a few
slightly coarser ones. All other species in the P. oculatus group have denser, coarser
punctate sculpture on this part of the head.

Pseudomyrmecine Ants

419

Biology. — I collected colonies of P. eduardi in dead twigs of Conocarpus
erectus and Avicennia germinans in mangrove in Venezuela. Other biological
collection data include: in tree fall, wet forest (Costa Rica), dead stem of beach tree
(Manaus, Brazil), in rain forest (Huanuco, Peru).

Material examined (CKIC, FFIC, INPA, IZAV , JTLC, KWJC, LACM, MCZC,
MHNG, MNHN, MZSP, PSWC, USNM, WPMC, WWBC).—

BOLIVIA Beni : Cavinas (W. M. Mann); La Paz: Lower Rio Madidi (W. M. Mann); Pando : Abuna (W.
M. Mann).

BRAZIL AM: Praia Grande, Rio Negro, Manaus (W. W. Benson); Rio Taruma Mirim-Igapo (J. Adis);
Taruma-Mirim (J. Adis); km. 27, ZF-3, near Manaus (W. W. Benson); MS: Imbirussu-Corumba (K. Lenko);
PA: Tucurui (R. B. Neto; W. L. Overal); RD: “Jl-Parana/Gleba G. Il-Perdida” (J. Arias).

COLOMBIA Cauca: Popayan (W. & E. MacKay); Cundinamarca: Medina (F. Fernandez); Magdalena:
Gairaca, Parque Tayrona (C. Kugler); Meta: Carimagua, 370 m (M. Com); Villavicencio (W. & E.
MacKay).

COSTA RICA “Costa Rica” (P. Serre); Heredia: 3 km S Pto. Viejo, 50 m (J. T. Longino).

JAMAICA Kingston (A. Forel); St. Eliz., Pedro Cross, 230 m (J. T. Longino); St. James, Porto Bello, 75
m (J. T. Longino); Trelawny, Green Park, 0-150 m (J. T. Longino).

PANAMA Panama: Marias Hernandez (N. L. H. Krauss).

PERU Huanuco: Panguana, 220 m (M. Verhaagh).

VENEZUELA Anzoategui: 1 km S El Hatillo, <5 m (P. S. Ward); Aragua: El Limon, 480 m (C. J.
Rosales); Bolivar: Ciudad Bolivar (N. A. Weber); Cojedes: Jabillal (H. Romero); Dto. Federal: Caracas,
Jardin Botanica (G. Morillo); Guarico: Est. Exp. Los Llanos, 27 km S Calabozo (C. R. F. Brandao); Sucre: 2
km W Cumana (D. H. Janzen); nr. Cumana (D. H. Janzen); T. F. Delta Amacuro?: “Orinoco Delta” (N. A.
Weber).

Pseudomyrmex elongatus (Mayr)
(Figures 6, 14, 15, 20, 21)

Pseudomyrma elongata Mayr, 1870:413. Syntype worker, S. Fe de Bogota, Colombia (Schaufuss) (NHMV)
[Examined]; here designated as LECTOTYPE.

Pseudomyrma elongata var. tandem Forel, 1906:228. Syntype workers, El Hiquito, near San Mateo, Costa
Rica (P. Biolley) (MHNG) [Examined] [Synonymy by Creighton, 1955: 18; here confirmed],
Pseudomyrma alliodorae Wheeler, 1942: 157. Syntype workers, queen, Ancon, Canal Zone, Panama (W. M.
Wheeler) (LACM, MCZC) [Examined]. One syntype worker in MCZC here designated LECTOTYPE.
Syn. nov.

Pseudomyrma allidora Enzmann, 1945:77. Syntype workers, queen, Ancon, Canal Zone, Panama (W. M.
Wheeler) (LACM, MCZC) [Examined] [Objective synonym of P. alliodorae Wheeler; Brown, 1949:42]

Worker measurements ( n-55 ). — hl 0.73-0.93, hw 0.56-0.68, MFC o.oi 1-0.027, ci
0.66-0.79, OI 0.50-0.59, REL 0.53-0.59, REL2 0.73-0.82, OOI 0.04-0.67, VI 0.75-0.90, FCI 0.019-0.047,
SI 0.42-0.48, SI2 0.54-0.63, FI 0.43-0.56, PDI 1.06-1.44, MPI 0.032-0.070, NI 0.54-0.65, PLI 0.79-1.09,
PWI 0.62-0.87, PPWI 1.09-1.48.

Worker diagnosis. — Similar to P. caeciliae ( q.v .), but averaging larger in size with longer eyes in
relation to head length (Fig. 6; compare HW, HL, and REL); ocellar distance variable (OD/HW
0.097-0.165). Clypeus as in Figs. 14 and 15. Basal face of propodeum tending to be less elevated above
level of mesonotum (compared to P. caeciliae) and usually meeting the declivitous face at a sharper angle.
Head densely punctate and predominantly opaque; punctures on upper half of head mostly contiguous,
separated by half their diameters or less (except for a thin shiny margin about 0.010 to 0.020 mm wide
immediately posterior to the compound eye). Mesosoma punctate to coriarious-imbricate, the central areas
more heavily sculptured and less shiny than in P. caeciliae.

Comments. — P. elongatus is a common and widespread species whose key
features are its relatively small size (HW 0.56-0.68); densely punctate and

Quaest. Ent., 1989, 25 (4)

420

Ward

predominantly opaque head; long eyes (REL > 0.53, REL2 > 0.73); and relatively
short, high petiole (PLI > 0.79). Distinctions between it and P. caeciliae, P.
cubaensis, and P. urbanus are discussed under those species.

I found the type worker of P. elongatus, which had previously eluded discovery
(Ward, 1985), during a visit to NHMV. It had been misplaced under Pseudomyrmex
elongatulus (Dalle Torre), a replacement name for the preoccupied P. elongatus (F.
Smith, 1877), which is a markedly different species (see Kempf, 1967).

Biology. — This species is a generalist inhabitant of dead twigs of woody plants.
It has been collected in mangroves, old fields, along roadsides, and in forested
habitats ranging from tropical dry forest to secondary and primary rain forest.
Specific nest-site records include dead twigs of Avicennia germinans, Rhizophora
mangle, Laguncularia racemosa, Baccharis halimifolia, Baccharis trinervis,
Baccharis sp., Ficus spp., Vernonia patens, Terminalia catappa, Tecoma stans,
Zanthoxylum culantrillo, Gliricidia sepium, Inga sp., Helicteres sp., Coccoloba sp.,
and Sapium sp. Also I collected a colony in a live branch of Pithecellobium saman,
and there are records from (apparently live) stems of Cecropia, Cordia alliodora ,
and Triplaris cumingiana. Thus, it appears that P. elongatus will occupy ant-plant
domatia opportunistically.

Material examined ( BMNH , CASC, CDAE, CELM, CNCC, CUIC, EBCC, FFIC,
GCWC, ICCM, INPA, IZAV, JTLC, KWJC, LACM, MCSN, MCZC, MHNG, MZSP,
NHMB, NHMV, PSWC, UCDC, USNM, UWEM, WPMC, WWBC).—

BELIZE El Cayo (N. L. H. Krauss).

BOLIVIA Beni: 46 km SSW San Borja, 300 m (P. S. Ward); Est. Biol. Beni, 42 km E San Borja, 210 m
(P. S. Ward); Huachi (W. M. Mann); Reyes (W. M. Mann); La Par. Espia Rio Bopi (W. M. Mann); Sta.
Helena (W. M. Mann).

BRAZIL AC: Porto Valter (P. L. Herbst); AM: Arquipelago d. Anavilhanas, Rio Negro (c.u.); Faz.
Dimona, 80 km N Manaus, 80 m (P. S. Ward); High Falls, Rio Taruma (W. L. Brown); Ig. Marianil, Rio
Branco Rd., 24 km NE Manaus (W. L. Brown); Manaus (C. R. Gon5alves; c.u.); Manaus, 60 m (P. S. Ward);
Manaus-AM.010, km. 10 (A. Y. Harada); Praia Grande, Rio Negro, Manaus (W. W. Benson); Reserva
Ducke (J. A. Rafael); Reserve Ducke, NE of Manaus (W. L. Brown); Rio Taruma Mirim-Igapo (J. Adis);
Taruma-Mirim (J. Adis); km. 24, ZF-3, near Manaus (W. W. Benson); km. 27, ZF-3, near Manaus (W. W.
Benson); BA: Salvador (N. L. H. Krauss); ES: Linhares (M. Alvarenga); GO: Araguacema (J. A. Rafael);
Jatai (F. M. Oliveira); MA: Bacabal (W. W. Kempf); MG: Pedra Azul, 800 m. (Seabra & Alvarenga); MS:
Faz. Canaa, Tres: Lagoas (F. Lane); Faz. Floresta, Mun. Tres: Lagoas (Exp. Depto. Zool.);
Imbirussu-Corumba (K. Lenko); Ladario (Gilbert & Muller); Serra do Urucum, Corumba (K. Lenko); Tres
Lagoas (Exp. Depto. Zool.; K. Lenko); MT: Sinop (Alvarenga & Roppa; M. Alvarenga); MT?\ “Matto
Grosso” (Germain); PA: Belem (P. Vanzolini; c.u.); Belem-Tapana (A. Y. Harada); C. Araguafa (A. Y.
Harada; J. A. Rafael); C. Araguafa, Campo Gramado (A. Y. Harada); Cach. (do) Breu (Sampaio);
Igarape-Agu (c.u.); Oriximina, Alcoa Miner. M. Branco (A. Y. Harada); Oriximina-Rio Trombetas, Alcoa
Miner. Campina (A. Y. Harada); Sa. dos Carajas (25 km W Camp N-l) (W. W. Benson); Santarem,
Taperinha (R. L. Jeanne); Tucurui (R. B. Neto; W. L. Overal); Utinga track, nr. Belem (K. Lenko); PE:
Tapera (B. Pickel); RD: Porto Velho (C. R. Gonsalves); Vilhena (M. Alvarenga); RJ: Cabo Frio (C. Gilbert);
Fonseca, Niteroi (C. R. Gon§alves); Guaratiba (A. Silva); I. Govemador (C. R. Gongalves); Marambaia (R.
Muller); Rio de Janeiro, Taquara (T. Borgmeier); Sao Bento (P. A. Telles); “Guanabara dos Bandeirantes”
(N. L. H. Krauss); SP: Adolfo (“P. de preia Rio Tiete”) (M. Dimiz); Agudos (C. Gilbert; R. Muller);
Anhembi, Faz. Barr. Rico (W. W. Kempf et al .); Barabosa-Tiete (J. Diniz [= missp. of M. Dimiz?]);
Cachoeira das Emas (EEBP), Mun. Piragununga (W. L. Brown); Campinas (W. W. Benson); Faz. Itaquere,
Nova Europa (K. Lenko); Franca (c.u.); S. Sebastiao (Fleddermann; W. W. Kempf); Ubatuba (P. C.
Montouchet); state unknown: Chapada (c.u.).

Pseudomyrmecine Ants

421

COLOMBIA Bolivar. Cartagena (c.u.); Cauca : 15 mi. S Corinto, 1140 m (E. I. Schlinger & E. S. Ross);
Cundinamarca : “S. Fe de Bogota” (Schaufuss); Anapoima (I. Zenner); El Colegio & Anapoima (I. Zenner);
Guajira : Puente Bomba, nr. Dibulla (W. L. & D. E. Brown); Rio Don Diego (C. Kugler & W. L. Brown);
Serrania de Macuira, W. of Nazareth (C. Kugler & W. L. Brown); Huila : 10 km S Campoalegre (W. & E.
MacKay); 10 km. SE Hobo (W. & E. MacKay); 10 km. W Palermo (W. & E. MacKay); 1 km S Neiva (W.
& E. MacKay); 4 km. S Tello (W. & E. MacKay); Colombia (W. & E. MacKay); La Plata (W. & E.
MacKay); Palermo (W. & E. MacKay); Magdalena : 2 km E Orihueca, 20 m (P. S. Ward); 2 km ESE Minca,
780 m (P. S. Ward); 5 km SE Rio Frio, 100 m (P. S. Ward); Bahia de Gairaca, Tayrona Pk, 20 km NE Santa
Marta (H.-G. Muller); Canaveral, 50 m (P. S. Ward); Canaveral, 5 m (P. S. Ward); Canaveral, <5 m (P. S.
Ward); Digrera, base of mts. nr. Santa Marta (C. Kugler & W. L. Brown); Minca (W. L. Brown); Rio Frio
(Darlington); Rio Frio, 20 m (P. S. Ward); Villa Culebra, nr. Bonda, 10 km E Sta. Marta (H.-G. Muller);
Magdalena?: locality illegible (A. Forel); Meta: Reserva Macarena, 560 m (F. Fernandez); Reserva
Macarena, Cano La Curia, 580 m (F. Fernandez); Valle: 6 km S Univ. Valle (W. L. Brown); Cali (N. L. H.
Krauss); Cali, in quarantine at San Francisco (c.u.); Dagua (W. & E. MacKay).

COSTA RICA “Costa Rica” (Alfaro; P. Biolley); Alajuela: 1 1 mi. N Florencia (D. H. Janzen); El
Hiquito near San Mateo, 250 m (P. Biolley); Guanacaste: 1 km SW Pto. Coyote, <5 m (P. S. Ward); 4 km N
Canas (D. H. Janzen); Heredia: 10°20'N, 84°04'W, 500 m (J. T. Longino); 3 km S Pto. Viejo, 50 m (J. T.
Longino); F. La Selva, 3 km S Pto. Viejo (H. A. Hespendeide); Finca La Selva (D. H. Janzen); Limon: La
Lola (D. H. Janzen; M. J. Stelzer; N. A. Weber); Zent (F. Lara); Puntarenas: 14 km E Palmar Norte, 120 m
(P. S. Ward); 14 km E Palmar Norte, 70 m (P. S. Ward); 1 km NE Tarcoles, 20 m (P. S. Ward); 5 km N
Ciudad Neily, 780 m (P. S. Ward); Golfito, 5 m (P. S. Ward); Guaria, 700 m (J. T. Longino); Llorona,
Corcovado Natl. Pk. (J. T. Longino); Llorona, Corcovado Natl. Pk., 10 m (P. S. Ward); Manuel Antonio
Natl. Pk., 5 m (P. S. Ward); Manuel Antonio Natl. Pk., <5 m (P. S. Ward); Monteverde, 1100 m (P. S.
Ward); Monteverde, 1200 m (P. S. Ward); Osa Penin., Corcovado Natl. Pk. (J. T. Longino); Palmar Sur (D.

H. Janzen); Reserva Biol. Carara, 10 m (P. S. Ward); Reserva Biol. Carara, 500 m (P. S. Ward); Sirena,
Corcovado Natl. Pk. (J. T. Longino); San Jose: 3.5 km NE Santiago de Pur (D. H. Janzen); 40 km S San
Isidro de General (D. H. Janzen).

ECUADOR Manabi: Puna Island (Ross & Michelbacher); Pichincha: 6 mi W Santo Domingo de los
Colorados (E. I. Schlinger & E. S. Ross).

EL SALVADOR San Salvador (N. L. H. Krauss); Santa Anna (N. L. H. Krauss).

GUATEMALA “Guatemala”, quarantine San Francisco (c.u.); Escuintla: Escuintla (W. M. Wheeler);
Escuintla?: San Jose (E. S. Ross; E. I. Schlinger & E. S. Ross); Guatemala: Guatemala City, quarantine San
Francisco (c.u.); Peten: Tikal (N. L. H. Krauss); Retalhuleu: 1.3 mi E Champerico (D. H. Janzen); 5 mi W
Retalhuleu, Hwy. CA-2, at Rio Nil (D. H. Janzen).

GUYANA Mazaruni-Potaro: Kartabo, Bartica Dist. (W. M. Wheeler); North West: Wauna &
Tokomabu (J. G. Myers); Rupununi: Dadanawa, 100 m (J. T. Longino).

MEXICO Is. Tres Marias: Maria Magdalena I. (H. H. Keifer; R. R. Snelling); Jal.: 14 km SSE Chamela,
20 m (P. S. Ward); 2 km E Chamela, 20 m (P. S. Ward); 32.8 mi. SW Autlan (D. H. Janzen); 6 km NW El
Tuito, 560 m (P. S. Ward); Mor.: Cocoyotla, munic. Coatlan del Rio, 1 120 m (G. Aleman); Cuzuhuatlan (G.
Aleman); Los Alcanfores, Mpo. Xochitepec (G. Aleman); Tetecala, 800 m (G. Aleman); Oax.: 19 km N San
Pedro Pochutla, 200 m (W. P. MacKay); S.L.P.: Rio Amahac, Tamazunchale, 300 ft. (W. S. Creighton);
Tamazunchale (W. S. Ross); Sin.: 4.6 mi E Chupaderos (E. I. Schlinger); Ver.: Cotaxtla Exp. Sta, Cotaxtla
(D. H. Janzen); Yuc.: 11 km S Progreso (E. O. Wilson).

NICARAGUA Matagalpa: El Tuma, 30 km E Matagalpa (C. Kugler & J. Hahn).

PANAMA Bocas del Toro: Changuinola Dist. (G. C. Wheeler); Canal Zone: 2 km W Gamboa, 30 m (P.
S. Ward); 5 km WNW Gatun Dam, 160 m (P. S. Ward); 6 km NW Gamboa, 50 m (P. S. Ward); 6 km NW
Gatun Dam, 75 m (P. S. Ward); Ancon (W. M. Wheeler); Barro Colorado I. (W. L. Brown & E. S.
McCluskey; H. Wolda; G. C. Wheeler; J. Zetek); Cerro Galera, 9 km W Panama City, 200 m (W. L. Brown
et a/.); Chiva Chiva (N. L. H. Krauss); Fort Sherman, Island P2 (D. Simberloff); Frijoles (G. C. Wheeler; W.
M. Wheeler); Miraflores (W. M. Wheeler); Ruta 1, 14 km W Panama City, 100 m (W. L. Brown et al); Ruta

I, Howard AFB, W of Panama City, 50 m (W. L. Brown et al .); vie. Pta de los Chivos, 3 km SW Gatun, 100
m (W. L. Brown et al.); Chiriqui: “Prov.Chiriqui” (F. M. Gaige); 10.7 mi ESE La Concepcion (D. H.
Janzen); Colon: Colon (E. Adams); Panama: Bella Vista (W. M. Wheeler); Cerro Azul, 800 m (E.
Mockford); Isla San Jose, Islas Perlas (C. Campbell); Matias Hernandez (N. L. H. Krauss); Pueblo Nuevo
(W. M. Wheeler); Rio Piedras (D. H. Janzen); Veraguas: 0.3 mi SW Santiago (D. H. Janzen); prov .unknown:
“Lilihe R” (W. M. Wheeler).

Quaest. Ent., 1989, 25 (4)

422

Ward

PERU Madre de Dios: 30 km SW Pto. Maldonado, 290 m (T. L. Erwin et al.)\ Cocha Cachu, Rio Manu
(D. Hunt); San Martin: Convento, 26 km NNE Tarapoto, 220 m (P. S. Ward); La Perla, 21 km NNE
Tarapoto, 220 m (P. S. Ward); Tarapoto, 350 m (P. S. Ward).

TRINIDAD Maracas Valley (N. A. Weber).

UNITED STATES Florida: Collier Co.: 6 mi S Naples (G. S. Walley); Everglade[s] (W. T. Davis);
Dade Co.: no specific locality (J. N. Knull); Cocoanut Grove (c.u.); Miami (c.u.); Miami, 10 m (P. S. Ward);
Rattlesnake Hammock, Homestead (R. Gregg); Shark Valley, Everglades Natl. Pk. (P. S. Ward); Highlands
Co.: Archbold Biol. Stn., L. Placid (T. C. Schneirla); Lee Co.: Ft. Myers (W. M. Barrows; D. E. Read);
Monroe Co.: Big Pine Key (E. O. Wilson); John Pennekamp St. Pk., <5 m (P. S. Ward); Key Largo (A.
Wetmore; c.u.); Key West (E. A. Schwartz); Lower Matecumbe Key (J. R. Vockeroth); N. Key Largo (R.
W. Klein); Plantation Key (E. O. Wilson).

VENEZUELA “Venezuela”, in quarantine at San Francisco (c.u.); Aragua: El Limon, 450 m (C. J.
Rosales); El Limon, 480 m (C. J. Rosales); Barinas: 17 km SSW Ciudad Bolivia, 240 m (P. S. Ward); 17 km
SW Ciudad Bolivia, 240 m (P. S. Ward); Bolivar: 49 km ENE Tumeremo, 200 m (P. S. Ward); Carabobo:
Puerto Cabello (W. L. Brown); Cojedes: 20 km W San Carlos (W. L. Brown); Dto. Federal: La Pastora,
Caracas (c.u.); La Trinidad, Caracas (M. E. Lopez); Guarico: Altagracia de Orituco (C. R. F. Brandao & L.

O. Campos); Hato Masaguaral (Y. D. Lubin); Merida: Merida (A. S. Menke & D. Vincent); T. F. Amazonas:

P. N. Duida-Marahuaka, Culebra, 250 m (K. Jaffe); T. F. Delta Amacuro: Palo Blanco (H. Romero); T. F.
Delta Amacuro?: “Orinoco Delta” (N. A. Weber); Trujillo: 19 km E Bocono, 600 m (J. T. Longino; P. S.
Ward); Zulia: El Tucuco, Perija (R. W. Brooks et al.).

Pseudomyrmex oculatus (F. Smith)

(Figure 7)

Pseudomyrma oculata Smith, 1855:159. Two syntype workers, Santarem, Brazil (Bates) (BMNH)
[Examined]. One syntype worker here designated LECTOTYPE.

Pseudomyrma advena F. Smith, 1855:157. [Objective synonym of P. oculatus', Forel, 1911:277. See
discussion by Forel (1911) and Kempf (1961)].

Pseudomyrma dolichopsis Forel, 1899:87. Syntype workers, Suerre, Jimenez, Costa Rica (Alfaro) (MHNG,
MCSN) [Examined] [Synonymy by Kempf, 1961:375; here confirmed].

Pseudomyrma dolichopsis var. implicata Forel, 1911:277. Two syntype workers, Amazonas (Bates)
(MHNG) [Examined] [Synonymy by Kempf, 1961:375; here confirmed]. One syntype worker here
designated LECTOTYPE (see comments below).

Pseudomyrma sericea var. altinoda Mann, 1916:427. Syntype workers Porto Velho, Rio Madeira, Brazil
(Manft & Baker) (MCZC) [Examined] Syn. nov.

Pseudomyrma wessoni Enzmann, 1945:97. Syntype queen, Peru, (MCZC) [Examined] Syn. nov.
Pseudomyrma wessoni var. tuberculata Enzmann, 1945:98. Holotype worker, Peru [Not examined] Syn. nov.
Pseudomyrmex oculatus (F. Smith); Kempf, 1961:375 [Redescription of worker and queen].

Worker measurements (n=24). — HL 0.94-1.09, HW 0.73-0.91, MFC 0.017-0.031, Cl
0.77-0.87, OI 0.57-0.61, REL 0.55-0.62, REL2 0.67-0.78, 00!-(-0.20)-0.37, VI 0.72-0.81, FCI
0.021-0.037, SI 0.46-0.50, SI2 0.62-0.72, FI 0.40-0.44, PDI 0.97-1.40, MPI 0.032-0.056, NI 0.72-0.81,
PLI 0.86-1.01, PWI 0.66-0.82, PPWI 1.35-1.61.

Worker diagnosis. — Medium to large species, for the oculatus group (HW 0.73-0.91), with broad
head (Cl 0.77-0.87) and with large, elongate eyes which tend to protrude conspicuously from the sides of
the head and which typically (but not always) reach the level of the median ocellus, when head is held in
full-face view; occipital margin flat to moderately concave; funicular segments II and III about as long as
broad, or nearly so. Fore femur relatively slender (FI 0.40-0.44); metanotal groove moderately impressed;
basal face of propodeum flat to weakly convex (lateral view), conspicuously elevated anteriorly above level
of mesonotum; basal face of propodeum usually exceeding length of the declivitous face and rounding into
the latter, which is laterally submarginate. Petiolar node (Fig. 7) high and subangulate, with distinctive
lateral profile: anterior face smoothly continuous with the flattened, posteriorly uplifted dorsal face which
itself rounds sharply into the vertical posterior face; anteroventral petiolar process conspicuous, variable in
shape (from bluntly rounded to slightly recurved and hook-like); postpetiole broader than long, in lateral
view appearing rather high, with the convex, gradually ascending anterodorsal face rounding into a more

Pseudomyrmecine Ants

423

steeply descending posterior face.

Head densely punctate and typically opaque; becoming sublucid on upper third of head in some
individuals, because of shiny interspaces between some of the punctures. Mesosoma punctate to
coriarious-imbricate and predominately opaque; petiole, postpetiole, and gaster subopaque, covered with
numerous fine piligerous punctures. Erect pilosity and appressed pubescence common on most of body,
including mesosoma dorsum. Medium to dark brown, the antennae, tibiae, and tarsi variably lighter, often
contrastingly so; pronotum and parts of head sometimes lighter in color than gaster; mandibles pale luteous.

Comments. — This common, widely distributed species shows considerable
geographical variation (particularly in size, sculpture, color, and head shape), but it
is at once recognizable by the distinctive shape of the petiolar node. The sudden,
subangulate juncture of the dorsal and posterior faces of the node (Fig. 7) is seen in
no other related species. Also characteristic of P. oculatus are the elongate eyes (see
REL and REL2 values), relatively broad head, the (typically) punctate-opaque
sculpture of the head, and the shape of the postpetiole in lateral view (summit of
postpetiolar node displaced posteriorly relative to other species in the P. oculatus
group, except P. schuppi ).

Lectotypes of both P. oculatus and P. implicatus are designated because the
apparent type series of these taxa contain more than one species. In the BMNH,
under P. oculatus , I found three species: (1) two syntype workers of what is here
interpreted to be P. oculatus, from Santarem, Brazil; (2) an alate queen of P. peperi
(Forel), a Central American acacia-ant labelled (obviously erroneously) “Santarem”;
and (3) four workers of P. filiformis , two of which (from Villanova, Brazil) were
labelled as types of P. oculatus. The type series of P. implicatus in MHNG contains
two species: (1) two workers of P. oculatus ; and (2) one worker of P. curacaensis.
For nomenclatural stability I have chosen as lectotypes of P. oculatus and P.
implicatus those workers which correspond to the species described (at least as a
queen) and illustrated (as a worker, under the name P. advena ) by F. Smith (1855).
The choice of lectotypes is also consistent with Kempf’s (1961) concept of P.
oculatus, which was based on an examination of material in the Hope Collection,
Oxford (OXUM) which Kempf considered to represent part of the type series of P.
oculatus.

The syntype queen of P. wessoni (Enzmann) is clearly conspecific with P.
oculatus. Unfortunately, the type of P. wessoni tuberculatus (Enzmann) cannot be
located, and second-guessing the identity of Enzmann’ s creations is an unsatisfying
task. Nevertheless there are enough features of the original description (broad head;
large eyes; densely punctate, opaque head; petiolar profile similar to that of P.
wessoni) to support synonymy under P. oculatus.

Biology. — P. oculatus is typically found nesting in dead twigs or branches, in
rain forest or at the edge of rain forest, less frequently in open or drier habitats.
Specific nest-site records include dead twigs of Ficus, Hampea, Inga, Vochysia, and
various unidentified lianas; I found one colony in the dead leaf sheath of a Scheelea
palm.

Quaest. Ent., 1989, 25 (4)

424

Ward

Material examined (BMNH, CASC, CNCC, CUIC, FFIC, GCWC, GHPC, ICCM,
INPA, JTLC, KSUC, LACM, MCSN, MCZC, MHNG, MZSP, PSWC, UCDC, USNM,
UWEM, WWBC). —

BELIZE Belize (c.u.); San Ignacio, Cayo, 60 m (S. E. Schoenig); Manatee (J. D. Johnson).

BOLIVIA Beni : 45 km SSW San Borja, 270 m (P. S. Ward); 46 km SSW San Borja, 300 m (P. S.
Ward); Cavinas (W. M. Mann); Est. Biol. Beni, 42 km E San Borja, 210 m (P. S. Ward); Ivon (W. M.
Mann); La Paz: Covendo (W. M. Mann); Ixiamas (W. M. Mann); Sta. Helena (W. M. Mann).

BRAZIL AC: Rio Branco (M. Alvarenga); AM: “Amazonas” (Bates); “Faz. NAFG, Est. Mr. Carac. km.
31” (c.u.); Arquipelago Anavilhanas, Rio Negro (R. Negrett); Benjamin Constant & vicinity (W. L. Brown);
Cucuf, Ponto 08 (D. Dias); E.E.S.T., km. 44, BR-174, [near] Manaus (c.u.); Faz. Dimona, 80 km N Manaus,
80 m (P. S. Ward); Faz. Esteio, 80 km NNE Manaus, 80 m (P. S. Ward); Humaita (P. da Silva & D. Araujo);
Ig. Jaraqui, Marg. esq., Rio Negro (Exp. Depto. Zool.); Ig. Marianil, Rio Branco Rd„ 24 km NE Manaus (W.
L. Brown); Manaus (A. Y. Harada; K. Lenko); Manaus-Caracaraf, km. 61 (W. W. Benson); Praia Grande,
Rio Negro, Manaus (W. W. Benson); Reserva Ducke, NE of Manaus (W. L. Brown); Rio Taruma
Mirim-Igapo (J. Adis); km. 24, ZF-3, Faz. Esteio, near Manaus (A. Y. Harada); km. 24, ZF-3, near Manaus
(W. W. Benson; A. Y. Harada); km. 27, ZF-3, near Manaus (W. W. Benson); km. 44, BR-174, near Manaus
(W. W. Benson); km. 44.5, BR-174, near Manaus (W. W. Benson); BA: Una (M. C. Alves); ES: Linhares
(M. Alvarenga); GO: Araguacema (A. Y. Harada; J. A. Rafael); Jatai (F. M. Oliveira); MT: Barra do
Tapirape (B. Malkin); Sinop (Alvarenga & Roppa; M. Alvarenga); Utiariti, Rio Papagaio, 325 m (Lenko &
Pereira); Vila Vera (M. Alvarenga); Xingu (Alvarenga & Wemer); PA: “Para” (Schulz.); Belem (K. Lenko;
N. Rosa); Belem-Tapana (A. Y. Harada); C. Araguaia (A. Y. Harada; J. A. Rafael); Cach. Breu (Sampaio);
Caldeirao, R. Itacaiunas (C. R. F. Brandao & W. W. Benson); Jacareacanga (M. Alvarenga); Oriximina (A.
Y. Harada); Oriximina, Alcoa C. Alta (A. Y. Harada); S. Norte, Carajas (C. R. F. Brandao & W. W.
Benson); Sa. dos Carajas (W. W. Benson); Santarem (Bates); Tucurui (W. L. Overal); Utinga tract, nr.
Belem (P. F. Darlington); RD: Porto Velho (J. Arias); Porto Velho, Rio Madeira (W. M. Mann & Baker);
Rio Madeira, Madeira-Mamore R. R. Co. Camp 39 (W. M. Mann & Baker); S. Antonio de Guapore (c.u.);
Vilhena (M. Alvarenga; A. Y. Harada); RR: Rio Uraricufra leha Manica (R. Negrett).

COLOMBIA Cundinamarca: Medina (F. Fernandez); Magdalena: 2 km ESE Minca, 780 m (P. S.
Ward); Naranjo (A. Forel).

COSTA RICA “Costa Rica” (Tonduz); Alajuela: 11 mi N Florencia (D. H. Janzen); Cartago: Turrialba
(D. H. Janzen); Guanacaste: 12 km N Liberia (D. H. Janzen); Finca Taboga, 6 mi. S, 6 mi. W Canas (D. H.
Janzen); Heredia: 10°20’N, 84°04'W, 500 m (J. T. Longino); F. La Selva, 3 km S Pto. Viejo (H. A.

Hespenheide); P. N. Braulio Carrillo, 500 m (J. T. Longino); Limon: Estrella Valley (G. C. Wheeler);

Hamburg* Farm, Santa Clara (F. Nevermann); Jimenez (Alfaro?); La Lola (N. A. Weber); Limon (N. L. H.
Krauss); Matina (A. Alfaro); Moin (D. H. Janzen); Parismina Br., Santa Clara (F. Nevermann); Portete (D.
H. Janzen); Suerre, Jimenez (Alfaro?); Puntarenas: 1 km NE Tarcoles, 20 m (P. S. Ward); 1 km N Palmar
Sur (D. H. Janzen); 20 km NE Palmar Sur (D. H. Janzen); 1-5 mi. NW Rincon (D. H. Janzen); Golfito (W.
S. Ross; Truxal & Menke); Llorona, Corcovado Natl. Pk., 100 m (P. S. Ward); Llorona, Corcovado Natl.
Pk., 10 m (P. S. Ward); Llorona, Corcovado Natl. Pk. (J. T. Longino); Manuel Antonio Natl. Pk. (L.

Masner); Manuel Antonio Natl. Pk., 10 m (P. S. Ward); Manuel Antonio Natl. Pk., 20 m (P. S. Ward);

Manuel Antonio Natl. Pk., 5 m (P. S. Ward); Manuel Antonio Natl. Pk., <5 m (P. S. Ward); Reserva Biol.
Carara, 40 m (P. S. Ward); Reserva Biol. Carara, 500 m (P. S. Ward); Sirena, Corcovado Natl. Pk. (J. T.
Longino); Sirena, Corcovado Natl. Pk., 0-100 m (J. T. Longino); Sirena, Corcovado Natl. Pk., 50 m (J. T.
Longino); San Jose: 3.5 km NE Santiago de Pur (D. H. Janzen); 40 km S San Isidro de General (D. H.
Janzen).

ECUADOR prov. unknown: Piedrero (M. Deyrup).

FRENCH GUIANA Cayenne (Pillault); Saul (G. H. Perrault).

GUATEMALA Alta Verapaz: Cacao Trece Aguas (Barber & Schwarz); Izabal: Lago Izabal, 1.5 km NE
El Estor (D. H. Janzen); Suchitepequez: Moca (W. M. Wheeler).

GUYANA East Demerara - West Coast Berbice: Timehri, 0-50 m (J. T. Longino); Mazaruni-P otaro:
Bartica (W. Beebe); Kartabo, Bartica Dist. (W. M. Wheeler); Oko R., Cuyuni trib. (N. A. Weber); Rupununi:
Karanambo, 100 m (J. T. Longino); Tutwau V., 100-300 m (J. T. Longino).

HONDURAS Atlantida: La Ceiba (F. J. Dyer); Tela (W. M. Mann); Colon: Corocito (c.u.); Cortes:
Lago Yojoa, 680 m (W. L. Brown).

Pseudomyrmecine Ants

425

MEXICO Chis.: 10 km S Palenque (M. Arrendondo); Q. Roo: San Miquel, Cozumel I. (N. L. H.
Krauss); Vallarta (A. Dejean); Tabasco: 3 mi. W Cardenas (D. H. Janzen); Ver.: “St. Lucrecia, Teh.” (W. M.
Mann); 6 mi. NW Salinas (E. Schlinger); Los Tuxtlas (R. L. Jeanne); Los Tuxtlas, 10 km NNW
Sontecomapan, 200 m (P. S. Ward); Los Tuxtlas, 10 km NNW Sontecomapan, 500 m (P. S. Ward).
NICARAGUA Matagalpa: El Tuma, 30 km E Matagalpa (C. Kugler & J. Hahn).

PANAMA Canal Zone: 3 km WNW Gamboa, 60 m (P. S. Ward); 5 km WNW Gatun Dam, 160 m (P. S.
Ward); 6 km NW Gamboa, 50 m (P. S. Ward); Barro Colorado I. (N. Banks; W. L. Brown & E. S.
McCluskey; H. Emerson; P. B. Kannowski, R. B. Root & W. L. Brown; R. B. & L. S. Kimsey; J. Reiskind;
C. W. Rettenmeyer; W. M. Wheeler; H. Wolda; c.u.); Cerro Galera, 9 km W Panama City, 200 m (W. L.
Brown et al.); Fort Kobbe (C. Campbell); Frijoles (W. M. Wheeler); Margarita Swamp (J. Zetek); Puma
Island (R. B. Kimsey); vie. Pta de los Chivos, 3 km SW Gatun, 100 m (W. L. Brown et al.); Summit (N. L.

H. Krauss); Chiriqui: Progreso (F. M. Gaige); Darien: Cana, 500 m (D. M. Olson); Panama: Bella Vista (N.
Banks); Cerro Campana, 800-950 m (G. B. Fairchild & W. L. Brown).

PERU “Peru” (c.u.); Junln: Chanchamayo, Anashirone R. (C. A. Portocarrero); El Campamiento, Col.
Perene (J. C. Bradley); Satipo (W. F. Walsh); Loreto: 15 km WSW Yurimaguas, 200 m (P. S. Ward); Yurac,
67 mi E Tingo Maria (E. I. Schlinger & E. S. Ross); Madre de Dios: 30 km SW Pto. Maldonado, 290 m (T.
L. Erwin et al.); San Martin: Convento, 26 km NNE Tarapoto, 220 m (P. S. Ward); Davidcillo, 30 km NNE
Tarapoto, 220 m (P. S. Ward).

TRINIDAD “Trinidad” (N. A. Weber; c.u.); Nariva Swamp (N. A. Weber); Palo Seco (H. Morrison).
VENEZUELA Barinas: 17 km SSW Ciudad Bolivia, 240 m (P. S. Ward); Bolivar: 44 km ENE
Tumeremo, 200 m (P. S. Ward); Campamento Rio Grande, 250 m (P. S. Ward); Rio Cuyuni, 66 km SSE El
Dorado, 250 m (P. S. Ward); Carabobo: San Esteban (W. L. & D. E. Brown); T. F. Delta Amacuro?:
“Orinico Delta” (N. A. Weber).

Pseudomyrmex pisinnus sp. nov.

(Figures 11a, lib)

Holotype worker. — BRAZIL, SP: Monte Aprazfvel, Faz. Bacuri [20°45'S,
49°42'W], 19.vii. 1974, in Eupatorium stem, M. Dimiz, no. 644 (MZSP). HW 0.51,
HL 0.78, EL 0.41, PL 0.30, PH 0.22.

Paratypes. — Same data as holotype, four workers (LACM, MCZC, MZSP,
PSWC). Additional, non-paratypic material is listed below.

Worker measurements (n=8). — HL 0.75-0.78, HW 0.47-0.51, MFC 0.009-0.016, Cl
0.61-0.68, OI 0.49-0.55, REL 0.52-0.54, REL2 0.78-0.86, OOI 0.73-1.58, VI 0.76-0.87, FCI 0.018-0.033,
SI 0.41-0.45, SI2 0.50-0.54, FI 0.46-0.52, PDI 1.46-1.69, MPI 0.029-0.047, NI 0.59-0.64, PLI 0.67-0.77,
PWI 0.54-0.65, PPWI 1.07-1.20.

Worker diagnosis. — Very small species (HW 0.47-0.51) with elongate head (Fig. 11a; Cl
0.61-0.68), and short scapes relative to eye length (SI2 0.50-0.54); sides of head subparallel; occipital
margin flat to slightly concave, in frontal view; funicular segments II and III notably broader than long (FLI

I. 16-1.48, n=4). Fore femur broad; metanotal groove rather weakly impressed; basal face of propodeum
rounding into the much shorter declivitous face. Petiolar node (Fig. lib) rather long and low, the dorsum
broadly rounded in lateral profile (PLI 0.67-0.77); postpetiole about as broad as long.

Head sublucid, covered with numerous piligerous punctures; punctures relatively coarse but appearing
somewhat effaced, and separated (at least on the upper half of the head) by shiny interspaces of one or more
diameters’ distance, especially in the region immediately posterior to the compound eye. Mesosoma weakly
punctate to coriarious-imbricate, sublucid dorsally. Petiole, postpetiole, and gaster subopaque to sublucid,
covered with numerous, fine piligerous punctures. Erect pilosity and appressed pubescence present on most
of the body; erect hairs present on the pronotum, mesonotum and propodeum, but relatively short, sparse,
and inconspicuous, except for one pair on the pronotal shoulders and a second pair above the juncture of the
basal and declivitous faces of the propodeum. Body light orange-brown to yellow-brown, with darker
infuscation on the anterior third of the first gastric (fourth abdominal) tergite, on the succeeding gastric
tergites, and on part of the hind femur; mandibles paler luteous.

Quaest. Ent., 1989, 25 (4)

426

Ward

Comments. — This small, slender species is easily recognized by the combination
of small size, elongate head, long basal face of propodeum, and low petiole (see
worker key and description above). The short scapes (relative to head length),
sublucid appearance, and light orange-brown color are also distinctive.

Biology. — The type series was collected in a dry stem of Eupatorium sp. The
Peruvian specimens are from primary rain forest.

Material examined. — Type material listed above plus the following (USNM,
PSWC):

PERU Huanuco : Panguana, 220 m, 9°37'S, 74°56'W (M. Verhaagh); Madre de Dios : Rio Tambopata
Res., 30 km SW Pto. Maldonado, 290 m, 12°50'S, 69°20’W (T. L. Erwin et al.).

Pseudomyrmex schuppi (Forel)

(Figure 8)

Pseudomyrma schuppi Forel, 1901a:298. Syntype workers, Sao Leopoldo, Rio Grande do Sul, Brazil

(Schupp) (MCSN, MZSP, NHMV) [Examined].

Pseudomyrma schuppi var. confusior Forel, 1901a:299. Syntype workers, Rio Grande do Sul, Brazil

(Schupp) (MHNG) [Examined] [Synonymy by Kempf, 1961:380; here confirmed],

Pseudomyrma schuppi var. geraensis Forel, 1912:23. Syntype workers, Juiz de Fora, Minas Gerais, Brazil

(Sampaio) (MHNG) [Examined] [Synonymy by Kempf, 1961:380; here confirmed].

Pseudomyrmex schuppi (Forel); Kempf, 1961:380 [Differentiating characters of worker and queen.]

Worker measurements (n = 14). — hl 0.96-1.18, hw 0.82-0.93, MFC 0.023-0.038, Cl
0.79-0.88, OI 0.55-0.64, REL 0.52-0.56, REL2 0.62-0.67, OOI 0.23-0.60, VI 0.75-0.84, FCI 0.026-0.042,
SI 0.45-0.48, SI2 0.68-0.76, FI 0.39-0.45, PDI 1.16-1.34, MPI 0.037-0.053, NI 0.53-0.62, PLI 0.94-1.07,
PWI 0.85-0.99, PPWI 1.32-1.60.

Worker diagnosis. — Relatively large species (HW 0.82-0.93), with broad head (Cl 0.79-0.89),
and eyes of moderate size; sides of head broadly convex, rounding into the occipital margin which varies
from flat to slightly concave; funicular segments II and III about as long as broad (FLI 1.71-2.12, n=9). Fore
femur, metanotal groove, propodeum, and postpetiole similar to those of P. oculatus (q.v.), except basal and
declivitous faces of propodeum tend to be more clearly differentiated. Petiolar node (Fig. 8) high and
broadly rounded in lateral view (PLI 0.94-1.07), very broad in dorsal view (PWI 0.85-0.99), with relatively
strong dorsolateral margination; anteroventral process of petiole conspicuous, tooth-like, often bluntly
recurved.

Head densely punctate, opaque; mesosoma obscurely punctate to coriarious-imbricate, opaque to
subopaque; petiole, postpetiole, and gaster subopaque, covered with numerous fine piligerous punctures.
Erect pilosity and appressed pubescence common and conspicuous on most parts of the body including the
mesosoma dorsum. Body medium to dark brown, the antennae and tarsi (and sometimes mandibles and
promesonotum) lighter in color.

Comments. — P. schuppi is diagnosed minimally by its relatively large size (HW
>0.81) and high, broad petiole (PLI > 0.93, PWI > 0.84) whose dorsum is smoothly
rounded in lateral profile. Also characteristic of P. schuppi are the conspicuous
elevation of the basal face of the propodeum above the level of the mesonotum and
the posterior displacement of the postpetiolar node as seen in lateral view. These last
two features are shared with P. oculatus , from which P. schuppi is distinguished by
petiole shape (see above and compare Figs. 7 and 8) and differences in relative eye
length (compare REL2).

Biology. — I have no field experience with this species and the museum material
which I have examined contains no biological information. Luederwaldt (1926)

Pseudomyrmecine Ants

427

recorded P. schuppi nesting under epiphytes.

Material examined (BMNH, ICCM, LACM, MCSN, MCZC, MHNG, MZSP,
NHMB, NHMV, PSWC, UCDC, USNM, WWBC).—

ARGENTINA Misiones : Iguazi (N. Kusnezov).

BRAZIL AC: Uranduique, Rio Branco (Machado & Pereira); BA: “Bahia” (c.u.); Encruzilhada, 960 m
(Seabra & Alvarenga); Encruzilhada, 980 m (Seabra & Alvarenga); DF: Parque Nac. da Brasilia (W. L. &
D. E. Brown); ES: Vila Velha (O. Seifert); MG: B. Horizonte (O. Monte); Juiz de Fora (Sampaio);
Monsenhor Paulo (V. dos Santos); Passo Quarto (Sampaio); Pedra Azul, 800 m (Seabra & Alvarenga); S.
Caraga (Engenho), 800 m (Martins & Silva); Tiradentes (R. L. Araujo); Tres Cora§oes (E. Azcanjo); MG?:
Tejuca (c.u.); PR: Col. Esperan§a, Arapongas (W. W. Kempf); Rolandia (W. W. Kempf); RJ: “Prov.Rio”
(Goldi); “Rio Janeiro” (c.u.); Corcovado (Alvarenga & Seabra; c.u.); Fonseca, Niteroi (C. R. Gon9alves);
Ilha Jaguanum (D. Hunt); Ilha da Gipoia, Angra dos Reis (T. Borgmeier); Monumento Rodoviario (C. R.
Gon9alves); Monumento Rodoviario, Via Dutra (C. R. Gon9alves); Petropolis (c.u.); Toco Seco (R. L.
Araujo); RS: “Rio Gr. do Sul” (Schupp); Laurentino (R. Muller); Pared Novo (Hansen); Sao Leopoldo
(Schupp); SC: Blumenau (R. L. Araujo; R. Muller; Reichensperger); Florianopolis (R. Muller); Gaspar (S.
Fontes; R. Muller); Hamonia (Leuderwaldt); Itajaf (S. Fontes); Nova Teutonia (F. Plaumann); Poco Grande,
Gaspar (R. Muller); Que9aba (T. Zimmermann); Rodeio (R. Muller); SP: “Mogy” (c.u.); Agudos (W. W.
Kempf; R. Muller); Anhembi, Faz. Barr. Rico (W. W. Kempf et al.; L. Travassos F.); Atibaia (W. W.
Benson); Barueri (K. Lenko); Botucatu (Goldi); Campo Limpo (W. W. Kempf); Caraguatatuba (Res. Flor.),
40 m (Exp. Depto. Zool.); Caraguatatuba (Res. Flor.), 680 m (K. Lenko); Embu (F. Lane); Est. Biol.
Boraceia, Salesopolis (K. Lenko); Faz. Itaquere, Boa Esperan9a do Sul (K. Lenko); Faz. Itaquere, Nova
Europa (K. Lenko); Faz. Itaquere, Tabatinga (K. Lenko); Guaratingueta (W. W. Kempf); H. Florestal, S.
Paulo (F. C. Val); Ilha do Cardoso (L. Foneris); Len9ois Pta. (R. Miiller); Rio Claro, Horto Florestal (V.
Gama); Rio Manso (c.u.); Rodov-Ctba, km. 40 (W. W. Kempf); S. Rogue (F. Lane); S. Sebastiao (W. W.
Kempf); S. Sebastiao, Bairro S. Francisco (W. W. Kempf); Sao Paulo (Parker); Ubatuba (P. C. Montouchet).
PARAGUAY Asuncion (P. Duelli); Pastoreo (P. Duelli); San Bernardino (Fiebrig).

Pseudomyrmex urbanus (F. Smith)
(Figures 12a, 12b, 28, 29, 30)

Pseudomyrma urbana F. Smith 1877:65. Syntype workers, Ega, Amazonas, Brazil (BMNH) [Examined];
one worker here designated as LECTOTYPE.

Pseudomyrma chodati Forel 1920:201. Syntype workers, Paraguay (Chodat) (MHNG) [Examined]. One
worker here designated LECTOTYPE. Syn. nov.

Pseudomyrma ogloblini Santschi, 1936:402. Syntype workers, Loreto, Misiones, Argentina (A. Ogloblin)
(MCZC, MZSP, NHMB) [Examined]. Syn. nov.

Worker measurements (n=21). — HL 0.76-0.95, HW 0.58-0.66, MFC 0.014-0.026, Cl
0.69-0.76, OI 0.52-0.58, REL 0.51-0.56, REL2 0.69-0.78, OOI 0.29-1.12, VI 0.76-0.85, FCI 0.022-0.041,
SI 0.44-0.48, SI2 0.58-0.67, FI 0.44-0.53, PDI 1.08-1.53, MPI 0.028-0.063, NI 0.51-0.65, PLI 0.76-0.88,
PWI 0.55-0.71, PPWI 1.06-1.48.

Worker diagnosis. — A relatively small species (HW 0.58-0.66), with moderately elongate eyes
(REL2 0.69-0.78); sides of head (Fig. 12a) subparallel to broadly convex, rounding somewhat gradually into
the flat to weakly concave occipital margin; second and third funicular segments broader than long (FLI
1.15-1.58, n=7). Fore femur relatively broad; metanotal groove well marked; basal face of propodeum
subequal to, or longer than, the declivitous face and usually rounding gently into the latter. Petiole (Fig. 12b)
longer than high, with soft dorsolateral margination, and a conspicuous anteroventral process; postpetiole
broader than long. Mandibles striato-punctate; head densely punctate, sublucid, the punctures often
separated by shiny interspaces, especially between the ocelli and compound eye, and immediately posterior
to the compound eye, where some punctures are separated by half their diameters or more. Mesosoma
punctate to coriarious-imbricate, tending to be sublucid dorsally and on the side of the pronotum, more
opaque elsewhere. Petiole, postpetiole, and gaster sublucid, the lustre dulled by numerous fine piligerous
punctures. Fine erect pilosity and appressed pubescence present on most of body, including mesosoma
dorsum. Body usually light to medium orange-brown, with varying degrees of infuscation of the gaster,

Quae st. Ent., 1989, 25 (4)

428

Ward

hindfemur, and (sometimes) head; mandibles and tarsi paler. In a few individuals (Dpto. Beni, Bolivia; Isla
Fernando Noronha, Brazil) entire body darker brown.

Comments. — P. urbanus belongs to a difficult species complex that also
includes P. caeciliae and P. cubaensis ( s.l. ). P. urbanus workers are characterized
by a combination of intermediate size (HW 0.58-0.66), relatively long eyes and
broad fore femur (see Figs. 28, 29), and (typically) orange-brown body. These
differences are detailed in the keys and in the discussions under P. caeciliae and P.
cubaensis. P. urbanus differs from P. elongatus (cf. Figs. 6 and 12a) by its shinier
appearance, particularly the presence of conspicuous sublucid areas on the head and
pronotum, and by its lighter color. The eyes of P. urbanus workers tend to be shorter
than those of P. elongatus , and the petiole is relatively lower.

Some problematic series are in the material listed below. These include
unusually dark brown workers from Dpto. Beni, Bolivia and large cubaensis- like
workers from Ecuador (Piedrero) and Venezuela (Edo. Zulia). The type specimens
of P. chodati (Forel) and P. ogloblini (Santschi) from Paraguay and Argentina,
respectively, appear to be rather typical P. urbanus insofar as they are in the
appropriate size range (HW 0.58-0.61) and have a light orange -brown, sublucid
integument; the eyes, however, are rather short (REL2 0.69-0.72). Future studies
may reveal more than one species here.

Biology. — Little biological information is available about P. urbanus.
Collections come from rain forest localities as well as areas typified by drier forest.
The type series of P. chodati was collected in cauline swellings of Cordia longituba,
that of P. ogloblini in the branches of “ Thecoma ipe ” [ = Tecoma obtusata ]
accompanied by coccids; both records suggest a propensity for this species to nest in
live plant cavities.

Material examined ( BMNH , CKIC, CUIC, GHPC, JTLC, KWJC, LACM, MCZC,
MHNG, MZSP, NHMB, PSWC, UCDC, USNM).—

ARGENTINA Misiones : Loreto (A. A. Ogloblin); Salta: Urundel (c.u.); Tucuman : Tucuman (W.
Weyrauch).

BOLIVIA Beni : 46 km SSW San Borja, 300 m (P. S. Ward); La Paz: Espia Rio Bopi (W. M. Mann).

BRAZIL AM: Ega (Bates?); FN: Ilha Fernando Noronha (O. Roppa); GO: Anapolis (W. W. Kempf);
Jatai (F. M. Oliveira); MA: Ilha de Balsas reg. (Westminster School Exp.); MG: Pedra Azul (Seabra &
Alvarenga); Pirapora (E. Garbe); Varginha (Macais); MS: Corumba (D. Hunt); Faz. Canaa, Tres Lagoas (F.
Lane); MT: Sinop (M. Alvarenga); Utiariti, Rio Papagaio, 325 m (K. Lenko); Vila Vera (M. Alvarenga); PA:
C. Araguafa (J. A. Rafael); Rio Trepecuru (Sampaio); RD: Vilhena (M. Alvarenga); SP: Agudos (C. Gilbert;
W. W. Kempf); Faz. Itaquere, Boa Esperanga do Sul (K. Lenko); Piracicaba (C. A. Triplehom).

COLOMBIA Guajira: Serr. de Macuira, 6-8 km S Nazareth, 70-200 m (W. L. Brown & C. Kugler);
Meta: R. Guayabero, Angostura 1, 270 m (C. Kugler); Vichada: Rio Tomo (W. W. Lamar).

ECUADOR Pichincha: Sto. Domingo (S. & J. Peck); prov. unknown: Piedrero (M. Deyrup).

FRENCH GUIANA 50 km S Cayenne (G. H. Perrault).

GUYANA Rupununi: Karanambo, 100 m (J. T. Longino).

PANAMA Canal Zone: Summit (N. H. L. Krauss); Chiriqui: “Prov.Chiriqui” (F. M. Gaige).

PARAGUAY: “Paraguay” (Chodat); Pastoreo (P. Duelli).

PERU Loreto: 15 km WSW Yurimaguas, 200 m (P. S. Ward); Madre de Dios: 30 km SW Pto.
Maldonado, 290 m (T. L. Erwin et al.).

SURINAM Moengo, Cottica R. (c.u.); Paramaribo (c.u.).

Pseudomyrmecine Ants

429

TRINIDAD Curepe (F. D. Bennett); Maracas Valley (N. A. Weber).

VENEZUELA Aragua : 2 km N Ocumare de la Costa (A. S. Menke & L. Hollenberg); Bolivar:
Campamento Rio Grande, 250 m (P. S. Ward); T. F. Amazonas: P. N. Duida-Marahuaka, Culebra, 250 m
(K. Jaffe); T. F. Delta Amacuro?: “Orinoco Delta” (N. A. Weber); Zulia: El Tucuco, Perija (R. W. Brooks et
al).

PSEUDOMYRMEX SUBTILISSIMUS GROUP

Introduction

Diagnosis (worker). — Small species (HW 0.48-0.73), with subopaque, densely
punctulate, and elongate head (Cl 0.56-0.73); mandibles as in the P. oculatus group;
frontal carinae distinctive; subcontiguous (FCI 0.023-0.066), raised anterodorsally,
and merging insensibly into the clypeus (rather than fusing with the antennal
sclerite) (Fig. 18), so that in lateral view they form a continuous curve with the
median clypeal lobe (Fig. 19); the latter tectiform and protruding, with the anterior
margin weakly flared and sharp-edged laterally, but not medially; in dorsal view,
median clypeal lobe broadly rounded anteriorly; scapes very short (SI2 0.44-0.59);
funicular segments II and III much broader than long (FLI 0.96-1.30); fore femur
notably broadened (FI 0.44-0.60); petiole relatively low, long, and slender (PLI
0.54-0.75, PWI 0.48-0.68), without a differentiated peduncle and node, appearing
dorsally flattened in lateral profile. Erect pilosity very sparse, lacking on the
antennae, legs, and most of head and mesosoma; a characteristic pair of rather long
setae present on the pronotal humeri, on the propodeum at the juncture of the basal
and declivitous faces, and on the petiolar dorsum. Pubescence covering most of the
body, exceptionally fine and inconspicuous in three of the four species. Palp
formula: 6,3.

Comments. — This is a small, discrete group of species, uniquely characterized
by the configuration of the frontal carinae and the clypeus. The elongate head, short
scapes, flattened apedunculate petiole, and patterns of pilosity are also distinctive.

Features shared with the P. oculatus group include the protruding, tectiform,
median clypeal lobe; palp formula of 6, 3; and elongate head and eyes. In addition
preliminary observations suggest similarities in the male genitalia which, together
with the shared worker characters, support the notion of a close relationship between
these two groups.

Synonymic List of Species

P. spiculus, sp. nov. Mexico to Peru, Bolivia
P. subtilissimus (Emery, 1890). Nicaragua, Costa Rica
P. tenuissimus (Emery, 1906), stat. nov. Mexico to Peru, Brazil
= P. culmicola (Forel, 1912), syn. nov.

P. villosus, sp. nov. Brazil

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430

Ward

Key to Species (workers and queens)

Note. — Only a few queens have been examined (unknown in P. villosus ) and the
ranges of measurements and indices for queens may be somewhat broader than the
known limits given here.

1 Small species (HW < 0.67) with elongate head and eyes

(worker Cl 0.56-0.71, queen Cl 0.43-0.61; worker REL2
0.76-0.92, queen REL2 0.84-1.15) (Figs. 32, 34, 36);

pubescence very short and inconspicuous, tightly
appressed to body 2

1' Larger species (HW 0.73, in unique worker type), with

relatively broad head and short eyes (Cl 0.73, REL2 0.72) (Fig.

38); body pubescence conspicuous, decumbent to suberect
(Brazil) P. villosus, sp. nov., p. 434

2 (1) Very small species (worker HW 0.48-0.53; queen HW

0.42-0.51, n=5), with elongate head (worker Cl 0.56-0.62;
queen Cl 0.43-0.53) and slender petiole (worker PLI

0.54-0.66; queen PLI 0.55-0.62) (Figs. 36, 37) (Mexico to

Peru, Brazil) P. tenuissimus (Emery), p. 432

2' Larger species (worker HW 0.55-0.66; qaeen HW 0.56-0.63,

n=6), with broader head (worker Cl 0.60-0.71; queen Cl
0.55-0.61); if worker HW < 0.60, then petiole generally rather
short (worker PLI 0.65-0.75) 3

3 (2') Smaller species (worker HW 0.55-0.59; queen HW 0.56, 0.57,

n=2) with more elongate head (worker Cl 0.60-0.65, queen Cl
0.55, 0.57) and with relatively short petiole (worker PLI
0.65-0.75, queen PLI 0.63, 0.68) (Figs. 34, 35) (Nicaragua,

Costa Rica) P. subtilissimus (Emery), p. 432

3' Larger species (worker HW 0.61-0.66; queen HW 0.61-0.63,

n=4), with broader head (worker Cl 0.66-0.71; queen Cl

-

0.60-0.61) and slender petiole (worker PLI 0.56-0.67; queen

PLI 0.56-0.61) (Figs. 32, 33) (Mexico to Peru, Brazil)

P. spiculus, sp. nov., p. 431

Species Accounts

Pseudomyrmex spiculus sp. nov.

(Figures 31, 32, 33)

Pseudomyrmex subtilissimus tenuissimus ; Kempf (nec Emery) 1972:224 (partim).

Holotype Worker. — COSTA RICA, Prov. Puntarenas: Reserva Biologica
Carara, 500m, 9°47'N, 84°36'W, 26.viii.1985, on low vegetation, old field/pasture,

Pseudomyrmecine Ants

431

P. S. Ward acc. no. 7668-9 (MCZC). HW 0.63, HL 0.89, EL 0.50, PL 0.37, PH

0.22.

Paratype Workers, Queens. — Same data as holotype, two workers; COSTA
RICA, Prov. Puntarenas : Manuel Antonio Natl. Park, 10m, 9°23'N, 84°09'W,
28.vii.1985, P. S. Ward acc. no. 7715-6, 1 worker; same locality, 40m, 27.vii.1985,
P. S. Ward acc. no. 7679, 1 dealate queen; Llorona, Corcovado Natl. Park, 10m,
10-14.V.1979, P. S. Ward acc. no. 3389-6, 1 worker; Sirena, Corcovado Natl. Park,
50m, 8°28'N, 83°35'W, 22.iv.1981, J. T. Longino, 1 worker; same locality,
20.xii. 1 98 1 , J. T. Longino, 1 worker; COSTA RICA, Prov. Limon : Portete,
13. ii. 1965, D. H. Janzen, 1 worker. (JTLC, LACM, MNCR, MZSP, PSWC,
USNM).

Type series is restricted to the above material from Costa Rica. Additional,
apparently conspecific material comes from Mexico, Panama, Colombia, Guyana,
Brazil, and Peru (see “Material examined” below).

Worker measurements (n=14). — HL 0.88-0.98, HW 0.61-0.66, MFC 0.022-0.042, Cl
0.66-0.71, OI 0.55-0.61, REL 0.53-0.57, REL2 0.76-0.83, OOI 0.04-0.67, VI 0.79-0.87, FCI 0.034-0.064,
SI 0.38-0.43, SI2 0.46-0.54, FI 0.48-0.55, PDI 1.20-1.61, MPI 0.029-0.059, NI 0.60-0.71, PLI 0.56-0.67,
PWI 0.48-0.59, PPWI 1.14-1.37.

Worker diagnosis. — Small, slender species (HW 0.61 - 0.66); head (Fig. 32) elongate, although
broader than that of P. tenuissimus (Cl 0.66-0.71); eyes elongate, almost reaching the level of the median
ocellus; occipital margin concave in full face, dorsal view; lateral margins of pronotum rounded; fore femur
short and broad (FI 0.48-0.55); metanotal groove narrow, shallow; basal face of propodeum convex,
declivitous face flat to weakly concave, the former about 1.5 times the length of the latter; basal face of
propodeum rounding into the declivitous face at a slight (obtuse) angle, marked by a pair of erect setae;
petiole (Fig. 33) slender, low, notably longer than wide (Fig. 31; PLI 0.56-0.67); postpetiole short, globose
in lateral view, subtriangular in dorsal view, about as wide as long, and twice as wide as the petiole (PWI2
0.51-0.57). Mandibles finely striate with scattered punctures, sublucid; head, mesosoma, and petiole densely
punctulate to coriarious-imbricate, opaque; postpetiole and gaster opaque, with numerous fine piligerous
punctures. Erect pilosity very sparse (except on apex of gaster), lacking on the antennae, sides of head, upper
half of gula, femora (except for one to several setae on the ventral surface of the fore femur), tibiae, tarsi,
and most of mesosoma. Two or three pairs of short erect setae on dorsum of head; a conspicuous pair of long
(ca. 0.15 mm) erect setae on the anterolateral comers of pronotum, at the juncture of the basal and
declivitous faces of the propodeum, and on the posterior dorsum of both the petiole and postpetiole; the last
two pairs often accompanied by a pair of shorter, posterolateral setae. Appressed pubescence common on
most of body, but rather fine and inconspicuous. Grey-brown; mandibles, fronto-clypeal complex, antennae,
tarsi, pronotum, petiole, and postpetiole paler luteous-brown.

Comments. — The combination of relatively large size, broad head, and slender
petiole serves to distinguish P. spiculus from the other species in the P. subtilissimus
group (see Figs. 31-39).

Biology. — The species is widespread but infrequently collected, and little is
known about its biology. Most collections are based on single individuals foraging
on low vegetation or tree-falls, in rain forest, rain forest edge, littoral forest, and
mangrove. In Costa Rica I collected a single dealate (colony-founding?) female in a
loose dead twig, hanging in low vegetation, in coastal rain forest. Type workers
from Reserva Biologica Carara were foraging on the trunk and associated vegetation
of a strangler fig tree ( Ficus oerstediana) and its unidentified host tree, in an old
pasture. At this location, workers of P. tenuissimus were present on the same

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432

Ward

vegetation, foraging in close proximity to those of P. spiculus. These two species
have been collected sympatrically elsewhere, and their ranges are broadly
coextensive.

Material examined. — Type material listed above, plus the following (INPA,
LACM, MCZC, MZSP, PSWC, UCDC):

BRAZIL AM: Manaus to Itacoatiara Hwy., km 34 (W. L. & D. E. Brown), MT: Sinop 55°37'W, 12°31'S
(M. Alvarenga); RD: Vilhena (A. Y. Harada).

COLOMBIA Magdalena'. 4km N San Pedro, 550m (P. S. Ward); Canaveral, < 5m (P. S. Ward).

GUYANA Mazaruni-Potaro: Kartabo (W. M. Wheeler); Rupununi : Upper Essequebo (J. G. Myers).

MEXICO Jal:. 14 km SSE Chamela, 20 m (P. S. Ward).

PANAMA Canal Zone'. 5 km WNW Gatun Dam, 160 m (P. S. Ward); Barro Colorado I. (N. Banks; R.
B. & L. S. Kimsey); Colon: Colon (E. Adams).

PERU Madre de Dios: Rio Tambopata Reserve, 30km SW Pto. Maldonado, 290m (T. L. Erwin et al.).

Pseudomyrmex subtilissimus (Emery)

(Figures 31, 34, 35)

Pseudomyrma subtilissima Emery 1890:65. Syntype workers, one dealate queen, Alajuela, Costa Rica

(Alfaro) (MCSN) [Examined]. One syntype worker here designated as LECTOTYPE.

Worker measurements (n=17, except for FI where n=16). — HL 0.86-0.95, hw
0.55-0.59, MFC 0.020-0.032, Cl 0.60-0.65, OI 0.52-0.59, REL 0.49-0.54, REL2 0.80-0.87, OOI
0.38-0.89, VI 0.79-0.88, FCI 0.035-0.055, SI 0.41-0.47, SI2 0.50-0.57, FI 0.44-0.53, PDI 1.60-1.88, MPI
0.015-0.039, NI 0.52-0.66, PLI 0.65-0.75, PWI 0.56-0.68, PPWI 1.18-1.35.

Worker diagnosis. — Similar to P. spiculus ( q.v .) except as follows: smaller, head (Fig. 34) more
elongate (HW 0.55-0.59, Cl 0.60-0.65), eyes shorter, not reaching the level of the median ocellus (REL
0.49-0.54); basal face of propodeum flatter, meeting the declivitous face at a more distinct angle; petiole
(Fig. 35) shorter, higher (PLI 0.65-0.75). Most of body uniformly light brown; gaster dark brown, with
contrasting light brown band on anterior half of abdominal tergite IV.

Comments. — A moderately elongate head (broader than P. tenuissimus, more
elongate than P. spiculus and P. villosus’, see Fig. 31), short petiole, and light brown
color (with banded gaster) characterize P. subtilissimus.

Biology. — This species has been collected only in, or on, swollen-thorn acacias,
in association with Pseudomyrmex flavicornis (F. Smith). The collection records
suggest that it is an obligate, non-aggressive acacia ant which somehow manages to
coexist with P. flavicornis.

Material examined ( BMNH , LACM, MCSN, MHNG, MZSP, NHMV, PSWC,
USNM). —

COSTA RICA Alajuela: Alajuela (Alfaro); Puntarenas: 1km NE Tarcoles, 20m (P. S. Ward); Est. Biol.
Palo Verde, 10 m (J. T. Longino); San Jose: 3.5km NE Santiago de Pur (D. H. Janzen); Rio Oro (D. H.
Janzen); prov. unknown: “Costa Rica” (Alfaro; Cameron; c.u.).

NICARAGUA Matagalpa: 15.8 mi. W Jebaca [= Sebaco] (D. H. Janzen).

Pseudomyrmex tenuissimus (Emery) stat. nov.
(Figures 18, 19, 31,36, 37)

Pseudomyrma subtilissima subsp. tenuissima Emery, 1906:119. Holotype (unique syntype) worker,
Corumba, Matto Grosso [do Sul], Brazil, 1 5.viii. 1 900, (MCSN) [Examined].

Pseudomyrma culmicola Forel, 1912:217. Holotype (unique syntype) worker, Trinidad (Forel) (MHNG)

Pseudomyrmecine Ants

433

[Examined], Syn. nov.

Pseudomyrmex subtilissimus', Wheeler & Wheeler (nec Emery), 1973:207 [description of larva].

Worker measurements ( n=30 , except for FI where n=29). — hl 0.82-0.93, HW
0.48-0.53, MFC 0.011-0.033, Cl 0.56-0.62, 01 0.50-0.58, REL 0.48-0.53, REL2 0.84-0.92, OOI
0.22-0.89, VI 0.85-0.90, FCI 0.023-0.066, SI 0.39-0.46, SI2 0.44-0.54, FI 0.52-0.60, PDI 1.58-2.14, MPI
0.022-0.055, NI 0.55-0.64, PLI 0.54-0.66, PWI 0.49-0.60, PPWI 1.05-1.34.

Worker diagnosis.- — Similar to P. spiculus (q.v.) except as follows: distinctly smaller, head (Fig.
36) more elongate (HW 0.48-0.53, Cl 0.56-0.62). Basal face of propodeum flatter, forming a more distinct
angle with the declivitous face. Body color more uniformly brown to grey-brown, the pronotum, petiole, and
postpetiole only slightly, or not at all, contrastingly lighter.

Comments. — P. tenuissimus can be distinguished from P. subtilissimus by its
darker color, smaller size, and longer petiole (compare Figs. 35 and 37). It differs
from P. spiculus by its smaller size and more elongate head (compare HW and Cl;
see Fig. 31). The unique syntype (holotype) of P. culmicola fits easily within this
concept of P. tenuissimus.

Biology. — Although P. tenuissimus is the most common member of the P.
subtilissimus group, most collections appear to based on scattered foragers. In Costa
Rica, Panama, and Colombia, I have encountered workers foraging on vegetation in
a variety of habitats including rain forest, rain forest edge, old field/pasture, and
tropical dry forest. I have seen nests only twice: one was a small colony containing a
single dealate queen, seven workers, and brood, in the dead, fibrous twig of a woody
liana, in tropical dry forest (northern Colombia); the other was a nest of 13 workers,
16 alate queens, and brood in the dead twig of a thorny vine at the edge of disturbed
second-growth rain forest (Edo. Barinas, Venezuela). Mann (1916:426) recorded a
colony from Maranhao, Brazil “taken from beneath a loose piece of bark”, an
unusual nest-site for Pseudomyrmex as Mann himself remarks.

Material examined (INPA, IZAV, JTLC, KWJC, LACM , MCSN, MCZC, MHNG,
MZSP, PSWC, UCDC, USNM, WPMC, WWBC).—

BELIZE Belize (N. L. H. Krauss).

BOLIVIA Beni: Cavinas (W. L. Mann).

BRAZIL AM: Faz. Esteio, 80 km NNE Manaus, 80 m (P. S. Ward); Manaus, 60 m (P. S. Ward); Rio
Taruma Mirim-Igapo (J. Adis); Taruma-Mirim (J. Adis); km. 27, ZF-3, near Manaus (W. W. Benson); CE:
Itapipoca (C. R. Gonsalves); ES: Vitoria, Penha (R. Muller); GO: Jatai (F. M. Oliveira); MA: “Maranhao”
(W. M. Mann; c.u.); Bacabal (W. W. Kempf); MG: Pedra Azul (F. M. Oliviera); Arassuahy (Thieman); MS:
Corumba (c.u.); Imbirussu-Corumba (K. Lenko); MT: Sinop (M. Alvarenga); Utiariti, Rio Papagaio, 325m
(K. Lenko); Vila Vera (M. Alvarenga); PA: Belem (C. R. Gonsalves); C. Araguaia (J. A. Rafael; A. Y.
Harada); Igarape-A§u (C. R. Gon9alves); S. Norte, Carajas (W. W. Benson); RJ: Fonseca, Niteroi (C. R.
Gongalves); RN: Natal (W. M. Mann); SP: Faz. Campininha, Mogi Gua$u (H.C.M.).

COLOMBIA Caqueta: Florencia (W. P. MacKay); Cundinamarca: El Colegio & Anapoima (I. Zenner);
Huila: 15 mi. S Neiva (W. & E. MacKay); 15 mi. W Campoalegre (W. & E. MacKay); Colombia (W. & E.
MacKay). Magdalena: 8km NE Cienaga, 40m (P. S. Ward); Canaveral, 50m (P. S. Ward); Magdalena?:
locality illegible (A. Forel); Meta: Carimagua (M. Com); Valle: Dagua (W. & E. MacKay).

COSTA RICA Alajuela: 1 1 mi. N. Florencia (D. H. Janzen); Heredia: F. LaSelva, 3km S Pto. Viejo (H.
A. Hespenheide); 3 km S Pto. Viejo (J. T. Longino); Limdn: Portete (D. H. Janzen); Puntarenas: Boca
Barranca (D. H. Janzen); Llorona, Corcovado Natl. Park, 0-1 00m (J. T. Longino); Llorona, Corcovado Natl.
Park, 10m (P. S. Ward); Reserva Biol. Carara, 500m (P. S. Ward); Sirena, Corcovado Natl. Park, O-lOOm,
50m (J. T. Longino).

ECUADOR prov. unknown: Piedrero (M. Deyrup).

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434

Ward

EL SALVADOR San Salvador: Lago de Ilopango, N. side (R. Silberglied).

FRENCH GUIANA Cayenne (Pillault).

GUYANA East Demerara-West Coast Berbice: Botanic Gardens, Georgetown (H. Morrison);
Mazaruni-Potaro: Kartabo (W. M. Wheeler).

HONDURAS Cortes : Lago Yojoa, 680m (W. L. Brown).

JAMAICA Kingston (N. L. H. Krauss); St. James: Great R., 0-80m (J. T. Longino); Trelawny: Green
Park, 0-1 50m (J. T. Longino).

MEXICO Oax:. 5 mi. E Temascal (D. H. Janzen); S.L.P.: Tamazunchale (W. S. Ross); Ver.: Pueblo
Nuevo, nr. Tetzonapa (E. O. Wilson); Rio Tonto (D. H. Janzen); 29.5 mi. NW Tuxpam on Hwy. 122 (D. H.
Janzen).

NICARAGUA Matagalpa: El Tuma, 30km E Matagalpa, Hwy. 5, 330m (C. Kugler & J. Hahn).
PANAMA Canal Zone : 3km SW Gatun, 100m (W. L. Brown et al.)\ 5km WNW Gatun Dam, 160m (P.
S. Ward); Barro Colorado I. (W. M. Wheeler; H. Wolda); Cerro Galera, 9km W Panama City (W. L. Brown
et al.)\ Mt. Hope, nr. Colon (W. M. Wheeler); Ruta 1, 14km W Panama City, 100m (Brown, W. L, et at);
Panama : Chilibra R. (W. M. Wheeler); Rio Piedras (D. H. Janzen); Taboga Island (A. Busck); prov.
unknown: Lilina R. (W. M. Wheeler).

PERU Huanuco : Panguana, 200 m (M. Verhaagh); Junin: Valle Chanchamayo, 800 m (Weyrauch);
dpto. unknown: Queb. Penaga, Anotape Mts. (Frizzell)

TRINIDAD Curepe (F. D. Bennett); Mayaro Bay (N. Weber); St. Augustine (N. Weber; Darlington);
“Trinidad” (Forel).

VENEZUELA Apure: Mantecal (H. Romero); Barinas: 10km WNW Santa Barbara, 280 m (P. S.
Ward); 17 km SSW Ciudad Bolivia, 240 m (P. S. Ward); Rio Capara Res. Stn., 32 km E El Canton (J. B.
Heppner); Carabobo: Hda. El Palmar, Las Trincheras, 850m (J. & B. Bechyne); T. F. Delta Amacuro?:
“Orinoco Delta” (N. A. Weber); Zulia: El Tucuco, Perija (R. W. Brooks et al.).

Pseudomyrmex villosus sp. nov.

(Figures 38, 39)

Holotype Worker. — BRAZIL, MT: Utiariti, Rio Papagaio [ = Rio Saueruina]
[13°02'S, 58°17'W], 325m, viii.1961, K. Lenko (MZSP).

Holotype measurements. — HL 1.00, HW 0.73, MFC 0.038, EL 0.53, PL 0.38, PH 0.27, Cl 0.73,
OI 0.60, REL 0.53, REL2 0.72, OOI 0.51, VI 0.81, FCI 0.052, SI 0.43, SI2 0.59, FI 0.50, PDI 1.50, MPI
0.035, NI 0.62, PLI 0.71, PWI 0.64, PPWI 1.39.

Worker diagnosis. — A large species (for the P. subtilissimus group) with broad head and
relatively short eyes (Fig. 38). Sides of head broadly convex, rounding into the distinctly concave occipital
margin. Fore femur broad; metanotal groove rather shallow; basal face of propodeum slightly convex (lateral
view), rounding into the much shorter declivitous face. Petiole (Fig. 39) relatively short and high, with a
distinct anteroventral tooth; postpetiole broader than long.

Mandibles striato-punctate, subopaque. Head densely punctulate, sublucid, with shiny interspaces
between the fine, partially effaced punctulae. Remainder of body punctulate to coriarious-imbricate, largely
sublucid. Erect pilosity sparse, distributed as in P. spiculus iq.v.). Pubescence common on most of body,
many hairs decumbent to suberect, and thus more conspicuous than in other members of the P. subtilissimus
group. Body brown, mandibles, tarsi, petiole, postpetiole, and (to a lesser extent) pronotum and antennae
lighter (luteous-brown) in color.

Comments. — Known only from the holotype worker, this species can be
recognized by its large size, broad head, short eyes, and rather conspicuous
(subdecumbent) pubescence. The short petiole and sublucid integument are also
distinctive.

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TAXONOMIC COMMENTS ON OTHER PSEUDOMYRMEX SPECIES

Introduction

The following notes are based on critical examination of type material in various
(mostly European) collections, supplemented by infusions of fresh material from
recent field work in Central and South America. Most of the new synonymies given
here are those that I consider relatively clear and straightforward. Much additional
synonymy is certain, but this cannot be documented reliably without detailed
species-level revisions of the groups concerned. Despite such synonymy, it should
also be pointed out that there are considerable numbers of undescribed species,
especially in the P. pallidus group.

Twenty-two species and 64 available names are formally dealt with below. The
valid species are listed alphabetically and described briefly, with related taxa
discussed or cross-referenced where appropriate. I have given particular attention to
species associated with ant-plants (. Acacia , Tachigali, Triplaris, etc.) because these
have received considerable ecological attention, often accompanied by taxonomic
inaccuracies. Grouped according to the plants which they inhabit, the following
plant-associated species of ants are discussed and diagnosed:

Acacia P. ferrugineus , P. flavicornis, P. nigrocinctus (under P. ferrugineus), P.

nigropilosus (under P . ferrugineus), P. spinicola, P. subtilissimus (above,
p. 432). See also note added in proof on P. peperi (p. 452).

Pithecellobium

P. perboscii

Tachigali P. concolor, P. malignus, P. tachigaliae

Triplaris P. dendroicus, P. triplaridis, P. triplarinus (under P. dendroicus), P.
viduus

P. viduus occurs in association with additional ant-plant genera ( Ocotea , Cordia,
Sapium, etc.), displaying a catholicity not found in the other Pseudomyrmex.

Species accounts

Pseudomyrmex concolor (F. Smith)
(Figure 40)

Pseudomyrma concolor F. Smith, 1860:70. Syntype dealate queen (unique?), “St. Paul” [Sao Paulo de
Olivenga], Amazonas, Brazil (BMNH) [Examined],

Pseudomyrma penetrator F. Smith, 1877:66. Syntype alate queen, “St. Paul” [Sao Paulo de Oliven^a],
Amazonas, Brazil (BMNH) [Examined] [Synonymy by Kempf, 1967:5; here confirmed],

Pseudomyrma latinoda Mayr, 1877:877. Holotype worker, Amazonas [probably Barreiras de Unahan, Rio
Purus; see Benson & Setz, 1985], Brazil (Trail) (NHMV) [Examined], Syn. nov.

Pseudomyrma damnosa Wheeler, 192 lb: 139. Syntype workers, queens, males, Kartabo, Guyana (W. M.
Wheeler) and Penal Settlement, Guyana (W. M. Wheeler) (MCZC) [Examined] [Synonymy, under P.
latinodus (Mayr), by Kempf, 1961:406; here confirmed].

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436

Ward

This is a widespread Tachigali-inhabiling species, usually going by the name
P. latinodus (Mayr). Comparison of the type queen of P. concolor with
worker-associated queens of P. latinodus from Brazil, Guyana, and Venezuela,
shows that the two are conspecific. Among the several species of Pseudomyrmex
associated with Tachigali, the queens and workers of P. concolor are recognized
by the following combination of features: upper third of the head predominantly
smooth and shining, with scattered punctures of variable density; median clypeal
lobe ventrally deflected and without lateral teeth or angles (Fig. 40); erect
pilosity conspicuous on most parts of the body, including the sides of the head;
light orange-brown in color.

The relationship of P. concolor to the four forms described as varieties of P.
latinodus , namely P. coronatus (Wheeler), P. endophytus (Forel), P. nigrescens
(Forel), and P. opacior (Forel), is unclear. Closely related but distinct species in
this complex include P. malignus (Wheeler) ( q.v .) and P. tachigaliae (Forel)
( q.v .), differing in pilosity and clypeal configuration (compare Figs. 40, 41, 42).

Pseudomyrmex dendroicus (Forel), stat. reval.

Pseudomyrma dendroica Forel, 1904a:40. Syntype workers, males, Rio Purus, Amazonas, Brazil (A. Goldi)
(MCSN, MHNG, MZSP) [Examined], One syntype worker in MHNG here designated LECTOTYPE.
Stat. reval. [Incorrectly synonymized with P. triplarinus (Weddell) by Kempf, 1961:408]

Pseudomyrma dendroica var. emarginata Forel, 1904b:684. Syntype workers, queen, Mavany Jurua,
Amazonas, Brazil (Ule) (MCSN, MHNG) [Examined] Syn. nov. [Incorrectly synonymized with P.
triplarinus (Weddell) by Kempf, 1961:408]

This ant is one of several Pseudomyrmex species which are obligate inhabitants
of trees of the genus Triplaris. Workers of P. dendroicus are distinguished from
those of the closely related species, P. triplarinus (Weddell) ( = P. arborissanctae
(Emery)), by the following features: (i) more widely separated frontal carinae; (ii)
dark brown head, contrasting with a lighter-colored mesosoma (head and mesosoma
more or less concolorous in P. triplarinus)’, (iii) subpetiolar process usually
subrectangular and recurved backwards to some degree (sub triangular in P.
triplarinus ); (iv) erect pilosity less abundant; fewer than 10 standing hairs in outline
on outer surface of hind tibia (erect pilosity more conspicuous and abundant in P.
triplarinus ).

See also descriptions of P. triplaridis (Forel) and P. viduus (F. Smith).

Pseudomyrmex ejectus (F. Smith)

Pseudomyrma ejecta F. Smith, 1858:157. Lectotype worker, “Brazil?” (BMNH) [Examined] [Lectotype
designation and discussion of type locality by Ward, 1985:231].

Ponera ( Ectatoma ) [sic] lincecumii Buckley, 1866:172. Syntype worker(s), “On trees in Central Texas”
[type(s) lost; not examined], Syn. nov.

In a recent review of the Nearctic species of Pseudomyrmex (Ward, 1985) I did
not consider this long-standing nomen dubium. The original description supports

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437

Smith’s (1951,1958) placement of Ponera lincecumii in the Pseudomyrmecinae.
Among the species of Pseudomyrmecinae occurring in central Texas, only
Pseudomyrmex ejectus (F. Smith) conforms to Buckley’s description with respect to
size and color (“length 0.15 inch. Upper surface of head and abdomen sub-piceous,
the rest reddish-yellow or pale yellow...”) and there is nothing in Buckley’s
description which precludes P. ejectus. As the Nearctic Pseudomyrmex species are
now relatively well-known (Ward, 1985), the synonymy seems reasonably secure.

Pseudomyrmex euryblemma (Forel)

Pseudomyrma euryblemma Forel, 1899:90. Syntype worker, (unique), Alajuela, Costa Rica (Alfaro)
(MCSN) [Examined],

Pseudomyrma kurokii Forel, 1906:226. Syntype worker (unique), Sierra Nevada de Santa Marta, Colombia
(Forel) (MHNG) [Examined], Syn. nov.

P. euryblemma is a common and widespread species, whose laterally angulate
median clypeal lobe (in workers and queens) and anteroventrally recurved pygidium
(in males), clearly place it in the P. pallidus group. Workers of this species have a
broad head (HW 0.81-0.98, Cl 0.86-0.92; n=14) whose puncticulate-coriarious
sculpture weakens towards the vertex (which is sublucid); wide but shallow
metanotal groove; long basal face of the propodeum; and pubescent fourth
abdominal tergite. The propodeal spiracles are rather salient in dorsal view, and
individuals are typically dark brown in color.

See also P. holmgreni (Wheeler) and P. rufiventris (Forel).

Pseudomyrmex ferrugineus (F. Smith)

(Figure 45)

Pseudomyrma ferruginea F. Smith, 1877:64. Syntype workers, Mexico (BMNH) [Examined]; one worker
here designated LECTOTYPE.

Pseudomyrma belti race fulvescens Emery, 1890:64. Syntype workers, Guatemala (Beccari) (MCSN)
[Examined] Syn. nov. One worker here designated LECTOTYPE.

Pseudomyrma canescens Wasmann, 1915:321. Syntype workers, Tampico, Mexico (Brakhoven) (MCSN,
MCZC) [Examined] Syn. nov. (Also preoccupied).

Pseudomyrma wasmanni Wheeler, 1921a:22. Replacement name, now unnecessary, for P. canescens
Wasmann, 1915 (nec F. Smith, 1877).

Pseudomyrma belti subsp. bequaerti Wheeler, 1942:164. Syntype workers, Puerto Castillo, Honduras (J.

Bequaert) (MCZC) [Examined] Syn. nov. One worker here designated LECTOTYPE.

Pseudomyrma belti subsp. saffordi Wheeler, 1942:162. Syntype workers, Chicoasen, Chiapas, Mexico (G.

N. Collins) (MCZC) [Examined] Syn. nov. One worker here designated LECTOTYPE.

Pseudomyrma belti subsp. vesana Wheeler, 1942:163. Holotype (unique syntype) worker, Cordoba, Mexico
(F. Knab) (MCZC) [Examined] Syn. nov.

Pseudomyrma belti subsp. bequaerti Enzmann, 1945:80. Syntype workers, Puerto Castillo, Honduras (J.

Bequaert) (MCZC) [Examined] [Objective synonym of P. belti bequaerti Wheeler; Brown, 1949:42].
Pseudomyrma kuenckeli var. hondurana Enzmann, 1945:87. Three syntype workers, Honduras (Bates)
(MCZC) [Examined]; one syntype here designated LECTOTYPE. Syn. nov. [Incorrectly synonymized
under P. kuenckeli (Emery) by Kempf, 1961:402],

P. ferrugineus (F. Smith) is an obligate inhabitant of swollen-thorn acacias,
distributed from Mexico to Honduras, whose interaction with the acacias was the

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438

Ward

subject of a detailed experimental study by Janzen (1967). The worker of P.
ferrugineus has a densely punctate head which is nevertheless weakly sublucid in
the area between the ocelli and upper margin of the compound eye; the median
clypeal lobe is anteroventrally subtended and without lateral angles or teeth (Fig.
45); and the body is variable in color, usually brown or dark brown, the mesosoma
sometimes lighter in color than the head and gaster.

The Costa Rican acacia-ant reported in the ecological literature as
“Pseudomyrmex ferruginea" (e.g., Janzen, 1983) is, in fact, P. spinicola (Emery)
{q.v.)- The two other common members of the P. ferrugineus group in Costa Rica
are P . flavicornis (F. Smith) ( q.v .) and the small (worker HW < 0.85), orange-brown
species, P. nigrocinctus (Emery). A fourth species of acacia-ant, P. nigropilosus
(Emery), is a member of the P. gracilis group and is recognizable by its large size
(worker HW > 1.16), elongate eyes which almost reach the level of the median
ocellus, laterally submarginate pronotum, and conspicuous black pilosity (further
description in Kempf, 1958). Like P. subtilissimus (see above, p. 432) P.
nigropilosus is a non-aggressive species, whose workers provide no protection to the
acacia that they occupy (Janzen, 1975).

Pseudomyrmex filiformis (Fabricius)

Formica filiformis Fabricius, 1804:405. Syntype dealate queen, Essequibo, Guyana (Smidt) (ZMUC)
[Examined]; here designated as LECTOTYPE. A second syntype dealate queen in ZMUC, lacking head,
metasoma, and a locality label, has been labelled paralectotype.

Pseudomyrma cephalica F. Smith, 1855:168. Dealate queen, labelled as type, Santarem, Brazil (Bates); two
workers, probable syntypes. Villa Nova, Brazil (Bates) (BMNH) [Examined] [Synonymy by Roger,
1862:289],

Pseudomyrma biconvexa Forel, 1899:95. Syntype workers, Pantaleon, Guatemala (Champion) (BMNH,
MHNG); Costa Rica (Tonduz) (BMNH) [Examined] [Synonymy by Wheeler, 1919:125],

Pseudomyrma biconvexa var. longiceps Forel, 1906:229. Syntype worker, Santa Marta, Colombia (A. Forel)
(MHNG) [Examined]. Syn. nov.

Pseudomyrma longiceps Stitz, 1933:69. Holotype queen, Macuto, near La Guayra, Venezuela (C. Gazgo)
(not in ZMUH, probably destroyed during World War II). Syn. nov. (Also preoccupied).

The syntype worker of P. longiceps (Forel) is simply a large P. filiformis worker,
with a rather elongate head and a conspicuous subpetiolar tooth. I have seen such
variant workers within nest-series of typical P. filiformis from elsewhere in Central
and South America. Although the unique type of P. longiceps (Stitz) is lost, the
original description (particularly as it pertains to head length, position of eyes, and
petiole shape) is closer to that of P . filiformis than any other species known to me.

Pseudomyrmex flavicornis (F. Smith )

Pseudomyrma flavicornis F. Smith, 1877:67. Three syntype workers, Nicaragua (BMNH) [Examined]. One
syntype here designated LECTOTYPE.

Pseudomyrma belti Emery, 1890:63. Syntype workers, queens, Alajuela, Costa Rica (MCSN) [Examined],
Syn. nov.

Pseudomyrma belti var. obnubila Menozzi, 1927:273. Syntype worker, San Jose, Costa Rica (H. Schmidt)

Pseudomyrmecine Ants

439

(NHMB) [Examined]. Syn. nov.

Pseudomyrma belti subsp. fellosa Wheeler, 1942:160. Syntype workers, Nicaragua (W. Fluck); Granada,
Nicaragua (C. F. Baker) (MCZC) [Examined], Syn. nov.

P. flavicornis is a widespread Central American acacia-ant, referred to in most
publications as P. belti. Because the taxonomic status of P. belti and its various
“subspecies” or “varieties” has never been fully clarified, I have little hesitation in
replacing P. belti with the less well-known senior synonym. Of the various
infraspecific names associated with P. belti (see Kempf, 1972), only two (P.
obnubilus and P. fellosus) are here considered synonyms of P. flavicornis. One of
the remaining names refers to an apparently distinct species ( P . veneficus Wheeler),
while the others are synonyms of P . ferrugineus ( q.v .). The worker of P. flavicornis
is distinguished from other obligate acacia-ants (P. ferrugineus group) by its dark
brown or black color (mesosoma sometimes contrastingly lighter); broad, densely
punctate, opaque head which lacks a sublucid area posteromesad of the compound
eye (in contrast to P. ferrugineus ); and narrow, anteroventrally subtended, median
clypeal lobe, whose lateral comers are rounded.

Pseudomyrmex gracilis (Fabricius)

Formica gracilis Fabricius, 1804:405. Syntype worker, Essequibo, Guyana (Smidt) (ZMUC) [Examined];
here designated LECTOTYPE.

Pseudomyrma variabilis F. Smith, 1877:62. Syntype worker, “Barbadoes” (BMNH) [Examined]; here
designated LECTOTYPE. Syn. nov.

The lectotype worker of P. gracilis , although lacking a head, seems to
correspond rather well to the concept of P. gracilis which has become prevalent in
publications. The mesosoma, postpetiole, and gaster are dark brown to black, the
petiole a contrasting light castaneous brown; fine, silvery (not black) pilosity covers
most of the body, and the associated piligerous punctures subdue the lustre of the
integument; the petiole is narrow and slender, with a long anterior peduncle; and the
pronotum is margined laterally but not sharply so. I do not attach much taxonomic
significance to the light-colored petiole. The P. gracilis lectotype worker agrees
well with material from Kartabo, Guyana (leg. Wheeler) in which there is variable
infuscation of the petiole. A second worker in the P. gracilis “type series” in
ZMUC, with a red “TYPE” label, but no locality or identification label, is in fact not
conspecific (it is a worker of P. maculatus (F. Smith)) and should be excluded from
consideration as type.

There is a bewildering and variable array of forms, variously described as
subspecies or “varieties” of P. gracilis, which require detailed taxonomic study. I
suspect that most of these will prove to be synonyms of a single polytypic species
(P. gracilis ), but at this stage there is insufficient information about the intra- and
inter-specific components of this variation. One unambiguous synonymy can be
established here: the lectotype worker of P. variabilis (F. Smith) in BMNH agrees
very closely with that of P. gracilis , the only substantial difference being that the P.

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440

Ward

variabilis petiole is black. I have designated a lectotype of P. variabilis because a
second worker glued to the same card (and bearing therefore the same type label as
P. variabilis ) is that of a different species - P. maculatus (F. Smith)! I have printed
a lectotype label for P. variabilis and marked the card shared by the two specimens
in such a way that the P. maculatus worker is clearly excluded as a type specimen of
P. variabilis.

Pseudomyrmex holmgreni (Wheeler)

Pseudomyrma holmgreni Wheeler, 1925:11. Two syntype workers, Chaquimayo, Peru (N. Holmgren)
(NHRS) [Examined], One worker here designated LECTOTYPE.

Contrary to statements in the original description, this widespread South
American species is not closely related to P . filiformis. Rather, it is a member of the
P. pallidus group, bearing some resemblance to P. euryblemma ( q.v .) but differing
in the following worker characters: metanotal groove wider and more deeply
incised; basal face of propodeum subequal in length to declivitous face (longer than
the declivitous face in P. euryblemma ); and head tending to be more elongate, with
the sides rounding more gradually into the occipital margin. The workers and
queens of P. holmgreni vary considerably in color, from concolorous orange to dark
brown, including intermediate forms with variable degrees of infuscation of the
body.

Pseudomyrmex laevifrons Ward, nom. nov.

Pseudomyrma laeviceps F. Smith, 1877:63. Two syntype workers, Para, Brazil (BMNH) [Examined]. One
syntype worker here designated LECTOTYPE. [Preoccupied by Pseudomyrma laeviceps F. Smith, 1859
= Tetraponera laeviceps (F. Smith)].

This is a small (worker HW 0.48-0.56, n=14) distinctive species with closely
contiguous, anterodorsally elevated frontal carinae; a smooth, shiny, puncticulate
head; long eyes; and, in the workers, an unusually shaped propodeum such that the
basal and declivitous faces meet at a sharp angle. The known range of P. laevifrons
extends from Costa Rica to Bolivia and Brazil.

Pseudomyrmex malignus (Wheeler)

(Figure 41)

Pseudomyrma maligna Wheeler, 1921b: 143. Syntype workers, males, queens, Kartabo, Guyana (W.M.
Wheeler) (MCZC, MZSP) [Examined].

Pseudomyrma maligna var. cholerica Wheeler, 1921b: 146. Syntype workers, Kartabo, Guyana (W.M.
Wheeler) (MCZC) [Examined] Syn. nov.

Pseudomyrma maligna var. crucians Wheeler, 1921b: 147. Syntype workers, Kartabo, Guyana (W.M.
Wheeler) (MCZC) [Examined] Syn. nov.

Pseudomyrma auripes Wheeler, 1922:5. Holotype queen, Trinidad, July 1920 (W.M. Wheeler) (MCZC)
[Examined] Syn. nov.

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441

Workers and queens of this Tachigali- associated ant species are distinguished
from those of P. concolor ( = P. latinodus) by the following features: median
clypeal lobe less ventrally deflected, laterally subangulate (Fig. 41); erect pilosity
sparser, lacking on sides of head above the eyes (frontal view); and anteroventral
process of petiole more prominently recurved, hook-like. In addition, the queen has
distinctive, basally geniculate mandibles ( e.g ., Wheeler, 1921b, Fig. 14). The color
and size variation reflected in the “varieties” cholericus and crucians can be seen
within single nest series.

See also the discussion under P. concolor and P. tachigaliae (Forel).

Pseudomyrmex osurus (Forel), stat. reval.

Pseudomyrma levigata [sic] subsp. osura Forel, 1911:279. Syntype worker, Costa Rica (Pittier) (MHNG)
[Examined], Stat. reval. [Incorrectly synonymized (provisionally) under P.faber (F. Smith) by Kempf,
1958:449],

Pseudomyrma laevigata subsp. insularis Enzmann, 1945:88. Holotype worker, Barro Colorado Island,
Panama (location unknown) [not examined], Syn. nov. [Incorrectly synonymized (provisionally) under
P.faber (F. Smith) by Kempf, 1958:449],

Examination of the syntype worker of P. osurus shows this to be a distinct
species, differing from P. faber (F. Smith) by the shape of the petiole (possessing a
distinct anterior peduncle and differentiated node, in lateral view), and the smooth,
shiny puncticulate head. Enzmann’s description of P. insularis fits P. osurus
tolerably well, and agrees more closely with P. osurus than with any other species in
the P. laevigatus complex known to me from Central America.

P. osurus is typically concolorous orange (head black in some Costa Rican and
Colombian workers, which I take to be this species), while P.faber is more variable,
with some workers (including the type) having a dark brown to black head and
mesosoma.

Pseudomyrmex pazosi (Santschi), stat. nov.

Pseudomyrma pazosi Santschi, 1909:309. Syntype workers, queens, Cuba (Pazosi) (NHMB) [Examined],
One syntype worker here designated LECTOTYPE.

Pseudomyrma flavidula var. pazosi Santschi; Wheeler, 1913:484.

Pseudomyrma flavidula var. jaumei Aguayo, 1932:217. Holotype worker, El Palenque, Cuba (M. Jaume)
(MCZ) [Examined] Syn. nov.

P. pazosi is a small (worker HW < 0.76) Cuban species in the P. pallidus group,
with orange head, mesosoma, petiole, and postpetiole, and a contrasting black
gaster; smooth, shiny puncticulate head; broad fore femur; and a gaster devoid of
dense pubescence. The P. jaumei holotype merely represents a faded, discolored P.
pazosi worker.

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Pseudomyrmex perboscii (Guerin)
(Figure 43)

Myrmex perboscii Guerin, 1844:428. Holotype queen, Baie de Campeche, Mexico (Perbosc) (not in MCSN,
MNHN, or ZSMC) [Not examined].

Tetraponera testacea F. Smith, 1852:45. Holotype dealate queen, [Rio] Napo, Peru (BMNH) [Examined]
Syn. nov.

Pseudomyrma perbosci [sic] Guerin; Forel, 1899:96 [Description of worker].

Pseudomyrma simoides Forel, 1911:281. Syntype worker, Amazonas, Brazil (Bates) (MHNG) [Examined]
Syn. nov.

Pseudomyrma icterica Wheeler, 1922:4. Holotype worker, Port of Spain, Trinidad (A. Busck) (MCZC)
[Examined] Syn. nov.

This medium-sized species (worker HW 1.09-1.44, n=15) has a truncate median
clypeal lobe, which is sharply rounded laterally (Fig. 43); relatively well-separated
frontal carinae (MFC subequal to distal scape width) and conspicuously protruding
median lobes of the antennal sclerites; a deeply incised metanotal groove (in the
worker); and a somewhat shiny, orange- to testaceous-brown, integument, the gaster
sometimes darker in color. The head and mesosoma have fine, puncticulate
sculpture, the punctulae varying in density, as do the minute, piligerous punctures
(and associated appressed pubescence) on the postpetiole and gaster. Standing
pilosity is fine and rather sparse (lacking on outer faces of the tibiae and on the
worker mesonotum).

Having examined the types of P. testaceus, P. simoides, and P. ictericus, I feel
fairly confident about the specific identity of these three. Moreover, they appear to
be conspecific with the “ Pseudomyrma perbosci ” worker described by Forel (1899)
from Costa Rica. Unfortunately, the type of P. perboscii could not be located, but
the original description, while scanty, contains enough information to justify the
above synonymy. Among the features mentioned by Guerin, the combination of the
elongate, subrectangular head (nearly twice as long as wide), globose postpetiole,
fawn-yellow body with dark gaster, finely shagreened to shiny appearance, and
large size (9.5mm long), fits the queen of no other species known to me.

In northern Colombia and Venezuela I collected colonies of this species in live
terminal branches of saman ( Pithecellobium saman) trees. The ants occupied
numerous unconnected cavities, 5-20 mm long (2-^1 mm internal diameter) in which
they kept brood and tended coccids. The cavities appeared to be intrinsic to the plant
since unoccupied cavities, without entrance holes, could be found. Although the
workers patrolled the foliage and would sting if molested, they were much less
aggressive than the Pseudomyrmex ants inhabiting Tachigali, Triplaris, or
swollen-thorn acacias. Thus the association may represent an early (or arrested?)
phase in the development of an ant-plant mutualism.

Pseudomyrmex rufiventris (Forel) stat. nov.

Pseudomyrma kurokii var. rufiventris Forel, 1911:275. Syntype queen, San Bernardino, Paraguay (K.
Fiebrig) (ZSMC) [Examined].

Pseudomyrmecine Ants

443

Although obviously closely related to P. euryblemma (= P. kurokii) ( q.v .), P.
rufiventris appears to be distinct enough to warrant specific status. The head of the
queen and worker is covered with dense, punctulate-coriarious sculpture which
renders it opaque; this may be contrasted with a weaker puncticulate sculpture and
sublucid appearance in P. euryblemma.

Pseudomyrmex rufomedius (F. Smith)

Pseudomyrma rufomedia F. Smith, 1877:66. Syntype alate queen (unique), Aceituno, Guatemala (BMNH)

[Examined],

Pseudomyrma stolli Forel, 1912:20. Syntype worker, queens, male, Retalhuleu, Guatemala (Stoll)(MHNG)

[Examined] Syn. nov.

P. rufomedius is a distinctive species, known only from Guatemala and southern
Mexico, workers and queens of which possess a broad head with relatively small
eyes; a flattened and laterally marginate pronotum; a more or less apendunculate
petiole, much longer than high or wide, whose dorsal face is also flattened and
laterally margined; very fine, pale, inconspicuous pilosity; and a bicolored body
(orange petiole contrasting with dark brown head and gaster; remainder of body
variably infuscated). An additional characteristic feature of the worker is the virtual
obliteration of the metanotal groove, the mesonotum and propodeum forming a
continuous, elevated, and laterally compressed plate.

The P. rufomedius type, although lacking a head, agrees closely with the
worker-associated queens of P. stolli (and other material which I have examined),
and there can be no doubt about the identity of the species (c/. Perrault, 1987).

Pseudomyrmex simplex (F. Smith)

In reviewing this species, as one of these occurring in the Nearctic region, I
interpreted incorrectly the type locality “St. Paul” to be in the state of Sao Paulo,
Brazil (Ward, 1985:238). In fact, “St. Paul” refers to Sao Paulo de Oliven^a, a Bates
collecting locality in the Brazilian state of Amazonas (Kempf, 1972).

Pseudomyrmex spinicola (Emery)

(Figure 44)

Pseudomyrma spinicola Emery, 1890:64. Syntype workers, queens, males, Alajuela, Costa Rica (Alfaro)

(MCSN) [Examined], One worker here designated LECTOTYPE.

P. spinicola is a common species of acacia-ant, ranging from Honduras to
Colombia, workers of which are distinguished from those of the more northerly
species, P. ferrugineus, by the following traits: median clypeal lobe with a concave
anterior margin and with lateral angles or teeth (Fig. 44); frontal carinae more
closely contiguous and median lobes of antennal sclerites correspondingly more
exposed; frons with more extensive shiny interspaces between the punctures;

Quaest. Ent., 1989, 25 (4)

444

Ward

propodeal spiracles more salient in dorsal view; petiole longer, with a tendency
towards more pronounced posterolateral angles in dorsal view.

Most of the forms described as subspecies or varieties of P. spinicola are
probably conspecific with it (with the exception of P. convarians (Forel)), but I
refrain from proposing formal synonymy until the variation is better understood.

Pseudomyrmex tachigaliae (Forel) stat. nov.

(Figure 42)

Pseudomyrma latinoda race tachigaliae Forel, 1904b:686. Syntype workers, queens, males, Tarapoto, Peru
(Ule) (MHNG) [Examined].

Pseudomyrma latinoda subsp. bradleyi Wheeler, 1942:169. Syntype workers, Perene, Peru (Bradley)
(MCZC) [Examined] Syn. nov.

Pseudomyrma bradleyi Enzmann, 1945:82. Syntype workers, Perene, Peru (Bradley) (MCZC) [Examined]
[Objective synonym of Pseudomyrma latinoda bradleyi Wheeler; Brown, 1949:42]

This Tachigali- ant is apparently confined to the western Amazon basin and
adjacent foothills. The worker is recognized by the laterally angulate and inflected
median clypeal lobe (Fig. 42); subcontiguous frontal carinae (MFC < basal scape
width); sparse and inconspicuous pilosity; and dense appressed pubescence which is
much better developed than in P. malignus or P. concolor.

Pseudomyrmex tenuis (Fabricius)

Formica tenuis Fabricius, 1804:405. Syntype worker, Essequibo, Guyana (Smidt) (ZMUC) [Examined];
here designated LECTOTYPE.

The lectotype worker of P. tenuis is concolorous testaceous brown, with a
densely punctate, opaque head; large elongate eyes; inflected, tectiform median
clypeal lobe; sharply margined pronotum; tall, thin, anterodorsally angulate
(pointed) petiole; and no erect pilosity on the mesosoma dorsum. This agrees with
the concept of P. tenuis which has been prevalent in the literature.

For a listing of synonyms see Kempf (1972), but note that P. nigriceps (F.
Smith) is now a junior synonym of P. termitarius (F. Smith) (Perrault, 1987).

Pseudomyrmex triplaridis (Forel)

Pseudomyrma triplaridis Forel, 1904b:684. Syntype workers, queens, male, Jurua Miry, Jurua, Amazonas,
Brazil (Ule) (MHNG) [Examined].

Pseudomyrma triplaridis subsp. boxi Wheeler, 1942:184. Syntype workers, queens, males, Blairmont,
Berbice, Guyana (H.E. Box) (MCZC) [Examined] Syn. nov. One syntype worker here designated
LECTOTYPE.

The worker of this Triplaris-associated ant has a sublucid, densely punctate head
which is distinctly broader than that of P. viduus (Cl 0.87-0.93, n=17); frontal
carinae more closely contiguous than in P. triplarinus or P. dendroicus but less so
than in P. viduus (FCI 0.071-0.109); a median clypeal lobe which is sharply
rounded laterally; and a conspicuous, recurved (hook-like) subpetiolar process.

Pseudomyrmecine Ants

445

I have designated a lectotype of P. boxi because the type series in the MCZC
contains a few workers of P. viduus ( q.v .). Wheeler (1942) confused these two
species; the main series of P. viduus from Blairmont, Guyana was described by him
as a second subspecies of P. triplaridis (P. triplaridis tigrinus).

Pseudomyrmex viduus (F. Smith)

Pseudomyrma vidua F. Smith, 1858:158. Syntype male (unique), Ega, Amazonas, Brazil (BMNH)
[Examined],

Pseudomyrma caroli Forel, 1899:89. Syntype workers, queens, Nicoya, Costa Rica (M. R. Alfaro) (MCSN,
MHNG) [Examined]. Syn. nov.

Pseudomyrma caroli var. sapii Forel, 1904b:688. Syntype workers, queens, Bom Fim, Jurua, Amazonas,
Brazil (Ule) (MHNG) [Examined], Syn. nov.

Pseudomyrma ulei Forel, 1904b:689. Syntype workers, Jurua Miry, Jurua, Amazonas, Brazil (Ule) (MHNG)
[Examined]. Syn. nov.

Pseudomyrma triplaridis subsp. baileyi Wheeler, 1942:185. Syntype workers, queens, Camaria, Cuyuni
River, Guyana (W. M. Wheeler) (MCZC) [Examined] Syn. nov.

Pseudomyrma triplaridis subsp. tigrina Wheeler, 1942:186. Syntype workers, males, Blairmont, Berbice,
Guyana (H. E. Box) (MCZC) [Examined] Syn. nov. One syntype worker here designated LECTOTYPE.
Pseudomyrma triplaridis subsp. biolleyi Enzmann, 1945:93. Syntype workers, queens, Camaria, Cuyuni
River, Guyana (W. M. Wheeler) (MCZC) [Examined] [Objective synonym of P. triplaridis baileyi
Wheeler; Brown, 1949:43].

Pseudomyrma triplaridis subsp. trigona Enzmann, 1945:94. Syntype workers, males, Blairmont, Berbice,
Guyana (N. E. Box) (MCZC) [Examined] [Objective synonym of P. triplaridis tigrina Wheeler; Brown,
1949:43],

The type of P. viduus is a damaged male lacking head, postpetiole, and gaster.
Nevertheless the shape of the petiole (in particular, the broad anterior peduncle;
MPW = 0.22, PWI3 = 0.61) and general habitus clearly place it in the cluster of
species which Kempf (1961) called the P. latinodus group (cf. Perrault, 1987). I
compared it with males of likely candidate species, namely P. concolor (= P.
latinodus ), P. nigrescens, P. tachigaliae, P. malignus, P. caroli, P. triplaridis, P.
symbioticus, P. triplarinus, and P. dendroicus. The P. viduus type agrees closely
with males of P. caroli with respect to size, pilosity, mesonotal sculpture, and
(especially) shape of the petiole, while disagreeing with the remaining species in
petiole shape and in one or more aspects of size, pilosity, or sculpture. Specifically,
P. viduus has a shiny punctate mesonotum, covered with both appressed pubescence
and fine, golden pilosity, the latter up to 0.20 mm long; conspicuous pilosity on the
legs and petiole (ca. 19 standing hairs in outline on the petiolar dorsum); an elongate
petiole (PLI = 0.52); and small absolute size (PL = 0.77, PH = 0.40, DPW = 0.36,
mesonotal width = 0.92). Only males of P. caroli agree with this combination of
characters. The other names listed under the synonymy of P. viduus ( P . sapii, P. ulei
P. baileyi, P. tigrinus , and Enzmann’s copycat names) appear to represent nothing
more than the normal variation of the species heretofore known as P. caroli.
Workers and queens of this species can be recognized by their elongate heads
(worker Cl 0.70 - 0.80, n=22; queen Cl 0.62 - 0.68, n=8) and closely adjacent
frontal carinae (worker FCI 0.048 - 0.087; queen FCI 0.062 - 0.085) compared to

Quaest. Ent., 1989, 25 (4)

446

Ward

other members of the erstwhile P. latinodus group (here referred to as the P. viduus
group).

As I presently understand it, P. viduus is a widespread, somewhat variable
species, and a generalist occupant of ant-plants. Colonies have been collected within
live branches or cavities of Ocotea, Cordia, Sapium, Coussapoa, and Triplaris.
Detailed biological studies of P. viduus may reveal several cryptic species,
associated with particular ant-plants, but the present material does not support any
clear division(s).

Summary of Taxonomic Changes

P. concolor (F. Smith), 1860.

= P. latinodus (Mayr, 1877), syn. nov.

P. dendroicus (Forel, 1904a), stat. reval.

= P. emarginatus (Forel, 1904b), syn. nov.

P. ejectus (F. Smith, 1858).

= P. lincecumii (Buckley, 1866), syn. nov.

P. euryblemma (Forel, 1899).

= P. kurokii (Forel, 1906), syn. nov.

P . ferrugineus (F. Smith, 1877).

= P.fulvescens (Emery, 1890), syn. nov.

= P. canescens (Wasmann, 1915), syn. nov. (preoccupied)

= P. wasmanni (Wheeler, 1921a) (replacement name for P. canescens
Wasmann).

= P. bequaerti (Wheeler, 1942), syn. nov.

= P. saffordi (Wheeler, 1942), syn. nov.

= P. vesanus (Wheeler, 1942), syn. nov.

= P. honduranus (Enzmann, 1945), syn. nov.

P.filiformis (Fabricius, 1804).

= P. longiceps (Forel, 1906), syn. nov.

= P. longiceps (Stitz, 1933), syn. nov. (preoccupied).

P . flavicornis (F. Smith, 1877).

= P. belti (Emery, 1890), syn. nov.

= P. obnubilus (Menozzi, 1927), syn. nov.

= P.fellosus (Wheeler, 1942), syn. nov.

P. gracilis (Fabricius, 1804).

= P. variabilis (F. Smith, 1877), syn. nov.

P. holmgreni (Wheeler). No taxonomic changes proposed.

P. laevifrons Ward, nom. nov.

= P. laeviceps (F. Smith, 1877) (preoccupied).

P. malignus (Wheeler, 1921b).

= P. cholericus (Wheeler, 1921b), syn. nov.

Pseudomyrmecine Ants

447

= P. crucians ( Wheeler , 1921b), syn. nov.

= P. auripes (Wheeler, 1922), syn. nov.

P. osurus (Forel, 1911), stat. reval.

= P. insularis (Enzmann, 1945), syn. nov.

P.pazosi (Santschi, 1909), stat. nov.

= P.jaumei (Aguayo, 1932), syn. nov.

P. perboscii (Guerin, 1844).

= P. testaceus (F. Smith, 1852), syn. nov.

= P. simoides (Forel, 1911), syn. nov.

= P. ictericus (Wheeler, 1922), syn. nov.

P. rufiventris (Forel, 1911), stat. nov.

P. rufomedius (F. Smith, 1877).

- P. stolli (Forel, 1912), syn. nov.

P. simplex (F. Smith). No taxonomic changes proposed.

P. spinicola (Emery). No taxonomic changes proposed.

P. tachigaliae (Forel, 1904b), stat. nov.

= P. bradleyi (Wheeler, 1942), syn. nov.

P. tenuis (Fabricius). No taxonomic changes proposed.

P. triplaridis (Forel, 1904b).

= P. boxi (Wheeler, 1942), syn. nov.

P. viduus (F. Smith, 1858).

= P. caroli (Forel, 1899), syn. nov.

= P. sapii (Forel, 1904b), syn. nov.

= P. ulei (Forel, 1904b), syn. nov.

= P. baileyi (Wheeler, 1942), syn. nov.

= P. tigrinus (Wheeler, 1942), syn. nov.

CONCLUDING REMARKS

The taxonomic findings presented in this paper illustrate two recurring patterns
in the genus Pseudomyrmex : the occurrence of closely related (sibling) species with
broadly overlapping geographical ranges and, at the same time (if my interpretation
of synonymy is correct), the existence of considerable geographical variation within
many species.

There are relatively few allopatric pairs of sister species in the genus (P.
curacaensis and P. cretus are one possible example). Pseudomyrmex queens of all
species are fully winged and capable of dispersal; such vagility may be responsible
for the relative rapidity with which recently speciated forms come into secondary
contact (assuming that initial differentiation occurred in allopatry). Moreover,
Pseudomyrmex species which are generalist nesters in dead twigs - and this is true
of the majority of species - are remarkably tolerant of other congeners in the same
habitat, with the result that allospecific colonies often share the same foraging space.

Quaest. Ent., 1989, 25 (4)

448

Ward

For example, thirteen species of Pseudomyrmex were observed foraging on the fig
tree and adjacent vegetation, where the type specimens of Pseudomyrmex spiculus
were collected.

Such close habitation is not typical of those Pseudomyrmex species which live in
plant domatia. Colonies of these species seldom overlap in foraging space; yet there
may be several species in a given locality. In the Tarapoto region of northern Peru,
Triplaris plants are occupied by four related species in the P. viduus group; similar
numbers of species co-occur on swollen thorn Acacia in parts of Central America.

The discrimination and identification of Pseudomyrmex species, which is
generally feasible for sympatric collections, is made more difficult on a large
geographical scale by the considerable variation which occurs within many taxa.
This is hardly a novel situation - examples of polytypy are common in ants (Ward,
1989) - but the situation is aggravated in some Pseudomyrmex where the
discriminatory value of characters varies from one region to another. Thus in Costa
Rica, workers of P. elongatus and P. caeciliae are consistently separable on head
sculpture, but in Guatemala and southern Mexico one encounters some individuals
exhibiting intermediate conditions of this character. In western Mexico, where there
appears to be a single taxon, which I take to be P. elongatus, individuals also tend to
be shifted towards an intermediate condition. Similar situations occur in other
species complexes. Whether the taxa concerned are truly discrete (but their identities
obscured by character shifts in allopatry) or are in fact reproductively isolated in
some regions but not others, cannot be determined without further population-level
studies.

ACKNOWLEDGMENTS

I am grateful to the following persons for loans of material or access to museum
collections: B. Bolton (BMNH); M. A. Tenorio (CASC); M. Wasbauer (CDAE); I.
Zenner-Polania (CELM); J. Denis (CNCC); J. Liebherr (CUIC); R. Ayala (EBCC);
D. Quintero Arias (GBFM); E. Mellini (IEGG); J. A. Rafael and A. Y. Harada
(INPA); J. Lattke (IZAV); R. Snelling (LACM; also very old loans to D. H. Janzen
from CASC, CISC, CUIC, GCWC, ICCM, INHS, KSUC, MCZC, SEMC, TAMU,
UCDC, UCRC, USNM, and UWEM, acquired by LACM along with Janzen’s
collection; these loans have now been returned to their respective institutions); R.
Poggi (MCSN); W. L. Brown, M. Moffett, and C. Vogt (MCZC); C. Besuchet
(MHNG); J. C. Weulersse (MNHN); C. R. F. Brandao (MZSP); C. Baroni Urbani
and M. Brancucci (NHMB); M. Fisher (NHMV); B. Gustafsson (NHRS); D. C.
Darling (OSUO); F. Fernandez (UNCB); D. Smith (USNM); F. Koch (ZMHB); O.
Lumholdt (ZMUC); D. R. Abraham (ZMUH); and E. Diller (ZSMC). Additional
useful material was received from John Brandbyge, Diane Davidson, Alain Dejean,
Jacques Delabie, Xavier Espadaler, Fernando Fernandez, Doug Gill, Henry
Hespenheide, Klaus Jaffe, Charles Kugler, Jack Longino, Bill MacKay, G. H.

Pseudomyrmecine Ants

449

Perrault, Fred Rickson, Cathy Toft, Manfred Verhaagh, and Woody Benson. I thank
Steve Shattuck for assistance with metric measurements and the preparation of
illustrations. Roberto Brandao helped clarify the geographical location of several
Brazilian collection sites. Useful discussions or comments on the manuscript were
provided by George Ball, Dan Janzen, Jack Longino, Steve Shattuck, and two
anonymous reviewers. This work was supported by NSF BSR-8507865.

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Ward, P. S. 1985. The Nearctic species of the genus Pseudomyrmex (Hymenoptera:

Quaest. Ent., 1989, 25 (4)

452

Ward

Formicidae). Quaestiones Entomologicae 21:209-246.

Ward, P. S. 1989. Genetic and social changes associated with ant speciation. In: M.
D. Breed and R. E. Page (Editors). The genetics of social evolution. Boulder,
Colorado: Westview Press, pp. 123-148.

Ward, P. S. 1990a. Phylogenetic analysis of pseudomyrmecine ants associated with
domatia-bearing plants. In: D. F. Cutler and C. R. Huxley (Editors). Interactions
between ants and plants. Oxford: Oxford University Press, in press.

Ward, P. S. 1990b. The ant subfamily Pseudomyrmecinae (Hymenoptera:
Formicidae): generic revision and relationship to other formicids. In prep.
Wasmann, E. 1915. Eine neue Pseudomyrma aus der Ochsenhomdomakazie in
Mexiko, mit Bemerkungen tiber Ameisen in Akaziendomen und ihre Gaste. Ein
kritischer Beitrag zur Pflanzen-Myrmekophilie. Tijdschrift voor Entomologie
58:296-325.

Wheeler, G. C. and J. Wheeler. 1973. Supplementary studies on ant larvae:
Cerapachyinae, Pseudomyrmecinae and Myrmicinae. Psyche 80:204-21 1.
Wheeler, W. M. 1913. The ants of Cuba. Bulletin of the Museum of Comparative
Zoology 54:477-505.

Wheeler, W. M. 1919. A singular neotropical ant {Pseudomyrma filiformis
Fabricius). Psyche 26:124-131.

Wheeler, W. M. 1921a. A new case of parabiosis and the “ant gardens” of British
Guiana. Ecology 2:89-103.

Wheeler, W. M. 1921b. The Tachigalia ants. Zoologica, New York 3:137-168.
Wheeler, W. M. 1922. The ants of Trinidad. American Museum Novitates 45:1-16.
Wheeler, W. M. 1925. Neotropical ants in the collections of the Royal Museum of
Stockholm. Arkiv forZoologi 17A(8):l-55.

Wheeler, W. M. 1942. Studies of neotropical ant-plants and their ants. Bulletin of
the Museum of Comparative Zoology 90:1-262.

Wilson, E. O. 1955. A monographic revision of the ant genus Lasitts. Bulletin of the
Museum of Comparative Zoology 113:1-201.

NOTE ADDED IN PROOF

Further study of Central American acacia-ants {P. ferrugineus group) permits
recognition of the following species:

Pseudomyrmex peperi (Forel).

Pseudomyrma peperi Forel, 1913:213. Syntype workers, Patulul, Guatemala (Peper) (MHNG) [Examined],
Pseudomyrma spinicola race convarians Forel, 1913:214. Syntype worker, Patulul, Guatemala (Peper)
(MHNG) [Examined] Syn. nov.

Pseudomyrma sabanica [sic] var. sajfirdi Enzmann, 1945:89. Syntype workers, Yerba Santa, Chiapas,
Mexco (G.N. Collins) (MCZC) [Examined] Syn. nov. [Incorrectly synonymized under P. belti saffordi
(Wheeler) by Brown, 1949:42],

An obligate acacia-ant, distributed from southern Mexico to Nicaragua, P. peperi
can be recognized by the following features in workers and queens: relatively small
size (worker HW < 0.92, queen HW < 0.86); light to medium brown (not black)

Pseudomyrmecine Ants

453

coloration; densely punctulate, opaque head (presenting a matte appearance); and
broad petiolar node with salient posterolateral angles.

Quaest. Ent., 1989, 25 (4)

454

Ward

Figures 1-5. Illustrations of some measurements and morphological terms used for pseudomyrmecine ants
(see also Figures 1^1 in Ward, 1985). Figs. 1, 2. Pseudomyrmex worker, right mandible: terminology and
associated measurements. Figs. 3, 4. Pseudomyrmex worker, fronto-clypeal complex: terminology and
associated measurements; right antenna (Fig. 3) and both antennae (Fig. 4) removed. Fig. 5. Pseudomyrmex
worker, lateral view of left hind tibia, illustrating the measurement LHT.

Pseudomyrmecine Ants

455

Figures 6-13. Various views of workers in the Pseudomyrmex oculatus group. Fig. 6. P. elongatus, lateral
view of worker head, antennae removed, showing the plane of view utilized for a full-face, frontal (dorsal)
view of head. Fig. 7. Lateral view of worker petiole, P. oculatus (Brazil). Fig. 8. Same, P. schuppi (Brazil).
Figs. 9-13. Paired illustrations of head (frontal view) without antennae or pilosity and petiole (lateral view)
of: P. cretus, holotype worker, Costa Rica (9a, 9b); P. curacaensis, worker, Colombia (10a, 10b); P.
pisinnus, holotype worker, Brazil (11a, lib); P. urbanus, worker, Brazil (12a, 12b); and P. alustratus,
holotype worker, Peru (13a, 13b). All drawn to same scale; scale line = 0.5 mm.

Quaest. Ent., 1989, 25 (4)

456

Ward

Figures 14-19. SEM views of the worker clypeus, in the Pseudomyrmex oculatus and P. subtilissimus
groups. The “dorsal” views (figures 14, 16, 18) are taken perpendicular to the clypeus, and are thus
“anterodorsal” relative to a full-face, dorsal view of the head (see figure 6). Fig. 14. P. elongatus (Mexico),
dorsal view. Fig. 15. P. elongatus (Colombia), oblique anterior view. Fig. 16. P. eduardi (Venezuela), dorsal
view. Fig. 17. P. eduardi (Venezuela), lateral view. Fig. 18. P. tenuissimus (Venezuela), dorsal view. Fig.
19. P. tenuissimus (Venezuela), lateral view.

Pseudomyrmecine Ants

457

Figures 20-25. SEM views of worker head sculpture in the Pseudomyrmex oculatus group. Figs. 20-23.
Supraocular sculpture: P. elongatus, Costa Rica (20); P. elongatus , Bolivia (21); P. cubaensis , Costa Rica
(22); P. caeciliae, Guatemala (23). Fig. 24. Sculpture on left half of frons, P. caeciliae, Costa Rica. Fig. 25.
Same, P. eduardi, Venezuela.

Quaest. Ent., 1989, 25 (4)

458

Ward

1.25
1.20
1.15
1.10
1.05

I

1.00
0.95
0.90
0.85
0.80

0.74 0.78 0.82 0.86 0.90 0.94 0.98

HW

Figure 26. Scattergram of HL by HW, for workers of Pseudomyrmex curacaensis and P. cretus.

0.66
0.62
0.58

_i
LlI

0.54
0.50

0.74 0.78 0.82 0.86 0.90 0.94 0.98

HW

• •

% •

o °

O o %

! •

• •

o

O o

• curacaensis
o cretus

•

• • •

o

o

<tt>

•

•

• .

• •

•

• •

•

o

o

o ° 0

o

• o

•

•

•

• curacaensis

•

o cretus

Figure 27. Scattergram of EL by HW, for workers of P. curacaensis and P. cretus.

FW

Pseudomyrmecine Ants

459

LU

0.56

0.52 -

0.48

0.44 -

0.40

>c P

cP o

O o

O &>°

o u

o o

o o

Qo

• ~ U u

J* & cPc

•

I _% • •

1 *1

o 9)

O O o
o

o o

■ coeci/iae
• urbanus
o cubaensis

HW

Figure 28. Scattergram of EL by HW, for workers of P. caeciliae, P. urbanus, and P. cubaensis.

0.33
0.31
0.29
0.27
0.25
0.23
0.31
0.19

0.50 0.54 0.58 0.62 0.66 0.70 0.74 0.78

HW

Figure 29. Scattergram of FW by HW, for workers of P. caeciliae, P. urbanus and P. cubaensis.

O ° O

oQd O O

°^8°

I.#

/ o8 &>

o

A*

\ ■ • * 8 O

i# m ° o

o

o

•

•

■■

■ caeciliae

•

• urbanus

o cubaensis

Quae st. Ent., 1989, 25 (4)

PLI

460

Ward

O.IO

0.09-

0.07 -•

0.06

• caeciliae
o urbanus

• •

•

o

o

o

oo o

• •

•. • • •

•

o

o o

_ m •

• °

o

• • • •

»

•

o°

o

o °
o

o

° 0

i ' i ■ i ■ i • i ' i ' i 1

0.52 0.54 0.56 0.58 0.60 0.62 0.64 0

HW

66

Figure 30. Scattergram of OD by HW, for workers of P. caeciiae and P. urbanus. The two largest P.
caeciliae workers come from Jamaica and are rather atypical.

0.76-

A

• spiculus

0.72-

A

a subtilissimus

AA a

A

m tenuissimus

A A

A A t

0.68-

A A

P' — ip

•

■

A A

0.64-

■■

•

m

. #«

• %

0.60-

%B .

« 9 •

%

■

■

i

• •

■

■

9 •

0.56-

■

V

■

•

0.52-

%

' 1 ' 1 ' r

' 1

0.48 0.52 0.56 0.60 0.64 0.66

HW

Figure 31. Scattergram of PLI (= PH/PL) by HW, for workers of P. spiculus, P. subtilissimus, and P.

tenuissimus.

Pseudomyrmecine Ants

461

Quaest. Ent., 1989, 25 (4)

Figures 32-39. Paired illustrations of the head (frontal view) without antennae and petiole (lateral view) of workers in the Pseudomyrmex subtilissimus group: P. spiculus, holotype
worker, Costa Rica (32, 33); P. subtilissimus , worker, Costa Rica (34, 35); P. tenuissimus, worker, Venezuela (36, 37); P. villosus, holotype worker, Brazil (38, 39). All drawings to same
scale; scale line = 0.5 mm.

■■I

462

Ward

Figures 40-45. SEM views of the worker clypeus of various Pseudomyrmex species. These “dorsal” views
are taken perpendicular to the clypeus, and are thus “anterodorsal” in relation to a full-face, dorsal view of
the head (see figure 6). 40. P. concolor, Brazil. 41. P. malignus, Venezuela. 42. P. tachigaliae, Peru. 43. P
perboscii, Venezuela. 44. P. spinicola , Costa Rica. 45. P.ferrugineus, Mexico.

Pseudomyrmecine Ants

463

INDEX TO NAMES OF TAX A
(Junior synonyms in italics)

FAMILY GROUP TAX A
Pseudomyrmecinae, 395, 437

GENERA AND SUBGENERA
Acacia, 435, 448
Anacardium, 411
Baccharis, 420
Cecropia, 420
Coccoloba, 420
Cordia, 435, 446
Coussapoa, 446
Croton, 413
Eupatorium, 425^-26
Ficus, 420, 423
Hampea, 423
Helicteres, 420
Inga, 420, 423
Ochroma, 410
Ocotea, 435, 446
Pithecellobium, 435
Prosopis, 41 1
Quercus, 411
Sapium, 420, 435, 446
Scheelea, 423
Schoepfia, 412-413
Tachigali, 435^136, 441-442
Tetraponera, 395, 398
Triplaris, 417, 435^136, 442, 444,
446, 448
Vochysia, 423

SPECIES AND SUBSPECIES
acaciarum (Wheeler),
Pseudomyrmex, 404
acanthobius (Emery),
Pseudomyrmex, 404
adustus (Borgmeier),

Pseudomyrmex, 404
advena (F. Smith), Pseudomyrmex ,
406, 422^123

alfari (Forel), Pseudomyrmex, 404
allidorus (Enzmann),

Pseudomyrmex, 406, 419
alliodora , Cordia, 420
alliodorae (Wheeler),
Pseudomyrmex, 406
altemans (Santschi),
Pseudomyrmex, 404
altinodus (Mann), Pseudomyrmex,
406, 422

alustratus sp. nov., Pseudomyrmex,
404, 406, 408-410
alvarengai Kempf,

Pseudomyrmex, 404
antiguanus (Enzmann),
Pseudomyrmex, 405
apache Creighton, Pseudomyrmex,
404

arborissanctae (Emery),
Pseudomyrmex, 436
argentinus (Santschi),
Pseudomyrmex, 404
atrinodus (Santschi),
Pseudomyrmex, 404
atripes (F. Smith), Pseudomyrmex,
404

atrox (Forel), Pseudomyrmex, 404
auripes (Wheeler),

Pseudomyrmex, 440, 447
bailey i (Wheeler), Pseudomyrmex,
445, 447

beccarii (Menozzi),
Pseudomyrmex, 404
belgranoi (Santschi),
Pseudomyrmex, 404
belti (Emery), Pseudomyrmex,

Quaest. Ent., 1989, 25 (4)

464

Ward

439, 446

bequaerti (Enzmann),
Pseudomyrmex, 437
bequaerti (Wheeler),

Pseudomyrmex , 437, 446
bicinctus (Santschi),
Pseudomyrmex, 404
bicolor (Guerin), Pseudomyrmex,

404

biconvexus (Forel),

Pseudomyrmex , 438
biolleyi (Enzmann),
Pseudomyrmex, 445
boopis (Roger), Pseudomyrmex,

405

boxi (Wheeler), Pseudomyrmex,
444^145, 447
bradleyi (Enzmann),
Pseudomyrmex, 444
bradleyi (Wheeler),

Pseudomyrmex, 444, 447
browni Kempf, Pseudomyrmex,
404

brunneus (F. Smith),
Pseudomyrmex, 404
brunnipes (Enzmann),
Pseudomyrmex, 405
caeciliae (Forel), Pseudomyrmex,
404-406, 409-411, 418-120,
428, 448

canescens (F. Smith),
Pseudomyrmex, 404
canescens (Wasmann),

Pseudomyrmex, 437, 446
caroli (Forel), Pseudomyrmex, 447
castus (Wheeler), Pseudomyrmex,
404

catappa , Terminalia, 420
cephalicus (F. Smith),
Pseudomyrmex, 438
championi (Forel),

Pseudomyrmex, 404

chodati (Forel), Pseudomyrmex,
406, 427-128
cholericus (Wheeler),

Pseudomyrmex, 440, 446
cladoicus (F. Smith),
Pseudomyrmex, 404
cocae (Santschi), Pseudomyrmex,
404

colei (Enzmann), Pseudomyrmex,
404

concolor (F. Smith),

Pseudomyrmex, 405, 435—136,
441, 444^146

confusior (Forel), Pseudomyrmex,
406, 426

convarians (Forel),

Pseudomyrmex, 444, 452
cordiae (Forel), Pseudomyrmex,
404

cordobensis (Forel),
Pseudomyrmex, 405
coronatus (Wheeler),

Pseudomyrmex, 405, 436
costaricensis (Enzmann),
Pseudomyrmex, 405
cretus sp. nov., Pseudomyrmex,
404, 406, 408, 4 1 2 — 4 1 3 ,
416-417,447
crucians (Wheeler),

Pseudomyrmex, 440, 447
cubaensis (Forel), Pseudomyrmex,
404, 406, 409—111, 413^115,
420, 428

culantrillo , Zanthoxylum, 420
culmicola (Forel), Pseudomyrmex,
429, 433

cumingiana , Triplaris, 420
curacaensis (Forel),

Pseudomyrmex, 404, 406, 408,
413—114, 416-417, 423, 447
damnosus (Wheeler),
Pseudomyrmex, 435

Pseudomyrmecine Ants

465

dendroicus (Forel),

Pseudomyrmex, 405, 435 — 436,
444-446

denticollis (Emery),
Pseudomyrmex, 405
depressus (Forel), Pseudomyrmex,
405

dimidiatus (Roger),
Pseudomyrmex, 404
distinctus (F. Smith),
Pseudomyrmex, 404
dolichopsis (Forel),

Pseudomyrmex, 406, 422
duckei (Forel), Pseudomyrmex,

405

eduardi (Forel), Pseudomyrmex,
404^106, 409,418-419
ejectus (F. Smith), Pseudomyrmex,
404, 436-437, 446
elongatulus (Dalle Torre),
Pseudomyrmex, 404, 420
elongatus (F. Smith),
Pseudomyrmex, 420
elongatus (Mayr), Pseudomyrmex,
404, 406, 408, 410—41 1,
414-415,419, 428,448
emarginatus (Forel),

Pseudomyrmex, 436, 446
endophytus (Forel),

Pseudomyrmex, 405, 436
erectus , Conocarpus, 419
ethicus (Forel), Pseudomyrmex,
404

euryblemma (Forel),

Pseudomyrmex, 404, 437, 440,
443, 446

excisus (Mayr), Pseudomyrmex,
404

faber (F. Smith), Pseudomyrmex,
404, 441

fellosus (Wheeler),

Pseudomyrmex, 439, 446

ferrugineus (F. Smith),

Pseudomyrmex, 403^104, 435,
437, 443, 446
fervidus (F. Smith),
Pseudomyrmex, 405
fiebrigi (Forel), Pseudomyrmex,
404

filiformis (Fabricius),

Pseudomyrmex, 405, 423, 438,
440, 446

flavicomis (F. Smith),

Pseudomyrmex, 404, 432, 435,
438—139, 446
flavidulus (F. Smith),
Pseudomyrmex, 404
fortis (Forel), Pseudomyrmex, 404
fulvescens (Emery),

Pseudomyrmex, 437, 446
fuscatus (Emery), Pseudomyrmex,
404

gaigei (Forel), Pseudomyrmex,

404

gebelli (Forel), Pseudomyrmex,

405

geraensis (Forel), Pseudomyrmex,
406, 426

germinans , Avicennia, 419-420
gibbinotus (Forel),

Pseudomyrmex, 404
glabriventris (Santschi),
Pseudomyrmex, 404
godmani (Forel), Pseudomyrmex,

404

goeldii (Forel), Pseudomyrmex,

405

gracilis (Fabricius),

Pseudomyrmex, 403-404, 439,
446

halimifolia , Baccharis, 420
haytianus (Forel), Pseudomyrmex,
405

heterogynus (Wheeler & Mann),

Quaest. Ent., 1989, 25 (4)

466

Pseudomyrmex, 404
holmgreni (Wheeler),

Pseudomyrmex, 404, 437, 440,
446

honduranus (Enzmann),
Pseudomyrmex , 437, 446
huberi (Santschi), Pseudomyrmex,
404

humboldi (Enzmann),
Pseudomyrmex, 404
ictericus (Wheeler),

Pseudomyrmex , 442, 447
implicatus (Forel), Pseudomyrmex ,
406, 422^123

incurrens (Forel), Pseudomyrmex,
404

infemalis (Wheeler),
Pseudomyrmex, 404
insularis (Enzmann),

Pseudomyrmex , 44 1 , 447
ita (Forel), Pseudomyrmex, 404
jaumei (Aguayo), Pseudomyrmex,
441,447

juliflora , Prosopis, 417
kuenckeli (Emery),
Pseudomyrmex, 405
kurokii (Forel), Pseudomyrmex,
437, 443, 446
laeviceps (F. Smith),

Pseudomyrmex, 440, 446
laeviceps (F. Smith), Tetraponera,
440

laevifrons Ward, Pseudomyrmex,
405, 440, 446
laevigatus (F. Smith),

Pseudomyrmex, 403^404
landolti (Forel), Pseudomyrmex,
404

lanuginosus , Pilosocereus, 417
latinodus (Mayr), Pseudomyrmex,
435, 441, 445-446
leptosus (Ward), Pseudomyrmex,

Ward

404

levivertex (Forel), Pseudomyrmex,
404

lincecumii (Buckley),

Pseudomyrmex ,436, 446
lisus Enzmann, Pseudomyrmex,
404

lizeri (Santschi), Pseudomyrmex,
404

longiceps (Forel), Pseudomyrmex,
438, 446

longiceps (Stitz), Pseudomyrmex,
438, 446

longior (Forel), Pseudomyrmex,
404

longituba , Cordia, 428
longus (Forel), Pseudomyrmex,
404

lynceus (Spinola), Pseudomyrmex,
404

maculatus (F. Smith),

Pseudomyrmex, 404, 440
major (Forel), Pseudomyrmex, 404
malignus (Wheeler),

Pseudomyrmex, 405, 435^136,
440, 445-446
mandibularis (Spinola),
Pseudomyrmex, 405
mangle , Rhizophora, 420
mexicanus (Roger),
Pseudomyrmex, 404
monochrous (Dalle Torre),
Pseudomyrmex, 404
niger (Donisthorpe),
Pseudomyrmex, 404
nigrescens (Forel),

Pseudomyrmex, 405, 436, 445
nigriceps (F. Smith),
Pseudomyrmex, 444
nigrocinctus Emery,

Pseudomyrmex, 404, 435, 438
nigropilosus (Emery),

Pseudomyrmecine Ants

467

Pseudomyrmex, 404, 435, 438
obnubilus (Menozzi),

Pseudomyrmex , 439, 446
obtusata , Tecoma, 428
oculatus (F. Smith),

Pseudomyrmex, 403-404,
406—407, 417, 422-423, 426
oerstediana , Ficus, 43 1
ogloblini (Santschi),

Pseudomyrmex, 406, 427-428
oki (Forel), Pseudomyrmex, 405
opacior (Forel), Pseudomyrmex,
405, 436

osurus (Forel), Pseudomyrmex,
404, 441,447

pallens (Mayr), Pseudomyrmex,

403- 404

pallidus (F. Smith),

Pseudomyrmex, 403—404
patens , Vemonia, 420
pazosi (Santschi), Pseudomyrmex,
404, 441,447
peltatus (Menozzi),
Pseudomyrmex, 404
penetrator F. Smith,
Pseudomyrma, 435
peperi (Forel), Pseudomyrmex,
404, 423, 452
perboscii (Guerin),

Pseudomyrmex, 405, 435, 442,
447

peruvianus (Wheeler),
Pseudomyrmex, 404
phyllophilus (F. Smith),
Pseudomyrmex, 404
pictus (Stitz), Pseudomyrmex, 404
pilosulus (F. Smith),
Pseudomyrmex, 404
pisinnus sp. nov., Pseudomyrmex,

404- 406, 408, 425

pupa (Forel), Pseudomyrmex, 404
racemosa , Laguncularia, 411, 420

rochai (Forel), Pseudomyrmex,
404

rubiginosus (Stitz),
Pseudomyrmex, 404
rufiventris (Forel),

Pseudomyrmex, 404, 437,
442—443, 447
rufomedius (F. Smith),

Pseudomyrmex, 405, 443, 447
rurrenabaquensis (Wheeler &
Mann), Pseudomyrmex, 405
saffordi (Enzmann),
Pseudomyrmex, 452
saffordi (Wheeler),

Pseudomyrmex, 437, 446, 452
salvini (Forel), Pseudomyrmex,
404

saman , Pithecellobium, 420, 442
santschii (Enzmann),
Pseudomyrmex, 405
sapii (Forel), Pseudomyrmex, 445,
447

scelerosus (Wheeler),
Pseudomyrmex, 404
schuppi (Forel), Pseudomyrmex,
404, 406, 423, 426
seminole Ward, Pseudomyrmex,
404

sepium , Gliricidia, 411, 413, 417,
420

sericatus (F. Smith),
Pseudomyrmex, 404
sericeus (Mayr), Pseudomyrmex,
403-404

simoides (Forel), Pseudomyrmex,
442, 447

simplex (F. Smith),

Pseudomyrmex, 404, 443, 447
simulans Kempf, Pseudomyrmex,
403-404

solisi (Santschi), Pseudomyrmex,
404

Quaest. Ent., 1989, 25 (4)

468

spiculus, sp. nov. , Pseudomyrmex,
405, 429—434, 448
spinicola (Emery),

Pseudomyrmex, 404, 435, 438,
443-444, 447
squamiferus (Emery),
Pseudomyrmex, 404
stans , Tecoma, 420
stolli (Forel), Pseudomyrmex, 443,
447

subater (Wheeler & Mann),
Pseudomyrmex, 405
subtilissimus (Emery),

Pseudomyrmex, 403, 405,
429-430, 432—433, 435, 438
symbioticus (Forel),

Pseudomyrmex, 405, 445
tachigaliae (Forel),

Pseudomyrmex, 403, 405,
435-436, 441, 444-445, 447
tandem (Forel), Pseudomyrmex,
406,419

tenuis (Fabricius), Pseudomyrmex,
403, 405, 444, 447
tenuissimus (Emery),

Pseudomyrmex, 405, 429—433
terminalis (F. Smith),
Pseudomyrmex, 404
termitarius (F. Smith),

Pseudomyrmex, 405, 444
testae eus (F. Smith),

Pseudomyrmex, 442, 447
thoracicus (Norton),
Pseudomyrmex, 404
tigrinus (Wheeler),

Pseudomyrmex, 445, 447
trigonus (Enzmann),
Pseudomyrmex, 445
trinervis , Baccharis, 420
triplaridis (Forel), Pseudomyrmex,
405, 435-436, 444-445, 447
triplarinus (Weddell),

Ward

Pseudomyrmex, 405, 435-436,
444-445

tuberculatus (Enzmann),
Pseudomyrmex, 406, 422
ulei (Forel), Pseudomyrmex, 445,
447

unicolor (F. Smith),
Pseudomyrmex, 404
urbanus (F. Smith),

Pseudomyrmex, 404—406, 409,
411,415,420, 427^128
variabilis (F. Smith),

Pseudomyrmex, 439, 446
velifer (Stitz), Pseudomyrmex, 404
veneficus Wheeler,

Pseudomyrmex, 404, 439
venustus (F. Smith),
Pseudomyrmex, 404
vesanus (Wheeler),

Pseudomyrmex, 437, 446
viduus (F. Smith), Pseudomyrmex,
403, 405, 435^136, 444-447
villosus sp. nov. , Pseudomyrmex,
405, 429—430, 432, 434
vinneni (Forel)), Pseudomyrmex,
404

virgo (Santschi), Pseudomyrmex,
404

vistanus (Enzmann),
Pseudomyrmex, 404
volatilis (F. Smith),
Pseudomyrmex, 404
voytowskii (Enzmann),
Pseudomyrmex, 405
wasmanni (Wheeler),

Pseudomyrmex, 437, 446
weberi (Enzmann),

Pseudomyrmex, 405
wessoni (Enzmann),

Pseudomyrmex, 406, 422—423
wheeled (Enzmann),
Pseudomyrmex, 405

Book Review

469

BOOK REVIEW

CAMPBELL, I., G. Theischinger & W.W.K. Houston, W.W.K. Houston & J.A.L.
Watson, F.B. Michaelis & C.M. Yule, and A. Neboiss. 1988. Zoological Catalogue
of Australia. Vol. 6. Ephemeroptera, Megaloptera, Odonata, Plecoptera, Trichoptera.
Bureau of Flora & Fauna, Canberra, ed. W.W.K. Houston. Australian Government
Publishing Service, GPO Box 84, Canberra, A.C.T. 2601. $Australian 29.95
(approx. $Canadian 27.00) incl. p. & p. ISBN 0 644 08269 3. xi+ 316 pp..

Not simply a list of names, synonymies, and literature associated with included
taxa, this book includes notes on general distribution of species, and bibliographical
annotations, where appropriate, by the authors of each ordinal section. Further, for
each order, and family within orders, there is an introduction which includes
comment on characteristics of the Australian fauna (endemism, etc.) of the taxon, a
brief description of each life stage of the taxon, and brief notes on taxonomic
history, biology, and distribution where available. The presentation, scope, and
extent of the information thus provided varies between the orders included,
depending, apparently, on the respective authors.

The orders are presented alphabetically, and the families within each are
arranged, as the Preface puts it, “phylogenetically”, whatever that may imply.

The genera, and species within genera are arranged alphabetically.

As the Preface points out, the nomenclature used for each taxon is governed by
the 3rd (1985) edition of the International Code for Zoological Nomenclature.

The geographical area encompassed by the catalogue is outlined by the only
illustration, a map of Australia and environs, on p. vi, and also by the Preface (p. vii)
which names the offshore islands involved (including the Australian Antarctic
Territory).

The Preface (incl. map) appears to be general for the series rather than specific to
vol. 6. For example, that part which explains the format of the catalogue at the
generic and specific levels employs an example from an obscure group of beetles,
the Carabidae!

The format of presentation comprises 1 1 points as follows (abbreviated):

1 . Genus valid name, author, year.

2. Genus available name, etc., if any.

3. Qualifying statement, if any.

4. Type species, associated information.

5. Bibliographical references to synonymy.

6. Extralimital distribution.

7. Species valid name, author, year.

8. Species available name, etc., if any.

9. Type data.

10. Distribution and ecology.

Quaest. Ent., 1989, 25 (4)

470

Book Review

11. Biological references (with brief notes on contents, as required, by the
present authors).

Depending on the circumstances attendant on each taxon, some of the above points
may be omitted.

The text, in common with other volumes of the series, is apparently generated
from a computer file, which is being constantly updated. The implication appears to
be that the next edition could appear 10 days after one acquires a current copy, or 10
years later. Indeed, this volume is so up to date that the existence of recognized but
as-yet-undescribed species is mentioned in passing, in some instances.

The strictly catalogue portion of the text is followed by three appendices (I -
Abbreviations and symbols; II - Museum acronyms; III - Taxonomic decisions
made in this work) and five taxonomic indexes, one for each order treated, in which
a species may be entered as many times as there are parts to the name ( e.g ., a species
will be entered under each of generic, subgeneric, and specific epithets, as
appropriate).

Ordinal and familial introductions and references (which are given in full) are
printed full page width, while generic and specific entries are in double columns.

As to the physical make-up of the book - the type face used is not easy on the
eye; not restful. The only word which comes to mind is ‘turgid’.

An oddity noted in passing is that, in the double column parts of the text, in order
to avoid having a family entry ending half way down a column on the left or right of
a page, the column will be split to give two parallel part columns, and the text
suddenly changes from single to double spacing, with the words of the double
spaced part-column themselves more widely spaced (the text margins are justified)!
This practice occurs in various situations, not just the specific one above (e.g., see
pp. 217, 234).

The volume is printed on matt-finish, high quality paper, and amazingly, in these
times, is strongly stitched in signatures. It won’t fall apart in the hand! It is
hard-bound in what appears to be a stout, institutional cloth. It’s meant to last.

In closing, I mention a curiosity found on p. 178 (Trichoptera section). Neboiss
writes “All publications containing original description have been sighted”.
Sighted?! I’m puzzled - does he mean cited; or seen? Is it merely a typographical
error, or have these publications been granted the boon of vision?

For those who work on, and take more than a local approach to, any of the five
orders included, this book is a must. At the price, it’s a bargain.

Andrew P. Nimmo
Department of Entomology
The University of Alberta

Book Notice

471

BOOK NOTICE

NICHOLS, S. W. (compiler), and R. T. SCHUH (Managing Editor). 1989. The
Torre-Bueno Glossary of Entomology. Revised Edition of a Glossary of
Entomology by J. R. de la Torre-Bueno, including Supplement A, by George S.
Tulloch. The New York Entomological Society and American Museum of Natural
History, xvii + 840 pp. ISBN 0-913424-13-7. Price $35.00 U. S.

“a-, prefix; wanting or without (T-B; Harris)”; “zymospecies-, a species
recognized soley |>ic] on the basis of characters obtained through enzymes
electrophoresis or comparable enzyme studies (Carlberg, pers. comm.)”. These
entries, respectively, begin and end the text of “The Torre-Bueno Glossary of
Entomology”.

Local entomological societies, at one time the backbone of entomology in North
America, have been displaced somewhat from a focal role by national organizations.
Nonetheless, their journals continue to provide outlets for first class publications,
especially about systematics, morphology and ecology. In 1937, one such
organization, the Brooklyn Entomological Society, published a work of inestimable
value to entomologists and destined to become a classic: “A Glossary of
Entomology”, by J. R. de la Torre-Bueno. In the course of the next 50 years, copies
of that publication, known to the initiated as simply “Torre-Bueno”, appeared on the
bookshelves of most North American entomologists, and on the bookshelves of
entomologists from elsewhere who had occasion to read or write English
entomological publications. As this work was becoming outdated, George S.
Tulloch provided Supplement A, in 1962- but with the passage of time, that
supplement became outdated, as well. Clearly, an extensive revision was required.

In the meantime, the Brooklyn Entomological Society and its assets became
incorporated with the New York Entomological Society, the Executive of which
determined to undertake a revision of the Torre-Bueno Glossary. Stephen W.
Nichols was employed, and together with Randall T. Schuh of the American
Museum, and some 50 editorial contributors, the work was undertaken, eventually
leading to publication of a volume more than twice the size of the original
“Torre-Bueno”.

Prefatory material includes: a list of names of the Editorial Contributors; Table
of Contents; Preface; Acknowledgements; Introduction to the Revised Glossary, to
the Original Glossary, and to Supplement A; and a list of the 32 names of hexapod
orders used in the Glossary. In the text, each main entry is in boldface type. For each
entry, the authority or authorities who proposed it are cited.

Following the 823 pages of text, references are cited: “Principal Sources for the
Current Edition”; “Additional Sources for the Current Edition”; “Non-English
Language Glossaries and Sources”; “Sources to English Common Names of
Insects”; and “Other Useful References”.

Quaest. Ent., 1989, 25 (4)

472

Book Notice

A thick, heavy book, the volume has a strong binding of an attractive shade of
brown buckram. Obviously, it is intended to last an entomologist for his working
life. All in all, it is an aesthetically pleasing book, prepared with care and thought.
This volume should be in the possession of every anglophone entomologist. At
$35.00, it is the entomological buy of the latter half of the 20th Century.

The New York Entomological Society and American Museum of Natural History
must be commended for undertaking publication of this volume. Those who worked
to produce the text, and especially the compiler and the Managing Editor deserve
high praise for their efforts. Perhaps in the fullness of time, entomologists will refer
to this volume as their beloved “Nichols-Schuh”, rather than “Torre-Bueno”.

f

473

EDITOR’S ACKNOWLEDGEMENTS AND COMMENTS

Because of forgetfulness and the pressures associated with preparation for and
departure on sabbatical leave, the Editor neglected to offer his thanks to those who
assisted in evaluation and preparation of the contributions for Volume 24.
Accordingly, the names of those who provided reviews for that volume are included
in the following list. The able assistance, promptly and cheerfully rendered by all of
the reviewers, is appreciated very much.

R. S. Anderson

Department of Entomology, Texas A & M University
B. Bolton

Department of Entomology, British Museum (Natural History)

T. L. Erwin

Department of Entomology, U. S. National Museum of Natural History
R. H. Gooding

Department of Entomology, University of Alberta
M. J. R. Hall

Department of Entomology, British Museum (Natural History)

A. T. & H. F. Howden

Department of Biology, Carleton University
D. H. Kavanaugh

Department of Entomology, California Academy of Sciences
J. K. Liebherr

Department of Entomology, Cornell University
M. W. Moffett

Department of Entomology, Museum of Comparative Zoology, Harvard

University
M. Sharkey

Biosystematics Research Centre, Agriculture Canada
M. V. H. Wilson

Department of Zoology, University of Alberta

During my absence, my colleague, Bruce S. Heming, acted as Editor. I am
pleased to have this opportunity to thank him for his thorough and enthusiastic
service.

French translations of abstracts were provided by J. C. Lacoursiere, of my
Department. J. S. Scott and D. Shpeley read proof, as requested, and assisted in
other ways, as well.

The Publications Manager, Mrs. S. Subbarao, provided excellent service, both in
production and in management of day-to-day operations of the office.

Quaest. Ent., 1989, 25 (4)

474

Our printers, Hallis Graphics Ltd., have served us well. I am mindful especially
of prompt turnaround of our submissions, and the air of cordiality that characterizes
our relationship with the President of Hallis, Mr. J. Haukedal.

With the publication of Volume 25, Quaestiones Entomologicae reached what is
generally regarded as a landmark of some significance, and recognized as a Silver
Anniversary. Although 25 years of publishing is an achievement, it seems
sufficiently commonplace not to warrant a formal celebration. Such can be reserved
for our 50th, or Golden Anniversary (in the year 2014), though the present members
of the publications staff are unlikely to be in attendance.

In passing, I note that Volume 1 contains a paper by Richard Freitag, about tiger
beetles, in the form of an analysis of the Cicindela maritima species group. Volume
25 contains a paper by the same author, about the Brazilian species of Cicindela.
And in Volume 24, there is a paper by Timothy G. Spanton, about the species of the
Cicindela sylvatica group, based on his Master’s thesis - which was supervised by
Freitag. Such connections through the passage of time indicate a stability that the
Editor finds satisfying and gratifying.

It seems appropriate to offer a note of appreciation in memory of the late founder
and first Editor of Quaest. Ent., Brian Hocking, who died in 1974. In his
introductory editorial, “words, words, words”. Hocking noted the trend of marked
increase in entomological publication, but saw it as the necessary consequence of
burgeoning effort in research. Implicitly, he expressed dismay about: 1, “mounting
page charges”; 2, the “publish or perish” blight; and 3, “the waning ability of
administrators to judge publications by anything beyond their number”. He
recognized that points 2 and 3 were contributing, undesirably, to the marked
increase in numbers of scientific publications.

He suggested that journals might help scientists keep abreast of the exponential
increase in published information by ensuring presentation of more comprehensive
papers (”It costs more in time, money, and effort to produce, file, store, retrieve, and
read ten one page papers than one ten page paper”), to “index and abstract
everything diversely”, and to “make full use of modem techniques... even
computers”. Quaest. Ent. was organized along such lines.

Hocking paid tribute to entomology’s “venerable indexing services” (Zoological
Record; H. A. Hagen’s Bibliotheca Entomologica; and W. Horn and S. Schenkling’s
Index Litteraturae Entomologicae). He held in high regard the contributions that
such publications make to improve retrieval of entomological information.

At the time that he wrote, indexing and abstracting publications endeavoured to
provide complete coverage of the entomological literature. This seems to be so no
longer. The more blatantly commercial of such publications pick and choose which
journals will be indexed or abstracted, and the status of journals in the scientific
community now seems to be determined in part by whether or not their offerings are
recorded by one or another commercial “service”. Paralleling this development is
the increasing trend of University administrators and granting agencies to evaluate

475

papers in terms of the journals in which they appear, rather than for their intrinsic
merit. Many scientists have bowed to such pressure.

Thus, scientific publication now seems to have more to do with gaining
individual recognition and prestige than with expressing desire to make known the
results of research. Indeed, the medium is becoming the message. Perhaps it was
ever thus, but Brian Hocking’s generous spirit would be repelled by the mean,
self-serving attitudes that have become commonplace in the scientific community.
Such attitudes, of course, are simply latter-day expressions of the “publish or perish”
syndrome that is a dark blight on the landscape of science.

Be that as it may, Quaest. Ent. has continued along the path that Hocking
charted. The journal has kept its covenant with its founder as best it could - or so it
seems to the present editor. Some subsequent Editor, who, metaphorically speaking,
will blow out the candles on the journal’s 50th Anniversary cake will be able to
offer a more objective opinion than the one proffered by the present Editor, who has
presided over this non-celebration of Quaest. Ent.’s 25th birthday.

George E. Ball
Editor

Quaest. Ent., 1989, 25 (4)

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